1b.marine Phytoplankton Vol2

MARINE PHYTOPLANKTON
OF KUWAIT’S WATERS
Volume II
Diatoms
Faiza Y. AL-YAMANI and Maria A. SABUROVA

Volume II
Diatoms
Kuwait Institute for Scientific Research

2019
MARINE PHYTOPLANKTON OF KUWAIT’S WATERS
Volume II. Diatoms
Published in Kuwait in 2019 by

Kuwait Institute for Scientific Research,
P.O. Box 24885, 13109 Safat, Kuwait
Copyright © Kuwait Institute for Scientific Research, 2019

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ISBN 978-99966-37-20-9
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.
Contents
Acknowledgments …………………………………………………………………………………. 5
Opening Remarks …………………………………………………………………………………… 7
Introduction …………………………………………………………………………………………. 8
Materials and Methods …………………………………………………………………………….. 10
General Identification Characteristics …………………………………………………………….. 13
Glossary………………………………………………………………………………………………. 14
SYSTEMATIC ACCOUNT: Marine Planktonic Diatoms

Phylum BACILLARIOPHYTA ……………………………………… ………………………….... 16
Subphylum Coscinodiscophytina ……………………………………………………………… 16
Class Coscinodiscophyceae ………………………………………………………………... 16
Order Rhizosoleniales …………………………………………………………….. 16
Order Triceratiales ………………………………………………………………… 34
Subclass Archaegladiopsophycidae …………………………………………………… 36
Order Stephanopyxales ……………………………………………………………. 36
Subclass Corethrophycidae ……………………………………………………………. 38
Order Corethrales ………………………………………………………………….. 38
Subclass Coscinodiscophycidae ……………………………………………………….. 40
Order Asterolamprales …………………………………………………………….. 40
Order Coscinodiscales …………………………………………………………….. 47
Order Stellarimales ………………………………………………………………… 72
Subclass Melosirophycidae ……………………………………………………………. 74
Order Melosirales ………………………………………………………………….. 74
Subphylum Bacillariophytina ………………………………………………………………….. 76
Class Mediophyceae ………………………………………………………………………... 76
Subclass Biddulphiophycidae …………………………………………………………. 76
Order Biddulphiales ……………………………………………………………….. 76
Order Briggerales ………………………………………………………………….. 82
Subclass Chaetocerotophycidae ………………………………………………………. 86
Order Chaetocerotales …………………………………………………………….. 86
Order Hemiaulales …………………………………………………………………. 136
Subclass Cymatosirophycidae …………………………………………………………. 144
Order Cymatosirales ………………………………………………………………. 144
Subclass Thalassiosirophycidae ………………………………………………………. 146
Order Eupodiscales ………………………………………………………………… 146
Order Lithodesmiales ……………………………………………………………… 153
Order Stephanodiscales ……………………………………………………………. 156
Order Thalassiosirales …………………………………………………………….. 164
Class Bacillariophyceae ……………………………………………………………………. 213
Subclass Urneidophycidae …………………………………………………………….. 213
Order Rhaphoneidales …………………………………………………………….. 213
Subclass Fragilariophycidae …………………………………………………………… 216
Order Licmophorales ……………………………………………………………… 216
Order Rhabdonematales …………………………………....…………………….. 218
Order Thalassionematales ………………………………………………………… 218
Subclass Bacillariophycidae …………………………………………………………… 222
Order Cocconeidales ………………………………………………………………. 222
Order Lyrellales ……………………………………………………………………. 228
Order Mastogloiales ……………………………………………………………….. 230
Order Naviculales ………………………………………………………………….. 231
Order Bacillariales …………………………………………………………………. 271
Order Surirellales ………………………………………………………………….. 305
Order Thalassiophysales ………………………………………………………….. 316
References …………………………………………………………………………………………… 322

Index to taxa ………………………………………………………………………………………… 333
“We should preserve every scrap of biodiversity as
priceless while we learn to use it and come to understand
what it means to humanity.”
E. O. Wilson
Coral reef surrounding Qaru Island

Photo by Dr. Igor Polikarpov
4 M A R I N E P H Y TO P L A N K TO N O F K U WA I T ’S WAT E R S . V O LU M E I I : D I ATO M S
ACKNOWLEDGMENTS
The authors acknowledge with appreciation the reviews provided
by the experts for the taxonomic part of this book. We would like
to acknowledge the long-term cooperation received from Drs. of Sc.
Elena Nevrova (A.O. Kovalevskiy Institute for Marine Biological
Research, Sevastopol, Crimea) and Ludmila Ilyash (Moscow State
University, Russia).
We acknowledge with thanks the assistance provided by Mr. Alan

Lennox and the cruise staff of the Environment and Life Sciences
Research Center (ELSRC) at the Kuwait Institute for Scientific
Research for obtaining the phytoplankton samples on which
this study was based. We are grateful to Mr. Noumaan Qureshi
(TESCAN Middle East, Dubai, UAE) and Mr. Andrey Averichkin
(TESCAN Ltd., Moscow Service Department, Russia) for their
advice and training in scanning electron microscopy. Particular
gratitude is expressed to Dr. Igor Polikarpov (ELSRC, KISR) for
providing support and help throughout the long period of this
research and the manuscript preparation.
The authors are grateful to the Kuwait Institute for Scientific

Research for providing financial support of the different research
projects, based on which the data for this book were obtained.
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 5
Shore in Khiran
OPENING REMARKS
This is the second volume of the book on Marine Phytoplankton of
Kuwait’s Waters, which deals with marine planktonic diatoms. The
present volume summarizes the results of more than a 10-year period
(2004-2017) of phytoplankton surveys in Kuwait’s waters occupying
the northwestern part of the Arabian Gulf. Through examination of
the samples collected throughout the coastal and offshore habitats
all the year round, we illustrate and document the diatom diversity
present in the studied habitats.
This book describes more than 250 taxa of marine diatoms belonging
to 47 families, 28 orders and 3 classes encountered in Kuwait’s waters
during the studied period. Each taxon is presented with a description
of cell morphology, its habitat and locality, and any known toxicity
and harmful effect. The morphological descriptions are illustrated
with various angles of the light photomicrographs for all species to
show key diagnostic characters. The nomenclatural species names
used are valid as of those published in 2017 (Guiry & Guiry, 2017
and references therein). The synonyms for species and intraspecific
taxa are listed according to the latest systematic and nomenclatural
changes. In addition, the global geographical distribution, and
local occurrence with seasonal abundance of species are indicated
based on data collected over the past decade. Each group of related
diatom species is preceded by an introductory taxonomical and
ecological overview covering the major diagnostic features of
the group. The relevant literature references are included in the
extensive bibliography, and notes in regard to some taxa are given,
where appropriate. The presented species are profusely illustrated
by means of 1427 high-resolution light and scanning electron
photomicrographs on 161 plates. The introductory overview of the
diatoms covers the major diagnostic features of their frustules.
The presented phytoplankton taxonomic account of the diatom

species in the north-western part of the Arabian Gulf provides
baseline information on phytoplankton composition. The book
constitutes an important reference and tool for further taxonomical
and ecological assessments, including studies on the regional
phytoplankton diversity, in the detection of invasive species, the
identification of potentially harmful species and other related
investigations. The book is intended to be a helpful source for
students and researchers in the field of the phytoplankton taxonomy
and ecology.
INTRODUCTION
Diatoms are single-celled photosynthetic eukaryotes universally distributed in all types of aquatic
environment. Diatoms form the majority of floating algae in marine and fresh waters, and also are an
important fraction in the benthic environments. They constitute a large component of aquatic biomass in
nutrient-rich coastal ecosystems, particularly during conspicuous seasonal phytoplankton blooms, and have
been estimated to contribute 40-45% oceanic primary productivity, which amounts to 20% of global carbon
fixation and oxygen production (Falkowski et al., 1998; Field et al., 1998; Mann, 1999; Malviya et al., 2016).
Diatoms multiply rapidly, maintaining a dynamic population of varying size. Their photosynthesis, biogenic
silica formation, environmental diversity and a tendency to dominate phytoplankton communities, have led
to the major involvement of diatoms in primary production, nutrient cycling, the biological carbon pump,
and at the base of the food chain (Mann, 1999; Malviya et al., 2016).
Diatoms are abundant, diverse, ubiquitous, and sensitive environmental indicators and, thus, have
an enormous ecological importance. This is one of the most species-rich groups of algae with the highest
estimates of around 200,000 extant species spread across all aquatic habitats (Mann & Droop, 1996). A large
number of ecologically sensitive species makes this group the ideal ecological indicator with a wide range
of applications, both as living organisms and fossils (Smayda, 1990; Round, 1993; John, 2000), and requires
accurate and consistent species identification.
The ecology and taxonomy of the phytoplankton in the north-western Arabian Gulf have been
studied during the past few decades (Enomoto, 1971; Al-Kaisi, 1976; Jacob et al., 1979; Al-Yamani, 1989;
Al-Yamani et al., 1997; Subba Rao et al., 1999) but involved very limited detailed taxonomical surveys.
A valuable review of phytoplankton diversity for the region has been conducted by Subba Rao and Al-
Yamani (1998) based on the available data on phytoplankton from the different parts of the Arabian Gulf.
Information about the distribution of some phytoplankton species occurring in Kuwaiti waters has been
published by Enomoto (1971), Al-Yamani et al. (2004), and Al-Kandari et al. (2009). There have only been
a few studies on the benthic diatom flora of the
north-western part of the Arabian Gulf (Hendey,
1970; Basson & Mohammed, 1989; Al-Handal,
2009; Al-Yamani & Saburova, 2011). The latest
estimation of phytoplankton diversity in Kuwaiti
waters was reported recently by Al-Yamani et
al. (2004) and by Al-Kandari et al. (2009) and
documented more than 320 taxa with a great
prevalence of diatoms.
As a whole, a pronounced numerical

prevalence of diatoms is typical for the
phytoplankton community in nutrient-rich
Kuwait’s waters throughout the year at most
of the sampled area (Subba-Rao & Al-Yamani,
1998; Al-Yamani et al., 2004; Polikarpov et al.,
2009, 2016). As the dominant group, diatoms
contribute 70-90% (up to 99.8%) to the total
phytoplankton abundance. Their prevalence
is at a maximum during the autumn-winter
period, especially in November and December,
and reduces during the spring and summer
(Polikarpov et al., 2009, 2016). Diatom-
dominated blooms are regularly observed
Fig. 1. Diatom-dominated phytoplankton bloom in Kuwait’s
waters.
in Kuwait’s waters (Fig. 1). The highest concentrations attained were more than 106-107 cells/l and are
associated with the inshore waters of Kuwait Bay (Al-Yamani et al., 2004; Polikarpov et al., 2009; Al-
Yamani et al., 2012). Among the common bloom-forming diatoms in Kuwait’s waters there are Chaetoceros
curvisetus, Chaetoceros pseudocurvisetus, and Chaetoceros socialis, Thalassiosira spp., Cyclotella spp., Skeletonema
grevillei, Guinardia flaccida, Eucampia zodiacus, and small-sized Nitzschia species.
The introduction of pennate diatoms, especially the species of typically benthic habitats from
periphyton, epipelon and epipsammon (genera Pleurosigma, Diploneis, Surirella, Trachyneis, Nitzschia,
Entomoneis, Plagiotropis) into the water column greatly influences the species composition of the phytoplankton
in Kuwait’s shallow waters. Resuspension of these benthic species from the bottom sediment to the surface
waters can occur via turbulent mixing including tidal forcing, winds or surface waves. Some pennate diatoms
(such as Pleurosigma spp., Surirella fastuosa, Trachyneis antillarum, and Nitzschia spp.) are occasionally found to
be quite abundant, particularly in inshore habitats along Kuwait’s coast.
Marina Crescent Beach, Salmiya

MATERIALS & METHODS
Sampling
The long-term phytoplankton survey covering Kuwait’s waters from the north to the south was conducted
during 2004-2017 (Fig. 2). A wide range of sampling sites were established to represent a variety of marine
habitats, including northern areas off Bubiyan and Failaka islands, Kuwait Bay, inshore waters along
Kuwait’s coast, offshore open waters, southern coastal waters, and surrounded waters of small offshore
islands including Kubbar and located at the extreme south of Kuwait’s marine waters Qaru and Umm Al-
Maradim islands.
Samples were obtained from the routine long-term phytoplankton surveys, which were supplemented
with occasional samples that were collected during red tide and fish kill events, which occurred in Kuwait’s
waters during the past decade. Monthly routine phytoplankton surveys were conducted; in addition, selected
localities were sampled on a biweekly or quarterly basis.
One-liter water samples from the surface layer (1 m depth) were collected using a 5-liter Niskin bottle
sampler and were carefully transferred to sampling containers through a flexible hose. In addition, vertical
net tows were conducted using a 20 μm-mesh plankton net. Samples were preserved with 4% acidified
Lugol’s solution. Replicate samples were taken at aech sampled site to obtain both preserved and living
materials simultaneously.
Sample Processing
Phytoplankton diatoms enumeration. To quantify the phytoplankton taxa, a 25 ml of water sample was
settled in a Utermöhl sedimentation chamber (Utermöhl, 1958) for at least 24 h before enumeration, and
examined with a Leica DM IL inverted microscope (Leica Microsystems GmbH, Wetzlar, Germany) at
200× or 400× magnification. Abundant phytoplankton species were counted on a variable number of
random fields until 400 counting units were recorded (Lund et al., 1958). In addition, the entire bottom of
the chamber was examined to obtain a more reliable evaluation of less abundant species. In case of very low
phytoplankton abundance, the whole volume of the sample was concentrated by sedimentation to 150-200
ml before enumeration. The abundance of each species/taxon was calculated as the number of cells per liter.
To detect the phytoplankton species that were overlooked in the analysis of bottle samples, 20 ml
aliquots of net samples were examined with an inverted microscope. The phytoplankton taxa exclusively
found from net samples were recorded as present but were not enumerated. Further, the species richness
was assessed based on the assembled data set on species occurrences from both bottle and net samples. This
strategy provided a better approach for taking more representative materials to study the phytoplankton
taxonomic composition in the studied areas.
A total of 470 Lugol-preserved water samples have been analyzed to assess the phytoplankton
composition and species abundances during the present study. This extensive data set was further utilized to
characterize the local occurrences, seasonal distributions and ranges of abundance for the described taxa.
Species identification. Species identifications were mostly based on live material obtained from both bottle
and net unpreserved samples. Nearly all of the taxonomic information on delicate diatoms came from
observing live cells. More rigid highly silicified diatoms were also examined alive whenever possible to
maintain the natural pigmentation, chloroplasts morphology, and general appearance of their cells.
For a detailed examination, diatom cells were isolated by micropipetting, and transferred to a glass
slide in preparation for high-magnification photomicroscopy. Cells were examined and photographed using
a compound Leica DM LM microscope (Leica Microsystems GmbH, Wetzlar, Germany) equipped with a
DFC425 color digital camera at 400-1000× magnifications. A range of microscope tools was used for cells
Fig. 2. Studied area and sampling site locations.
examination, including brightfield (BF) illumination to show the natural color of the organisms and to
discern the chloroplasts arrangement and nucleus location, and Nomarski differential interference contrast
(DIC) or phase contrast optic to visualize a three-dimensional appearance of cellular content and frustule
structure.
Identifications were principally made at the species level whenever possible, except for the small-
sized fragile diatoms and morphologically similar taxa, for which only the genus or group was recorded
(e.g., small and morphologically similar Thalassiosira and Nitzschia species) due to the difficulties of accurate
identification. Typically, such cells did not possess enough morphological characters for identification at a
lower taxonomic level without the aid of more complex diagnostic tools. Morphological features of species
belonging to genera such as Chaetoceros, Thalassiosira, Coscinodiscus, Navicula and Pseudo-nitzschia needed to be
investigated with the electron microscope in order to be accurately identified.
Most diatom species were examined from cleaned material. The diatom frustules have a light
refraction close to that of water, but by removing the organic material and embedding the cleaned silica parts
in a medium with a different light refraction, many details, which are important for identification, could be
revealed under the light microscope.
For cleaning the highly silicified diatom frustules, a mixture of sulfuric acid with potassium
dichromate was used. More delicate weakly silicified species were treated with a solution of hydrogen
peroxide. Subsamples were placed in vials where organic coatings were solubilized for 24-48 h in these
mixtures. After complete organic matter solubilization, the diatom frustules were rinsed with 4-5 changes of
distilled water by centrifugation. The cleaned material was mounted for examination under the light (LM)
and scanning electron (SEM) microscopes.
For LM examination, the cleaned diatom frustules were dried up on a cover glass and mounted in a
medium of a high refractive index (Melmount). The cleaned frustules from the permanent slides were observed
using the Axiovert-200 microscope (Carl Zeiss) with phase contrast optic and photographed using the high-
resolution AxioCamHR digital camera. For SEM observations, cleaned diatom frustules were mounted
directly on metal stubs and dried from the distilled water in which they were stored. The mounted samples
were then coated with gold-palladium and examined by either JEOL’s JSM-6300 (Electron Microscopy Unit
(EMU) at Kuwait University) or TESCAN Vega3 SBU (KISR) scanning electron microscope.
Estimation of cell sizes. Size is an important criterion for the identification of the different phytoplankton
species. Cell dimensions were measured by light microscopy using a calibrated ocular micrometer or from
calibrated images using Leica Application Suite v. 3.7 (Leica Microsystems (Switzerland) Ltd) software. In
addition, some morphometric measurements were conducted from micrographs using AxioVision 3.0 (Carl
Zeiss) software as well as from SEM micrographs. Measurements were mainly based on several imaged
specimens. Measurements without a range sometimes came from a single specimen found.
Taxonomic treatment
In this book, for practical purposes, the high-level classification of the described diatom taxa and verification
of currently accepted taxonomic names followed AlgaeBase (Guiry & Guiry, 2017), which is constantly
updated and can be considered as a practical compromise among diverse opinions and usages, as well as
when encountering conflicting evidence about certain morphological and phylogenetic questions important
for defining the boundaries between and ranks of major taxa. Along with true planktonic diatoms, we have
included presumably benthic taxa that were occasionally found in the plankton samples, which were most
probably due to resuspension from the bottom sediments.
GENERAL IDENTIFICATION CHARACTERISTICS
Diatom Taxonomy
Generally, there is currently no consensus among the world’s taxonomists concerning which classification
scheme to use for the overall hierarchy of life (e.g., Cavalier-Smith, 1981; Ruggiero et al., 2015). The
classifications of the different microalgal groups are constantly being reviewed and revised with the
rearrangement of systematic entities due to ever-increasing knowledge on the diversity of this group. Based
on the light microscopy for a long time, the algal systematic was largely descriptive, whereas over the past
decades, ultrastructural- and molecular-based revisions of classification at nearly all taxonomic levels have
occurred (e.g., Round et al., 1990; Ruggiero et al., 2015; Adl et al., 2005). Diatom taxonomy is in a state
of revolutionary change involving splitting of genera, the creation of new ones and revision of criteria for
classification. With molecular genetics and breeding as added tools, the taxonomy of diatoms has become
a dynamic field of research (Round et al., 1990; Medlin & Kaczmarska, 2004; Adl et al., 2005; Sims et al.,
2006; Williams & Kociolek, 2007).
The siliceous cell wall has been the main focus of attention for classification of diatoms. Diatom
valves are ornamented by a variety of siliceous structures: porefields, areolae, striae, spines, labiate
processes, and strutted processes. The diatom taxonomy has been almost exclusively based for a long time
upon frustule characteristics: shape, size, symmetry, structure and density of striae, nature of raphe and its
position, copulae and processes on the valves. However, increasingly, there is a trend towards ultrastructure
and features of living cells such as chloroplastids, life cycle pattern and reproductive biology to be considered
in the systematic (e.g., Round et al., 1990; Medlin & Kaczmarska, 2004).
In the early days of diatom systematics, the classification of different species was based exclusively
on features visible using light microscopy. The simplest classification approach was broadly based on the
distinction in the valve symmetry between the radially symmetric centric and bilaterally symmetric pennate
diatoms (Karsten, 1928).
The advent of scanning electron microscopy in the 1960s-1970s revolutionized diatom systematics,
revealing taxonomically important ultra-structural features not visible with the LM. A further more
complicated classification scheme by Round, Crawford and Mann recognized diatoms as a division
BACILLARIOPHYTA with three major classes corresponded to three main types of valve organization:
Coscinodiscophyceae (centric diatoms, generally radially-symmetric forms with no raphe), Fragilariophyceae
(araphid pennate diatoms, bilaterally-symmetric forms without raphe), and Bacillariophyceae (raphid
pennate diatoms, bilaterally-symmetric/asymmetric forms bearing raphe at least in one valve) (Round et al.,
1990).
More recently, molecular information has increased in importance in diatom systematics, and has
been applied, along with morphological data, to a new classification scheme by Medlin & Kaczmarska
(2004), which proposed a replacement for the traditional view suggesting two new subdivisions,
COSCINODISCOPHYTINA and BACILLARIOPHYTINA. The COSCINODISCOPHYTINA
included one class, Coscinodiscophyceae, composed of radially symmetric centric diatoms. The
BACILLARIOPHYTINA included two classes, Mediophyceae (radially symmetric diatoms with fultoportulae
and many bi- to multipolar diatoms previously included among the centric diatoms) and Bacillariophyceae
(araphid and raphid pennate diatoms). Further, the latest classification scheme was adopted by Adl et al.
(2005), who treated, however, both Coscinodiscophyceae and Mediophyceae as unnatural paraphyletic
groups (Adl et al., 2005; Williams & Kociolek, 2007). Supplementary, several alternative molecular trees have
been proposed, none appearing to support the monophyly of either Coscinodiscophytina or Mediophyceae
(Williams & Kociolek, 2007).
Frustule Morphology
The most obvious feature of diatoms is their siliceous cell wall (frustule) made of two intricately sculptured
halves (valves). A diatom cell under the microscope may be seen in different views. The term ‘valve view’
refers to the view of the ‘face’ of a diatom frustule. The side view (profile) is referred to as the girdle view.
The symmetry and structure of the frustule is extremely important in the systematic of diatoms and
basically related to their axes (John, 2000). Diatoms are found in two general structural types: cells with
radial symmetry and those with bilateral symmetry. The centric diatoms exhibit symmetry about a central
axis and most frequently, but not always, are circular. Pennate diatoms are generally elongated, and many
exhibit bilateral symmetry, and even helical symmetry is possible (Round et al., 1990). Genera with complex
shapes include Entomoneis, Petrodictyon, Surirella, Amphora, and Rhopalodia.
The upper lid of the frustule (epivalve) overlaps the lower lid (hypovalve). The epivalve and hypovalve
are connected by girdle bands termed copulae made of silica. The structure and type of bands are significant in
the taxonomy of diatoms. In some species of diatoms, the copulae have inwardly directed septa. In the genus
Mastogloia the valvocopula has internal chambers termed partecta. The majority of the pennate diatoms have
a longitudinal slit termed raphe. The pennate diatoms with raphe are referred to as ‘raphids’ and without
raphe as ‘araphids’ (Round et al., 1990; John, 2000).
The siliceous structure of the valve consists of transverse and short lateral ribs separated by tiny
‘pores’ or ‘areolae’ (puncta). These little holes arranged in rows are termed striae. Secondary deposition of
silica may transform the areolae into complex ‘chambers’ or locules as seen in centric diatoms (Round et al.,
1990; John, 2000).
Diatom valves may have spines, hair-like structures (setae) and horns which may be solid or hollow.
These siliceous appendages from the frustules provide linkage between cells to form chains. In addition to these,
many species have special processes termed ‘portules’. There are two types of portules: fultoportulae (strutted
processes) and rimoportulae (labiate processes), the former is confined to the centric order Thalassiosirales and
the latter wide spread in centric and pennate diatoms (Round et al., 1990; John, 2000).
GLOSSARY
alveoli - transversely linear chambers in the valve with a small opening externally and large opening internally.
apex - pole (end) of diatom valve.
apical axis - axis connecting the two poles of a valve.
apiculate - pole (apex) of valve tapering to a point.
araphid - diatoms without any raphe.
arcuate - strongly curved.
areolae - chambered pores in the diatom valve.
axial area - the area between the raphe and striae on the valve face.
bifurcate - divided into two branches.
biraphids - diatoms with raphe on both valves.
biseriate - two rows of pores or areolae.
canal raphe - raphe lying in a distinct groove.
capitate - pole (end) swollen.
central area - the clear area at the center of the valve face.
central ending - the external groove of the raphe slit at the center of the valve.
central nodule - the thickly silicified area in between the central endings of the raphe slits.
cingulum - girdle.
convergent striae - striae converging (angling) towards the poles.
copulae - girdle bands.
costae - distinctly thickened ribs on the valve face.
cruciform - cross-shaped.
cuneate - wedge-shaped.
dorsal margin - the convex margin.
eccentric raphe - raphe deviating from the apical axis.
elliptical - shape of valve ellipsoid.
fascia - a distinct central area extending to the margins of the valve, without any marking.
fibula(e) - siliceous bridge on either side of a raphe.
foot pole - narrow half (basal) of a club-shaped valve.
frustule - the cell walls of a diatom connected by girdle bands.
fultoportula(e) - strutted process, a tubular process through the valve of some centric diatoms. The
fultoportula consists of a central tubular process surrounded by two or more satellite pores.
Externally, the fultoportula appears as either a tube or a simple pore in the valve wall.
girdle view - side view of cell with girdle bands.
head pole - broad (upper) half of a club-shaped valve.
helictoglossae - internal, distal termination of the raphe in the shape of a pair of lips or a rolled tongue.
heteropolar - poles (apices) of valve being dissimilar.
hyaline area - clear area (siliceous) without any marking.
isopolar - poles of a valve being similar shape and size.
keel wing - a raised part of a valve face eccentrically located to the apical axis.
labiate process - see rimoportula.
lanceolate - lance shaped.
lenticular - lens shaped.
linear - long and narrow almost uniform in width.
linear-elliptic - elongated with inflated ellipsoid margins.
longitudinal striae - striae parallel to the longitudinal axis (apical axis) of the valve.
lunate - crescent shaped.
mantle - the side (outermost) part of valve bending downwards from the surface to the girdle.
monoraphid - diatom with a raphe only on one valve.
mucilage - viscous polysaccharide secreted by diatoms.
multiseriate - many rows of pores or areolae.
orbicular - spherical.
ovate - oval shaped.
panduriform - shaped like a fiddle (‘8’ shaped).
parallel striae - striae arranged more or less parallel to each other.
pennate - valve with laterally symmetrical.
polar nodule - thickened silicified area of the pole where the terminal ends of the raphe are located.
puncta - poroid opening of the siliceous valve – a traditional term for areolae.
radiate striae - striae pointing to the center of the valve.
raphe - a longitudinal slit (groove) in the valve face located between the two polar nodules, with two raphe
branches.
raphe branch - refers to the raphe branch from the center to the pole. A raphe consists of two raphe branches.
raphe-sternum - thickened hyaline axial area in pennate diatom (pseudoraphe).
rimoportula(e) - labiate process, a tubular process through the valve of some diatoms. On the internal
valve face, the rimoportula opening has the shape of a pair of lips. On the external valve face, the
rimoportula opening may be a simple, round aperture at the valve surface or it may be a tube extending
out from the valve.
rostrate - the apex of a valve ending like a beak.
septum - the inward expanded plate from the valvocopula (girdle band).
sigmoid - shaped like ‘S’ with the ends curving in opposite direction.
stauros - thickened central nodule extending to the margin of valve.
stria(e) - row of pores in valve.
strutted process – see fultoportula.
terminal endings (terminal fissures) - polar endings of raphe.
transapical axis - axis through the center connecting the margins across.
transverse striae - striae running across the valve face.
uniseriate - single row of pores or areolae.
valve face - the surface of valve surrounded by the mantle.
valve margin - the edge of valve face.
valvocopula - copula (girdle element) closest to the valve – different in structure from the rest of the cingulum.
ventral - margin (straight or concave) opposite to the dorsal side.
SYSTEMATIC ACCOUNT
Marine Planktonic Diatoms
Empire EUKARYOTA Chatton

Kingdom CHROMISTA Cavalier-Smith
Phylum BACILLARIOPHYTA Dillon
Subphylum Coscinodiscophytina Medlin et Kaczmarska
Class Coscinodiscophyceae Round et Crawford
Order Rhizosoleniales Silva
Family Probosciaceae Nikolaev et Harwood
Genus Proboscia Sundström
The generic type, Proboscia alata, was first described as Rhizosolenia alata with a series of forms. The different
forms are now regarded as separate species (Throndsen et al., 2007; Guiry & Guiry, 2017). Species of the
genus Proboscia are distinguished by the subconical valve extended into a shorter or longer tube with truncate
tip (termed ‘proboscis’), and by the absence of internal processes. In chain-forming cells, the distal part of
proboscis fits into a groove on the adjacent valve of linked cells (Sundström, 1986; Hasle & Syvertsen, 1997;
Throndsen et al., 2007).
Proboscia alata (Brightwell) Sundström

Plate 1: a, b
Pavillard, 1925: Fig. 38 (as Rhizosolenia alata); Hustedt, 1930: Fig. 344 (as Rhizosolenia alata); Allen & Cupp, 1935: Fig.
43 (as Rhizosolenia alata); Cupp, 1943: Fig. 52A (as Rhizosolenia alata); Crosby & Wood, 1958: Pl. 38: 62a (as Rhizosolenia
alata); Hendey, 1964: Pl. II: 2 (as Rhizosolenia alata f. alata); Simonsen, 1974: p. 28 (as Rhizosolenia alata); Priddle &
Fryxell, 1985: p. 76-77; Ricard, 1987: Figs. 202-205 (as Rhizosolenia alata f. alata); Hasle & Syvertsen, 1997: Pl. 30;
Sunesen & Sar, 2007a: Figs. 82-88, 98; Throndsen et al., 2007: p. 157; Hoppenrath et al., 2009: Fig. 28 a-d; Yun & Lee,
2011: Fig. 1; Lobban et al., 2012: Pl. 9: 1, 2.
Synonymized names:
Rhizosolenia alata Brightwell 1858
Rhizosolenia alata var. gracillima (Cleve) Grunow ex Van Heurck 1880
Rhizosolenia gracillima Cleve 1881
Rhizosolenia alata f. gracillima (Cleve) Grunow 1882
Rhizosolenia alata f. gracillima (Cleve) Gran 1905
Rhizosolenia alata f. genuina Gran 1908
Cells are solitary or occur in short 2-4-celled chains, narrow cylindrical, bilaterally symmetrical, and circular
in cross section, 380-550 µm long, 17-20 µm in diameter. Valves are subconical, asymmetrical, tapering to
an eccentric, tube-like and oblique process (proboscis) with truncate tip; with depression at base of tube
into which apex of adjoining cell fits. Contiguous area is convex towards the valve surface, distally limited
by asymmetric claspers. Intercalary bands are scale-like, rhombic, arranged in two dorsoventrally rows.
Cell wall is thin, weakly silicified, finely striated, 21-23 puncta in 10 µm. Chloroplasts are numerous, small,
golden-brown, discoid.
Distribution:
The biogeographical limits of Proboscia alata are not clear because synonyms were used for probably
all taxa included in the genus (Sundström, 1986). The species has been considered as distributed from
tropical to subtropical waters by Hernández-Becerril (1995). However, Jordan and Ligowski (2004) stated
that Proboscia alata is not cosmopolitan, being represented by a complex cryptic species with different
geographic distribution. The species has been previously reported (as Rhizosolenia alata) from the Arabian
Gulf (Simonsen, 1974) and from Kuwait’s waters (Al-Yamani et al., 2004; Al-Kandari et al., 2009).
COSCINODISCOPHYCEAE
Local occurrence:
Proboscia alata is of low occurrence in Kuwait’s waters, being recorded from less than 3% of all collected
samples. Its records were mostly restricted to the period from December to April. The species was most
abundant (1.1-1.5∙103 cells/l) in the southern inshore waters off Khiran in January-March 2008.
Harmful Effect:
Proboscia alata is known to form nontoxic high biomass harmful blooms, which have been implicated to
mortality of fish, shrimp, and shellfish resulting in the reduction of fish mariculture outputs. Harmful
effects associated with blooms of Proboscia alata have been recorded in 1983 within South China Sea
(Yan et al., 2002). Bloom of this species causing pale brown water discoloration has been reported from
the coastal Indian waters off Bekal in 2009 (Thomas et al., 2014).
Proboscia indica (Péragallo) Hernández-Becerril

Plate 1: c-i
Pavillard, 1925: Fig. 39 (as Rhizosolenia indica); Hustedt, 1930: Fig. 346 (as Rhizosolenia alata f. indica); Allen & Cupp,
1935: Fig. 45 (as Rhizosolenia alata f. indica); Cupp, 1943: Fig. 52C (as Rhizosolenia alata f. indica); Crosby & Wood, 1958:
Pl. 38: 62c (as Rhizosolenia alata f. indica); Hendey, 1964: Pl. II: 4 (as Rhizosolenia alata f. indica); Simonsen, 1974: p. 28
(as Rhizosolenia alata f. indica); Ricard, 1987: Figs. 200, 201 (as Rhizosolenia alata f. indica); Sunesen & Sar, 2007a: Figs.
89-97, 99; Hoppenrath et al., 2009: Fig. 28 e-l; Yun & Lee, 2011: Fig. 2.
Synonymized names:
Rhizosolenia indica Péragallo 1892
Rhizosolenia alata f. indica (Péragallo) Ostenfeld 1901
Rhizosolenia alata f. indica (Péragallo) Gran 1905
Cells are solitary or occur in short 2-4-celled chains, cylindrical, bilaterally symmetrical, circular in cross
section, 390-710 µm long, 45-50 µm in diameter. Valves are subconical, asymmetrical, tapering to an
eccentric, tube-like and strongly curved process (proboscis); with depression at base of tube into which apex
of adjoining cell fits. Contiguous area is convex towards the valve surface, distally limited by asymmetric
claspers. Intercalary bands are scale-like, rhombic, arranged in columns. Cell wall is finely striated,
longitudinal striae are converging towards the apex. Numerous chloroplasts are small, golden-brown, and
discoid.
Distribution:
Proboscia indica is a commonly reported species from warm temperate to subtropical and tropical waters
(e.g., Hendey, 1964; Simonsen, 1974; Guiry & Guiry, 2017). It has been previously reported from the
Arabian Gulf (Simonsen, 1974) and from Kuwait’s waters (Al-Kandari et al., 2009 as Proboscia alata f.
indica).
Local occurrence:
Proboscia indica is one of the most important species with respect of its contribution to the phytoplankton
standing crop (Polikarpov et al., 2009). This species was frequently observed across Kuwait’s waters
throughout the year, being recorded from 49% of all collected samples. The species was most abundant
(0.6-2.1∙104 cells/l) during the period from December to March.
Plate 1. Proboscia spp.
a, b – Proboscia alata: detail of cell ends; c-i – Proboscia indica: c – complete live cell; d, e – detail of cell end;
f – collapsed girdle segments; g – valve in ventral view showing the contiguous area; h – isolated intercalary
band; i – detail of intercalary band showing areolation.
LM, BF illumination (a, c-f), phase contrast (b); SEM (g-i); scale bars: a, b, f – 20 μm; c-e – 25 μm; g-i – 10 μm.
COSCINODISCOPHYCEAE
Family Rhizosoleniaceae De Toni
Genus Dactyliosolen Castracane
Species of the genus Dactyliosolen are characterized by the presence of numerous half-collar-shaped
intercalary bands, somewhat spirally twisted and with their wedge-shaped thin ends fitting together in a
tooth-like manner. The ends of the intercalary bands do not always lie in an even line in the pervalvar
direction, but sometimes form a spiral line. Species within the genus are distinguished by shape of colonies,
cells and valves, and by morphology of intercalary bands (Hasle & Syvertsen, 1997; Throndsen et al., 2007;
Yun & Lee, 2011).
Dactyliosolen phuketensis (Sundström) Hasle

Plate 2: g, h
Hasle & Syvertsen, 1997: Pl. 31; Throndsen et al., 2007: p. 155; Yun & Lee, 2011: Fig. 6 E-G.
Synonymized names:
Rhizosolenia phuketensis Sundström 1980
Cells are cylindrical, bilaterally symmetrical, occur solitary or in curved chains, 85-94 µm long, 38-42 µm in
diameter. Valves are slightly convex with obtuse, short tube-like external process in the valve marginal part.
Girdle bands are well visible with LM in water mounts and composed of half bands with poroid areolae.
Numerous chloroplasts are small, golden-brown, and elliptical.
,
Distribution:
Dactyliosolen phuketensis occurs in temperate to warm water regions (Sundström, 1986; Hasle & Syvertsen,
1997; Throndsen et al., 2007).
Local occurrence:
Dactyliosolen phuketensis is recorded for the first time in Kuwait’s waters, however, its occurrence and
seasonality was not defined as the species could have been misidentified with Guinardia striata (see below)
during the previous routine phytoplankton surveys.
Genus Guinardia Péragallo
Species of the genus Guinardia are characterized by more or less long cylindrical cells that often occur
in straight to twisted close-set chains, with numerous ligulate intercalary bands usually distinct with LM,
and with circular valves bearing an asymmetrical external tube-like process. Species within the genus are
distinguished by the shape of cells (straight or curved), shape of chains (loose or close set), shape of valves
(flat or convex), and by shape and location of external process (central or marginal) (Cupp, 1943; Hasle &
Syvertsen, 1997; Throndsen et al., 2007).
Guinardia delicatula (Cleve) Hasle

Plate 2: a-f
Pavillard, 1925: Fig. 40 (as Rhizosolenia delicatula); Hustedt, 1930: Fig. 328 (as Rhizosolenia delicatula); Cupp, 1943: Fig.
44 (as Rhizosolenia delicatula); Crosby & Wood, 1958: Pl. 39: 71 (as Rhizosolenia delicatula); Hendey, 1964: Pl. IV: 2 (as
Rhizosolenia delicatula); Hasle & Syvertsen, 1997: Pl. 31; Throndsen et al., 2007: p. 156; Hoppenrath et al., 2009: Fig.
29 i-k; Yun & Lee, 2011: Fig. 5 A, B.
Synonymized names:
Rhizosolenia delicatula Cleve 1900
Cells are small, cylindrical, and bilaterally symmetrical, commonly occur in closely set, straight chains, 22-
27 µm long, 11-17 µm in diameter. Intercalary bands are usually difficult to see with LM. Valves are flat,
rounded only slightly on margins, bearing an asymmetrical, thin and short lateral external tube-like process
at the valve margin that is oblique to the pervalvar axis. In chain, external process fits into a depression on
the adjacent valve. The cytoplasm contains several (usually 2-4) plate-shaped chloroplasts with crenated
margins, arranged along the girdle wall.
Plate 2. Guinardia delicatula and Dactyliosolen phuketensis.
a-f – Guinardia delicatula: a – live single cell and 2-celled chain showing external process in valve marginal
part (arrowhead); b-d – detail of chains; e, f – 2-celled chain at two focal planes with external processes
(arrowheads); g, h – Dactyliosolen phuketensis: complete live cell at two different focal planes, arrowhead points
to external process.
LM, BF illumination; scale bars: 10 μm.
COSCINODISCOPHYCEAE
Distribution:
Guinardia delicatula is a neritic cosmopolitan species widely distributed in temperate to tropical waters
(Hasle & Syvertsen, 1997; Throndsen et al., 2007). The species has been previously reported from
Kuwait’s waters (Al-Kandari et al., 2009).
Local occurrence:
Guinardia delicatula was frequently observed across Kuwait’s waters throughout the year, being recorded
from 49% of all collected samples. The species was most abundant (0.5-1.5∙105 cells/l) in Kuwait Bay
during the spring months of April and May.
Harmful Effect:
Guinardia delicatula is known to form nontoxic high biomass blooms worldwide. No harmful incident has
been reported, but species may be regarded as potentially harmful during its blooms (Fryxell & Hasle,
2004).
Guinardia flaccida (Castracane) Péragallo

Plate 3
Pavillard, 1925: Fig. 36; Hustedt, 1930: Fig. 322; Allen & Cupp, 1935: Fig. 28; Cupp, 1943: Fig. 40; Crosby & Wood,
1958: Pl. 39: 79; Hendey, 1964: Pl. V: 5; Simonsen, 1974: p. 30; Ricard, 1987: Figs. 241-249; Hasle & Syvertsen, 1997:
Pl. 32; Throndsen et al., 2007: p. 156; Hoppenrath et al., 2009: Fig. 29 l, m; Yun & Lee, 2011: Fig. 5 C-E.
Synonymized names:
Rhizosolenia flaccida Castracane 1886
Rhizosolenia castracanei Cleve 1889
Henseniella baltica Schütt 1894
Guinardia baltica Schütt 1896
Cells are large, cylindrical, one and a half to several times longer than broad, occur solitary or united in
straight or slightly curved close chains by whole valve surface, 60-75 µm long, 33-44 µm in diameter. Valves
are nearly flat or very slightly concave, with short tube-shaped external processes that are not visible with
LM. Girdle segments are numerous, composed of open bands with poroid areolae. Nucleus is usually lying
in a central plasma mass, suspended by cytoplasmic strands extending to the cell walls. The cytoplasm
contains numerous small, round to asymmetrically star-shaped chloroplasts lying near the cell wall.
Distribution:
Guinardia flaccida is a cosmopolitan neritic species widely distributed from temperate to tropical waters
(Cupp, 1943; Hasle & Syvertsen, 1997). The species has been previously reported from the Arabian Gulf
(Simonsen, 1974) and from Kuwait’s waters (Al-Yamani et al., 2004; Al-Kandari et al., 2009).
Local occurrence:
Guinardia flaccida is one of the most important species with respect to its contribution to the phytoplankton
standing crop (Polikarpov et al., 2009). This species was commonly observed throughout Kuwait’s
waters, being recorded from 63% of all collected samples. Guinardia flaccida was most abundant (1.1-
19.7∙104 cells/l) in the northern waters during the winter season in January and February. A bloom of
this species (0.8∙106 cells/l) was recorded in the coastal waters of Bubiyan Island in February 2005 (Al-
Yamani et al., 2012).
Harmful Effect:
Guinardia flaccida is known to form nontoxic high biomass harmful blooms in South China Sea, which
have been implicated to mortality of fish and marine invertebrates, resulting in the reduction of fish
mariculture output and tourism (Yan et al., 2002). Due to its capacity to form blooms, Guinardia flaccida
has been included in the list of potentially harmful species for Kuwait’s marine environment (Al-Yamani
et al., 2012).
Plate 3. Guinardia flaccida.
a, b – live cells connected to long chains; c – single cell in girdle view; d, e – detailed girdle bands and
chloroplasts.
LM, BF illumination (a-c), DIC optic (d, e); scale bars: a, b – 25 μm; c-e – 15 μm.
COSCINODISCOPHYCEAE
Guinardia striata (Stolterfoth) Hasle
Plate 4
Pavillard, 1925: Fig. 42 (as Rhizosolenia stolterfothii); Hustedt, 1930: Fig. 329 (as Rhizosolenia stolterfothii); Allen & Cupp,
1935: Fig. 29 (as Rhizosolenia stolterfothii); Cupp, 1943: Fig. 45 (as Rhizosolenia stolterfothii); Crosby & Wood, 1958: Pl.
39: 77 (as Rhizosolenis stolterfothii); Hendey, 1964: Pl. IV: 5 (as Rhizosolenia stolterfothii); Ricard, 1987: Figs. 209-211 (as
Rhizosolenis stolterfothii); Hasle & Syvertsen, 1997: Pl. 31; Throndsen et al., 2007: p. 156; Hoppenrath et al., 2009: Fig.
29 n-p; Yun & Lee, 2011: Fig. 5 F-H.
Synonymized names:
Eucampia striata Stolterfoth 1879
Rhizosolenia stolterfothii Péragallo 1888
Cells are large, cylindrical, bilaterally symmetrical, uniformly curved along pervalvar axis, commonly occur
in curved and often spiraling chains, 65-78 µm long, 28-33 µm in diameter. Cell wall is weakly silicified.
Valves are flat and rounded at the edges, bearing a thin, spine-like and hook-shaped to the pervalvar axis
external marginal process, fits into depression on adjacent cell. Girdle bands are numerous, composed of
open bands with tapering ends and poroid areolae. Nucleus is located near the cell wall. The cytoplasm
contains numerous small, round to elliptical chloroplasts lying along the cell wall.
Distribution:
Guinardia striata is a cosmopolitan neritic species widely distributed worldwide from temperate (except
the polar regions) to tropical waters (Hasle & Syvertsen, 1997). The species has been previously reported
(as Rhizosolenia stolterforthii) from Kuwait’s waters (Al-Yamani et al., 2004; Al-Kandari et al., 2009).
Local occurrence:
Guinardia striata was commonly observed across Kuwait’s waters throughout the year, being recorded
from 63% of all collected samples. The species was most abundant (0.5-1.1∙105 cells/l) during the period
from October to December.
Genus Neocalyptrella Hernández-Becerril et Meave del Castillo
Neocalyptrella robusta (Norman ex Ralfs) Hernández-Becerril et Meave

Plate 5: a-d
Pavillard, 1925: Fig. 43 (as Rhizosolenia robusta); Hustedt, 1930: Fig. 330 (as Rhizosolenia robusta); Allen & Cupp, 1935:
Fig. 31 (as Rhizosolenia robusta); Cupp, 1943: Fig. 46 (as Rhizosolenia robusta); Crosby & Wood, 1958: Pl. 39: 75 (as
Rhizosolenia robusta); Hendey, 1964: Pl. II: 5 (as Rhizosolenia robusta); Simonsen, 1974: p. 29 (as Rhizosolenia robusta);
Ricard, 1987: Figs. 212-214 (as Rhizosolenia robusta); Hasle & Syvertsen, 1997: Pl. 30 (as Rhizosolenia robusta); Sunesen
& Sar, 2007a: Figs. 62-67; Throndsen et al., 2007: p. 157; Hoppenrath et al., 2009: Fig. 37 a-c; Yun & Lee, 2011: Fig. 3.
Synonymized names:
Rhizosolenia robusta Norman ex Ralfs 1861
Calyptrella robusta (Norman ex Ralfs) Hernández-Becerril et Meave del Castillo 1996
Cells are large, solitary or sometimes paired, bilaterally symmetrical, elliptical in cross section, crescent
shaped in lateral view and of sigmoid form in dorsiventral view, 420-580 µm long, 100-110 µm wide. Valves
are conoidal with a rounded apex, tapering into a small external cylindrical tube-like process, straightened
towards the distal part and merging with the calyptra structure. Otaria, claspers, and contiguous areas are
absent. Girdle segments are oriented in a straight line and arranged in two dorsiventral columns. Cell wall
is thin, but rather strongly silicified, delicately punctated. Puncta are arranged in three-line, self-crossing
system (quincunx). Nucleus is located close to the concave cell wall. The cytoplasm contains numerous
small, round to elliptical chloroplasts lying along the cell wall.
Plate 4. Guinardia striata.
a-c – 2-celled arc-shaped chains; d – live single cell in girdle view; e-g – detail of chains, arrowheads point to
external marginal processes.
LM, BF illumination (e-g), DIC optic (a, b, d), and phase contrast (c); scale bars: a-e – 20 μm; f, g – 10 μm.
COSCINODISCOPHYCEAE
Plate 5. Neocalyptrella robusta and Pseudosolenia calcar-avis.

a-d – Neocalyptrella robusta: complete live cells in broad (a, b, d) and narrow (c) girdle views; e, f – Pseudosolenia
calcar-avis: e – live cells connected to 2-celled chain; f – detail of chain with contiguous area of the sibling cells.
Distribution:
Neocalyptrella robusta is an oceanic to neritic species widely distributed from temperate to tropical regions,
being more frequently reported from warm waters (Hernández-Becerril & Meave del Castillo, 1996;
Hasle & Syvertsen, 1997; Guiry & Guiry, 2017). The species has been previously reported (as Rhizosolenia
robusta) from the Arabian Gulf (Simonsen, 1974) and from Kuwait’s waters (Al-Yamani et al., 2004; Al-
Kandari et al., 2009).
Local occurrence:
Neocalyptrella robusta is one of the most important species with respect to its contribution to the
phytoplankton standing crop (Polikarpov et al., 2009) due to its large size rather than abundance. This
species was occasionally observed across Kuwait’s waters throughout the year, being recorded from 29%
of all collected samples but never in a large number. The highest abundances of this species (200-818
cells/l) were restricted to the northern inshore waters.
Genus Pseudosolenia Sundström
Pseudosolenia calcar-avis (Schultze) Sundström

Plate 5: e, f
Pavillard, 1925: Fig. 52 (as Rhizosolenia calcar avis); Hustedt, 1930: Fig. 339 (as Rhizosolenia calcar avis); Allen & Cupp,
1935: Fig. 41 (as Rhizosolenia calcar-avis); Cupp, 1943: Fig. 51 (as Rhizosolenia calcar avis); Crosby & Wood, 1958: Pl. 38:
65 (as Rhizosolenia calcar avis); Hendey, 1964: Pl. IV: 3 (as Rhizosolenia calcar-avis); Hendey, 1970: p. 121 (as Rhizosolenia
calcar-avis); Simonsen, 1974: p. 28 (as Rhizosolenia calcar-avis); Hasle & Syvertsen, 1997: Pl. 30; Sunesen & Sar, 2007a:
Figs. 68-81; Throndsen et al., 2007: p. 158; Yun & Lee, 2011: Fig. 4.
Synonymized names:
Rhizosolenia calcar-avis Schultze 1858
Cells are solitary or sometimes paired, elongated, cylindrical, bilaterally symmetrical, and circular in cross
section, 450-640 µm long, 35-48 µm in diameter. Valves are sub-conical, asymmetrical, with the ventral
part slightly longer than the dorsal part, tapering into a prominent claw-shaped curved process, tapered
towards the distal part. Otaria and claspers are absent. Girdle segments are rhomboidal, arranged in two
(in small individuals) or multiples of two columns. Cell wall is thin and weakly silicified, very delicately
punctated. Nucleus is sub-centrally located, suspended by cytoplasmic strands extending to the cell walls.
The cytoplasm contains numerous small elliptical chloroplasts.
Distribution:
Pseudosolenia calcar-avis is a circumglobally distributed oceanic to neritic species (Sundström, 1986)
frequently reported from warm waters and occasionally observed in temperate regions (Hasle &
Syvertsen, 1997). The species has been previously reported (as Rhizosolenia calcar-avis) from the Arabian
Gulf (Simonsen, 1974), and from Kuwait’s shore (Hendey, 1970) and coastal waters (Al-Yamani et al.,
2004; Al-Kandari et al., 2009).
Local occurrence:
Pseudosolenia calcar-avis was occasionally observed in the northern waters of Kuwait but never in large
numbers, being recorded from 8% of all collected samples. The species was most abundant (120-220
cells/l) during the period from September to January.
COSCINODISCOPHYCEAE
Genus Rhizosolenia Brightwell
Species of the genus Rhizosolenia are characterized by cylindrical cells with greatly elongated pervalvar axis,
elliptical or circular in cross section, with very numerous rhombic, trapezium-like, or scale-shaped intercalary
bands usually distinct with LM, and with symmetrically or eccentric cone-shaped or hood-shaped valves,
characteristically terminated by an eccentric process, short or bristle-like elongated, blunt or sharp, solid or
hollow. Species within the genus are distinguished by shape and arrangement of girdle intercalary bands
(segments, copulae), by shape of valve and process, and by shape, position, size and extension of otaria
(pair of membranous wing-shaped costae that occur opposite each other at or near the base of the external
process) (Cupp, 1943; Sundström, 1986; Hasle & Syvertsen, 1997; Throndsen et al., 2007).
Rhizosolenia bergonii Péragallo

Plate 6: a-d
Pavillard, 1925: Fig. 45; Hustedt, 1930: Fig. 327; Cupp, 1943: Fig. 43; Crosby & Wood, 1958: Pl. 38: 64; Hendey, 1964:
Pl. III: 4; Simonsen, 1974: p. 28; Hasle & Syvertsen, 1997: Pl. 29; Sunesen & Sar, 2007a: Figs. 4-15.
Synonymized names:
Rhizosolenia amputata Ostenfeld 1902
Cells are solitary, large, elongated, cylindrical, bilaterally symmetrical, and circular in cross-section, 320-
450 µm long, 45-55 µm in diameter. Valves are acute and conical, gradually tapering into a short, straight,
centrally located apical process, traversed by a canal in center, cut off abruptly at end. Internal structure of
the process is visible by LM, with basal lumen spindle-shaped, abruptly narrowed in a tubular canal and
expanded into a funnel-shaped aperture at the tip. Claspers and otaria are absent. Intercalary bands are
rhomboidal, arranged in several columns, with distinct imbrication lines. Chloroplasts are small, numerous,
round to elliptical, distributed along the entire cell wall, but especially massed in the girdle zone around the
nucleus.
Distribution:
Rhizosolenia bergonii is an oceanic warm water species, distributed from south temperate to tropical
regions (Sundström, 1986; Hasle & Syvertsen, 1997). The species has been previously reported from the
Arabian Gulf (Simonsen, 1974) and from Kuwait’s waters (Al-Yamani et al., 2004; Al-Kandari et al.,
2009).
Local occurrence:
Rhizosolenia bergonii was occasionally observed throughout Kuwait’s waters but never in large numbers,
being recorded from 21% of all collected samples. The species was most abundant (400-840 cells/l)
during the summer month of June.
Rhizosolenia (Pseudosolenia?) cochlea Brun

Plate 6: e-g
Ostenfeld, 1902: Fig. 5 (as Rhizosolenia calcar avis var. cochlea); Simonsen, 1974, p. 29; Santhanam & Krishnamurthy,
1975: Figs. 1, 2.
Synonymized names:
Rhizosolenia calcar-avis var. cochlea (Burn) Ostenfeld 1902
Cells are solitary or occur in short chains, elongated, cylindrical, bilaterally symmetrical, and circular in
cross section, 350-400 µm long, 60-65 µm in diameter. Valves are regularly conical, distinctly curved at apex,
tapering into a prominent claw-shaped curved process, gradually tapered towards the distal part. Otaria and
claspers are absent. Cell wall is thin and weakly silicified. The cytoplasm contains numerous small elliptical
chloroplasts.
Plate 6. Rhizosolenia spp.
a-d – Rhizosolenia bergonii: a, b – complete live cells; c – detail of cell end; d – detailed girdle bands; e-g –
Rhizosolenia cochlea: e – complete live cell; f – detail of cell end; g – detail of chain with contiguous area of
the sibling cells.
LM, BF illumination (a-c, e-h) and DIC optic (d); scale bars: a, b – 50 μm; c-g – 25 μm.
COSCINODISCOPHYCEAE
Distribution:
Rhizosolenia cochlea is a scarcely reported warm water species. It was characterized by Simonsen (1974)
as a tropical (most probably neritic) species typical for the Indian Ocean and the westernmost part
of the Pacific. Rhizosolenia cochlea has been reported from the Indian coastal waters (Santhanam &
Krishnamurthy, 1975), from Asian region including China, Siam, Hong-Kong, and Japan (Santhanam
& Krishnamurthy, 1975; Liu, 2008). The species has been previously reported from the Arabian Gulf
(Simonsen, 1974), and from Iraqi (Maulood et al., 2013) and Kuwait’s waters (Al-Yamani et al., 2004;
Al-Kandari et al., 2009).
Local occurrence:
Rhizosolenia cochlea is one of the most important species with respect to its contribution to the
phytoplankton standing crop (Polikarpov et al., 2009). This species was frequently observed across
Kuwait’s waters throughout the year, being recorded from 47% of all collected samples. The highest
abundances (1.1-4.3∙103 cells/l) were restricted to the northern inshore waters around Bubiyan Island.
Rhizosolenia hyalina Ostenfeld

Plate 7
Hasle & Syvertsen, 1997: Pl. 28; Sunesen & Sar, 2007a: Figs. 16-24.
Synonymized names:
Rhizosolenia pellucida Cleve 1901
Cells are solitary or occur in more or less short straight two-eight-celled chains, elongated, cylindrical,
bilaterally symmetrical, and circular in cross section, 180-215 µm long, 40-47 µm in diameter. Valves are
sub-conical, asymmetrical, with the ventral part longer than the dorsal part and characteristically undulated
outline, tapering into a thin, long, and slightly curved process, tapering needle-shaped towards the distal
part. Otaria are narrow, extend along the process, rounded at the tip, parallel to long axis of process. Girdle
segment are numerous, rhomboidal to trapezoidal. The nucleus is sub-centrally located, elongated in a girdle
view. The cytoplasm contains numerous small round chloroplasts.
Distribution:
Rhizosolenia hyalina is a warm water species (Sundström, 1986; Hasle & Syvertsen, 1997) scarcely
reported from subtropical and tropical areas of the Atlantic and Pacific Oceans (Guiry & Guiry, 2017).
Local occurrence:
Rhizosolenia hyalina is recorded for the first time in Kuwait’s waters. This species was occasionally
observed across Kuwait’s waters throughout the year, being recorded from 11% of all collected samples
but never in large numbers. The highest abundances of this species (100-342 cells/l) were restricted to
the southern waters during the summer month of June.
Rhizosolenia imbricata Brightwell

Plates 8; 9: e-h
Hustedt, 1930: Fig. 331; Allen & Cupp, 1935: Fig. 35; Crosby & Wood, 1958: Pl. 39: 74a; Hendey, 1964: Pl. III: 1;
Hendey, 1970: p. 121; Simonsen, 1974: p. 29; Ricard, 1987: Figs. 216-218; Hasle & Syvertsen, 1997: Pl. 29; Sunesen &
Sar, 2007a: Figs. 48-61; Throndsen et al., 2007: p. 162; Hoppenrath et al., 2009: Figs. 6, 26 f-l.
Synonymized names:
Rhizosolenia striata Greville 1864
Rhizosolenia shrubsolei Cleve 1881
Rhizosolenia imbricata var. striata (Greville) Grunow 1882
Rhizosolenia imbricata var. shrubsolei (Cleve) Schröder 1906
Cells are long, cylindrical, occur solitary or in short two-six-celled chains, bilaterally symmetrical, and circular
in cross section, 350-420 µm long, 45-58 µm in diameter. Valves are obliquely conical and furnished with a
strong marginal spine-like process, which appears as a continuation of the dorsal side of the valve. Process is
short, swollen at the base and narrowing abruptly towards the distal part. Otaria are small, extending along
Plate 8. Rhizosolenia imbricata. COSCINODISCOPHYCEAE
a – 2-celled chain; b – part of live cell; c – detail of cell end showing process and otaria (arrowheads); d – detail
of chains with contiguous area of the sibling cells; e – detailed girdle bands with striation pattern in empty
frustule in dorsal view; f – valve showing the process with otaria; g – detailed girdle bands with striation
pattern in lateral view.
LM, BF illumination (a-e, g); SEM (f); scale bars: a – 50 μm; b-e, g – 25 μm; f – 10 μm.
Plate 7. Rhizosolenia hyalina.

a, b – live cells united to long chains; c, d – single complete cells in girdle view; e, f – detail of chains with
contiguous area of the sibling cells; g – part of cell; h – detail of cell end showing long process; i – valve
showing the process with otaria.
LM, BF illumination (a-h); SEM (i); scale bars: a, b – 50 μm; c-h – 25 μm; i – 10 μm.
the swollen part of the process. Girdle is composed of numerous intercalary scale-like segments with oblique
ends arranged in two dorsiventral columns. Intercalary scales are coarsely striated; striae are arranged in
oblique rows, converging upon a line or hyaline break which runs up the middle of the girdle. The nucleus
is sub-centrally located, elongated in a girdle view. The cytoplasm contains numerous small plate-shaped
chloroplasts distributed along the entire cell wall.
Remarks:
In taxonomic literature, the smaller form of Rhizosolenia imbricata has been treated as a separate taxon
(Rhizosolenia shrubsolei Cleve) or a variety of Rhizosolenia imbricata (Rhizosolenia imbricata Brightwell v.
shrubsolei (Cleve) Schröder) for a long time (e.g., Hustedt, 1930; Allen & Cupp, 1935; Cupp, 1943; Crosby
& Wood, 1958; Hendey, 1964; Priddle & Fryxell, 1985; Ricard, 1987). However, currently this name is
regarded as a synonym of Rhizosolenia imbricata (Guiry & Guiry, 2017). Rhizosolenia shrubsolei shares
morphological characters with larger Rhizosolenia imbricata except the cell diameter that is less than 40
µm in Rhizosolenia shrubsolei (e.g., Cupp, 1943; Hendey, 1964).
In Kuwait’s waters, both large- and small-sized forms of Rhizosolenia imbricata co-occur (Plates 8 and
9: e-h, respectively; see also Al-Kandari et al., 2009), but never in one sample. In routine phytoplankton
monitoring, the larger and smaller forms of Rhizosolenia imbricata were encountered separately with
respect to their different contribution to the total phytoplankton biomass.
Distribution:
Rhizosolenia imbricata is a cosmopolitan species widely reported worldwide from polar to tropical waters
(Simonsen, 1974; Guiry & Guiry, 2017). The species has been previously reported from the Arabian
Gulf (Simonsen, 1974), and from Kuwait’s shore (Hendey, 1970) and coastal waters (Al-Yamani et al.,
Local occurrence:
Rhizosolenia imbricata is a common species occurring across Kuwait’s waters throughout the year, being
recorded from 38% of all collected samples and attained concentrations up to 1.2-6.2∙103 cells/l. The
smaller form was more frequent (54% of all collected samples) and abundant (1.0-13.8∙103 cells/l),
particularly during the period from October to January.
Rhizosolenia setigera Brightwell

Plate 9: a-d
1958: Pl. 39: 76; Hendey, 1964: Pl. IV: 1; Simonsen, 1974: p. 29; Ricard, 1987: Fig. 226; Hasle & Syvertsen, 1997: Pl.
30; Sunesen & Sar, 2007a: Figs. 25-34; Throndsen et al., 2007: p. 161; Hoppenrath et al., 2009: Fig. 27 d-l.
Synonymized names:
Rhizosolenia japonica Castracane 1886
Rhizosolenia hensenii Schütt 1900
Cells are long, narrow, solitary or occur in pairs, cylindrical, bilaterally symmetrical, circular in cross-
section, 290-347 µm long, 16-19 µm in diameter. Valves are sharply sub-conical, tapering into a very long,
needle-shaped process. The process is almost straight, wider at the base or for some distance from base and
gradually tapering along the apex. Otaria are absent. Girdle segments are rhomboidal and arranged in two
dorsiventral columns. The cytoplasm contains numerous small elliptical chloroplasts.
Remarks:
Rhizosolenia setigera is morphologically closely related to Rhizosolenia hebetata f. semispina, however, the
species is distinguished from R. hebetata by the lack of otaria.
Distribution:
Rhizosolenia setigera is a cosmopolitan species of preferably neritic habitats widely reported worldwide
from temperate to tropical waters (Simonsen, 1974; Hasle & Syvertsen, 1997; Guiry & Guiry, 2017). The
COSCINODISCOPHYCEAE
species has been previously reported from the Arabian Gulf (Simonsen, 1974) and from Kuwait’s waters
(Al-Yamani et al., 2004; Al-Kandari et al., 2009).
Local occurrence:
Rhizosolenia setigera was frequently observed across Kuwait’s waters throughout the year, being recorded
from 39% of all collected samples. This species was most abundant (0.3-4.8∙103 cells/l) during the warm
season from May to October.
Plate 9. Rhizosolenia spp.

a-c – Rhizosolenia setigera: a – complete cell; b – 2-celled chain; c – detail of cell end showing process; d-h
– small-sized Rhizosolenia imbricata (var. schrubsolei): d – detail of chain with contiguous area of the sibling
cells; e, f – detail of cell end in dorsal (e) and lateral (f) view showing process with small otaria (arrowhead); g
– collapsed frustule showing valve with process and intercalary bands, arrowhead points to otaria; h – isolated
intercalary band showing areolation.
LM, BF illumination (b-f) and phase contrast (a); SEM (g, h); scale bars: a, b – 25 μm; c, d, g, h – 10 μm; e,
f – 5 μm.
Order Triceratiales Round et Crawford
Family Triceratiaceae (Schütt) Lemmermann
Genus Triceratium Ehrenberg
Species of the genus Triceratium are characterized by generally tripolar (rarely bi- or tetrapolar) valves with
elevations at the corners bearing ocelli, poroidal areolae with cribra composing the valve structure, mantle
projected outwards in the valvar margin, and simple and finely punctate girdle (Hendey, 1964; Fernandes &
Souza-Mosimann, 2001).
Triceratium dubium Brightwell

Plate 10: h, i
Hustedt, 1930: Fig. 469; Hendey, 1970: Pl. 6: 67; Simonsen, 1974: p. 27; Ricard, 1977: Pl. 9: 11; Foged, 1984: Pls. XXII:
5, XXIII: 6; Podzorski & Håkansson, 1987: Pl. 5: 5, 7; Ricard, 1987: Figs. 421-423; Witkowski et al., 2000: Pl. 8: 4, 5;
Al-Handal, 2009: Figs. 30, 31; Al-Yamani & Saburova, 2011: Pl. 13: f, g.
Synonymized names:
Triceratium bullosum Witt 1873
Triceratium bicorne Cleve 1878
Biddulphia bicornis Cleve 1878
Biddulphia dubia Cleve 1883
Amphitetras bicornis De Toni 1907
Cells are small, solitary, rectangular in girdle view. Valves are hexagonal with undulated margins and concave
sides, and with tree larger corners possessing the ocellii and three smaller corners. The length of side from
ocellus to ocellus varies between 26 and 31 μm. Valve surface is irregularly areolated. Areolae are polygonal
and distinctly poroid, larger at the center and smaller towards the margins, radiating from the center, number
5-5.5 in 10 μm. Girdle is coarsely punctated.
Distribution:
Triceratium dubium is widespread along the coasts of marine warm water areas. The species has been
previously reported from northern neritic habitats of the Arabian Gulf (Simonsen, 1974), from Kuwait’s
Mina Al-Ahmadi shore by Hendey (1970), from the bottom sediments in Shatt Al-Arab Estuary, Iraq
(Al-Handal, 2009), and from Kuwait’s intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Triceratium dubium is a typically benthic species (Al-Yamani & Saburova, 2011), which occasionally
becomes stirred up from the bottom sediments and carried into the water column. Frustules of this
species were sporadically found in plankton net hauls in Kuwait’s inshore waters.
Triceratium robertsianum Greville

Plate 10: a-e
Péragallo & Péragallo, 1897-1908: Pl. CI: 4; Hustedt, 1930: Fig. 466; Crosby & Wood, 1958: Pl. 33: 36; Hendey, 1970:
p. 118; Simonsen, 1974: p. 28; Navarro, 1981: Fig. 62; Foged, 1984: Pl. XXIII: 5; Al-Yamani & Saburova, 2011: Pl. 13
a-c.
Synonymized names:
Biddulphia robertsiana Boyer 1900
Triceratium robertsianum var. macracantha Schmidt 1885
Cells are large, solitary, strongly silicified, and rectangular in girdle view. Valves are triangular with strongly
convex margins. The length of side from ocellus to ocellus varies between 53 and 94 μm. The angles of the
valve are furnished with prominent stout processes. Valve surface is areolated. Areolae are hexagonal and
distinctly poroid, number 2-2.5 in 10 μm. Girdle surface is coarsely punctated. Numerous discoid golden-
brown chloroplasts are arranged alongside the cell wall.
Distribution:
Triceratium robertsianum is a preliminary benthic species widely distributed mainly in the warmer
latitudes of all oceans. It has been previously reported from the Indian Ocean (Simonsen, 1974), from
COSCINODISCOPHYCEAE
Kuwait’s Mina Al-Ahmadi shore (Hendey, 1970), and from Kuwait’s intertidal sediments (Al-Yamani
& Saburova, 2011).
Local occurrence:
Triceratium robertsianum is a typically benthic species (Al-Yamani & Saburova, 2011), which occasionally
becomes stirred up from the bottom sediments and carried into the water column. Frustules of this
species were occasionally found in plankton samples from Kuwait’s inshore waters. Live cells were
observed occasionally (fewer than 200 cells/l) in samples from Kuwait Bay and adjacent waters during
the warm season from April to October.
Plate 10. Triceratium spp. and Schuettia annulata var. minor.

a-e – Triceratium robertsianum: a, b – live cells in valve (a) and girdle (b) view; c, d – acid cleaned frustules in
valve (c) and girdle (d) view; e – detail of valve surface showing areolation; h, i – Triceratium dubium: acid
cleaned frustules in valve (h) and girdle (i) view; f, g – Schuettia annulata var. minor: acid cleaned valve at
different focal planes.
LM, BF illumination (a, b), DIC optic (d, f-i), and phase contrast (c, e); scale bars: a-d – 25 μm; f, g – 20 μm;
e, h – 10 μm.
Subclass Archaegladiopsophycidae Nikolaev et Harwood
Order Stephanopyxales Nikolaev
Family Stephanopyxidaceae Nikolaev
Genus Stephanopyxis (Ehrenberg) Ehrenberg
Species of the genus Stephanopyxis are distinguished by strongly developed pervalvar axes, primarily circular
valve outline, high and more or less curved valve mantle, presence of marginal ring of labiate processes, and
chain formation. Species within the genus differ by the character of valve areolation (Cupp, 1943; Hasle &
Syvertsen, 1997).
Stephanopyxis palmeriana (Greville) Grunow

Plate 11
Pavillard, 1925: Fig. 3; Hustedt, 1930: Fig. 147; Allen & Cupp, 1935: Fig. 2; Cupp, 1943: Fig. 4; Crosby & Wood, 1958:
Pl. 35: 2; Simonsen, 1974: p. 7; Tynni, 1983: Pl. 1: 8, 9; Hoppenrath et al., 2009: Fig. 17 k-o.
Synonymized names:
Creswellia palmeriana Greville 1865
Pyxidicula palmeriana (Greville) Strelnikova et Nikolajev 1986
Cells are oblong, with the slight narrowing of the cylindrical part of the valve against the margin, occur
in more or less short and straight chains, 45-62 µm in diameter. Valves are circular and areolated. Areolae
are hexagonal and smaller near the girdle line than on the rest of the valve, number 1.5-2.5 in 10 µm at the
valve center, 3.5-4 on the upper part of mantle, and 5-5.5 near the girdle line. Cells are united in chain by
10-22 hollow spines arranged in a circle alongside the valve margin. The nucleus is centrally located. The
cytoplasm contains numerous small plate-like chloroplasts.
Distribution:
Stephanopyxis palmeriana is a neritic species widely distributed species in warm water. The species has
been previously reported from the Arabian Gulf (Simonsen, 1974), from the bottom sediments of Shatt
Al-Arab Estuary, Iraq (Tynni, 1983), and from Kuwait’s waters (Al-Kandari et al., 2009).
Local occurrence:
Stephanopyxis palmeriana is of low occurence in Kuwait’s waters throughout the year, being recorded
from less than 3% of all collected samples and never in a large number. The species was most abundant
(170-433 cells/l) during February-March 2005.
COSCINODISCOPHYCEAE
Plate 11. Stephanopyxis palmeriana.

a, b – detail of chains; c – detail of valve surface showing areolation; d-f – collapsed sibling cells showing
marginal ring of hollow spines.
LM, BF illumination (a-d) and phase contrast (e, f); scale bars: a, b – 20 μm; c-f – 10 μm.
Subclass Corethrophycidae Round et Crawford
Order Corethrales Round et Crawford
Family Corethraceae Lebour
Genus Corethron Castracane
Species of the genus Corethron are characterized by dome-shaped valves bearing a crown of long thin spines
or setae at the margin directed outward at an angle (Cupp, 1943; Hasle & Syvertsen, 1997).
Corethron histrix Hensen

Plate 12
Hustedt, 1930: Fig. 311; Cupp, 1943: Fig. 34 A-C; Throndsen et al., 2007: p. 146; Hoppenrath et al., 2009: Fig. 30 a-h.
Cells are usually solitary, small, cylindrical, weakly silicified, with arched hemispherical valves, 28-33 µm
long (pervalvar axis), 19-21 µm in diameter. Valves are convex, circular, possessing a circle of long slender
setae alongside the edge of valve radiating out from center of cell. After cell division, setae are enclosed
within the girdle zone, parallel to the pervalvar axis. Setae differ in morphology and orientation; longer
barbed setae are orientated downwards and shorter hooked setae are orientated upwards. Girdle is composed
of numerous intercalary bands which are barely visible with LM. The cytoplasm contains numerous small
elongated chloroplasts.
Distribution:
Corethron histrix is a temperate oceanic species (Cupp, 1943; Throndsen et al., 2007; Guiry & Guiry,
2017).
Local occurrence:
Corethron histrix was occasionally recorded at inshore localities in the northern Kuwait’s waters around
Bubiyan Island but was rare elsewhere, being found from 22% of all collected samples. The highest
abundances of this species (0.3-3.7∙103 cells/l) were observed during the period from November to
February.
COSCINODISCOPHYCEAE
Plate 12. Corethron histrix.

a, d-f – live cells in oblique (a, e), girdle (d) and valve (f) view; b – two cells after division; c – frustule in girdle
view; g, h – valves in oblique (g) and apical (h) view showing long marginal setae.
LM, BF illumination (a, b, d, e), DIC optic (f), and phase contrast (c, g, h); scale bars: a-c, g, h – 10 μm; d-f
– 20 μm.
Subclass Coscinodiscophycidae Round et Crawford
Order Asterolamprales Round
Family Asterolampraceae Smith
Genus Asteromphalus Ehrenberg
Species of the genus Asteromphalus are characterized by partially areolated valve surface with hyaline rays
extending from a nonareolate central area and terminate short of the valve margin in a labiate process
(rimoportula), with one of the hyaline rays narrower than the others, and by characteristic pattern of the
central area traversed by a number of straight, zigzagged, or branched lines. Species within the genus are
distinguished by valve outline, number and shape of hyaline rays, position, shape, size and pattern of central
area, shape of areolated sectors, and size of areolae (Cupp, 1943; Hasle & Syvertsen, 1997).
Asteromphalus cleveanus Grunow

Plate 13
Allen & Cupp, 1935: Fig. 23; Hendey, 1970: Pl. 4: 42.
Cells are solitary, small, undulate in girdle view and oviform in valve view, the major axis is 32-44 µm,
and the minor axis is 26-29 µm. There are six to nine hyaline radial rays alternating with areolated sectors
and terminating at the margin in a distinct opening. The ordinary rays are straight, slightly wider at their
base than at the valve margin. The singular ray is much thinner than the ordinary rays. Areolae are small,
hexagonal, arranged in radial rows, number 16-17 in 10 µm. The cribra pattern is a hexagonal array of
slits around each central pore. The central area is hyaline, covers about half the minor axis, divided by the
straight separating lines. The cytoplasm contains numerous plate-shaped ellipsoid chloroplasts.
Distribution:
Asteromphalus cleveanus is a neritic warm water species primarily distributed in the Indian and Pacific
Ocean, but rarely recorded in the Atlantic. The species has been previously reported from Kuwait’s
shores (Hendey, 1970) and Iraqi waters (Maulood et al., 2013)
Local occurrence:
Asteromphalus cleveanus is a rare species in Kuwait’s waters, being recorded from less than 1% of all
collected samples, and never in large numbers.
Asteromphalus flabellatus (Brébisson) Greville

Plate 14: a-c
Pavillard, 1925: Fig. 27; Hustedt, 1930: Fig. 279; Allen & Cupp, 1935: Fig. 22; Hendey, 1970: p. 115; Simonsen, 1974:
p. 25; Ricard, 1987: Fig. 58; Hasle & Syvertsen, 1997: Pl. 24.
Synonymized names:
Spatangidium flabellatum Brébisson 1857
Cells are solitary, small, undulate in girdle view and almost circular in valve view, 53-65 µm in diameter.
There are 9-11 hyaline radial rays alternating with areolated sectors and terminating at the margin in a
distinct opening. The ordinary rays are straight, club-shaped, and wider at their base than at the valve margin.
The singular ray is markedly narrower than the ordinary rays. Areolae are small, hexagonal, arranged in
radial rows, number 16-18 in 10 µm. The central area is hyaline, slightly eccentric, covers about 0.4-0.6 the
diameter, divided by the straight separating lines. The cytoplasm contains numerous plate-shaped ellipsoid
chloroplasts with slightly crenated edges.
Distribution:
Asteromphalus flabellatus is a warm water species with predominant occurrence in subtropical and tropical
regions (Simonsen, 1974). The species has been previously reported from the Arabian Gulf (Simonsen,
1974), and from Kuwait’s shores (Hendey, 1970) and coastal waters (Al-Kandari et al., 2009).
COSCINODISCOPHYCEAE
Plate 13. Asteromphalus cleveanus.

a – live cell in valve view; b, c – frustules in girdle view; d – internal valve view; e – external valve view; f, g –
detail of valve showing morphology of cribra and ray holes; h – internal surface of valve showing radial ray.
LM, BF illumination (a); SEM (b-h); scale bars: a, b, d, e – 10 μm; c, f-h – 5 μm.
Local occurrence:
Asteromphalus flabellatus was occasionally recorded across Kuwait’s waters throughout the year, being
found from 12% of all collected samples, but never in large numbers. The species was found to be more
abundant (120-360 cells/l) during the period from October to December in Kuwait Bay and adjacent
waters.
Asteromphalus heptactis (Brébisson) Ralfs

Plate 15
Hustedt, 1930: Fig. 277; Cupp, 1943: Fig. 32; Hendey, 1964: Pl. XXIV: 5; Hendey, 1970: p. 115; Simonsen, 1974: p. 25;
Priddle & Fryxell, 1985: p. 116-117 (A, E); Ricard, 1987: Figs. 54-57.
Synonymized names:
Spantangidium heptactis Brébisson 1857
Spatangidium ralfsianum Norman 1859
Asteromphalus reticulatus Cleve 1873
Asterolampra ralfsiana (Norman) Grunow 1876
Asterolampra areolata A. Mann 1925
Cells are solitary, small, undulate in girdle view and oviform in valve view, the major axis is 40-43 µm,
and the minor axis is 35-38 µm. There are seven hyaline radial rays alternating with areolated sectors and
terminating at the margin in a distinct opening. The ordinary rays are straight, club-shaped. The singular ray
is markedly narrower than the ordinary rays. Areolae are large, hexagonal, arranged in radial rows, number
6-6.5 in 10 µm. The central area is hyaline, covers about quarter to third the minor axis, divided by bent
separating lines. The cytoplasm contains numerous plate-shaped chloroplasts.
Distribution:
Asteromphalus heptactis is a cosmopolitan oceanic species widely distributed worldwide (Simonsen, 1974;
Guiry & Guiry, 2017). It has been previously reported from the Arabian Gulf (Simonsen, 1974), and
from Kuwait’s shores (Hendey, 1970) and coastal waters (Al-Yamani et al., 2004).
Local occurrence:
Asteromphalus heptactis was occasionally recorded across Kuwait’s waters throughout the year, being found
from 10% of all collected samples, generally fewer than 100 cells/l. The species was more abundant (0.3-
3.2∙103 cells/l) during the period from July to November in the inshore waters.
Asteromphalus sarcophagus Wallich

Plate 14: d-f
Simonsen, 1974: p. 22: 3-6; Semina, 2003: Pl. 27: 3, 4.
Cells are solitary, small, obovate in valve view and slightly asymmetrical, the major axis is 27-28 μm, and the
minor axis is 11-11.5 μm. There are six hyaline radial rays alternating with areolated sectors and terminating
at the margin in a distinct opening. The arrangement of rays is asymmetrical. Two longest rays extend
the central length of valve and weakly sigmoid. Lateral rays are much shorter, curved towards the valve
periphery at their distal ends. Areolae are large, irregularly round, number 10-11 in 10 μm.
Distribution:
Asteromphalus sarcophagus is scarsely reported marine warm water species (Simonsen, 1974; Guiry
& Guiry, 2017). It has been previously reported from the Arabian Sea off the southern Indian coast
(Simonsen, 1974).
Local occurrence:
Asteromphalus sarcophagus is recorded for the first time in Kuwait’s waters. Only two specimens were
found from the samples collected in Kuwait’s inshore southern waters during the spring of 2008.
Plate 14. Asteromphalus spp. COSCINODISCOPHYCEAE
a-c – Asteromphalus flabellatus: a – live cell in valve view; b, c – acid cleaned valves; d-f – Asteromphalus
sarcophagus: d – two valves in external view; e – valve in internal view; f – detail of internal valve surface
showing morphology of cribra and radial rays.
LM, BF illumination (a) and phase contrast (b, c); SEM (d-f); scale bars: a-e – 10 μm; f – 5 μm.
Plate 15. Asteromphalus heptactis.
a, b – live cell in valve (a) and girdle (b) view; c – acid cleaned valve; d – frustule in valve view; e – collapsed
valve; f – detail of valve surface showing morphology of cribra and radial ray.
LM, BF illumination (a, b) and phase contrast (c, e); SEM (d, f); scale bars: a-e – 10 μm; f – 5 μm.
COSCINODISCOPHYCEAE
Order Coscinodiscales Round
Family Aulacodiscaceae (Schütt) Lemmermann
Genus Schuettia De Toni
Schuettia annulata var. minor (Grunow) De Toni

Plate 10: f, g
Van Heurck, 1883: Pl. 124: 13; Hendey, 1957: Pl. III: 3 (as Schuettia annulata); Simonsen, 1974: Pl. 20: 6; Podzorski
& Håkansson, 1987: Pl. 3: 2 (as Actinoptychus annulatum); Yim & Li, 2000: Fig 3a: 13 (as Actinoptychus annulatus); Al-
Handal, 2009: Fig. 29 (as Symbolophora cf. trinitatis Ehrenberg); Lee & Park, 2015: Fig. 9.
Synonymized names:
Actinoptychus annulatus var. minor Grunow ex Van Heurck 1883
Triceratium annulatum Wallich 1858
Actinoptychus annulatus (Wallich) Grunow ex Van Heurck 1883
Cymatogonia annulata (Wallich) Grunow 1883
Schuettia annulata (Wallich) De Toni 1894
Cymatogonia annulata (Wallich) Millis 1934
Valves are triangular with broadly rounded apices and concave sides. The length of side from angle to angle
is 37-38 μm. The angles of the valve are furnished with small short tube-like processes (rimoportulae). Each
angle bears a prominent mid-rib or sectorial fold lying from the valve center to the margin. Valve surface
is undulated and finely areolated. Areolae are arranged in rows following valve undulation, 8-9 in 10 μm.
Central area is hyaline surrounded by hyaline concentric ridges.
Distribution:
Schuettia annulata and its variety are warm water taxa scarcely reported in taxonomic literature. Based on
documented records of these diatoms, Simonsen (1974) has restricted their distribution to the tropical
regions of the West Pacific and Indian Ocean. The species/variety has been reported from the Yellow
Sea off Korea, South China Sea, Taiwan, Philippines, Java, Sumatra, Ganges Delta, Indian West coast,
West African shore, Gulf of Suez, and Strait of Hormuz (Hendey, 1957; Simonsen, 1974; Podzorski &
Håkansson, 1987; Yim & Li, 2000; Lee & Park, 2015; Guiry & Guiry, 2017). In the Arabian Gulf, Al-
Handal (2009) isolated this species from the Shatt Al-Arab Estuary, Iraq (Al-Handal, 2009: Fig. 29), but
identified it as Symbolophora cf. trinitatis Ehrenberg.
Local occurrence:
Schuettia annulata var. minor is recorded for the first time in Kuwait’s waters. The species is very rare;
several frustules were found in plankton hauls from Kuwait’s northern waters in the vicinity of Bubiyan
Island.
Family Coscinodiscaceae Kützing
Genus Coscinodiscus Ehrenberg
The genus Coscinodiscus is regarded as one of the largest and diverse marine planktonic diatom genera
with more than 400 validly described taxa (VanLandingham, 1968; Hasle & Syvertsen, 1997). The recent
taxonomic revisions resulted in great number of taxa formerly assigned to Coscinodiscus were transferred into
other genera including Thalassiosira, Actinocyclus, and Azpeitia, however, despite this reduction, Coscinodiscus
continues to be characterized by a considerable diversity with more than 150 taxa (Hasle & Syvertsen, 1997;
Guiry & Guiry, 2017).
The species belonging to the genus Coscinodiscus are characterized by circular valves with radial pattern of
areolae and presence of marginal ring of small labiate processes (rimoportulae) alongside the valve margin
with two larger macroprocesses (macrorimoportulae) located asymmetrically at an angle of more than 90°
and less than 180°. Species within the genus are distinguished by size and shape of cells in valve and girdle
view, size and pattern of areolae, presence and structure of central area, presence and width of hyaline
interstriae between rows of areolae, size, shape and position of macrorimoportulae, distance between small
marginal rimoportulae, and presence of rimoportulae on valve surface (Cupp, 1943; Hasle & Syvertsen,
1997).
Some species of Coscinodiscus may be identified by their gross morphology as seen in water mounts,
however, reliable identification of most species requires permanent mounts of acid cleaned frustules
combined with phase or differential interference contrast (DIC) optic or electron microscopy tools.
Coscinodiscus asteromphalus Ehrenberg

Plate 16
Hustedt, 1930: Fig. 250; Allen & Cupp, 1935: Fig. 14; Hendey, 1964: Pl. XXIV: 2; Hendey, 1970: p. 111; Simonsen,
1974: p. 14; Tynni, 1983: Pl. 4: 34, 35; Priddle & Fryxell, 1985: p. 128-129; Hasle & Syvertsen, 1997: Pl. 15.
Synonymized names:
Coscinodiscus radiatus var. asteromphalus (Ehrenberg) Ehrenberg 1854
Coscinodiscus asteromphalus var. conspicua Grunow 1883
Coscinodiscus asteromphalus var. genuina Grunow 1884
Cells are large, solitary, and circular in valve view, 280-320 µm in diameter. Valves are slightly convex, with
a small central depression and gently sloping mantle. Valve surface is strongly areolated, with a distinct
central rosette composed of larger areolae enclosing a small hyaline area. Areolae are large, polygonal, with
cribra, arranged in radial rows with secondary spiral rows, somewhat unequal, attaining their maximum
size at distance equal to half the radius, 3-4.5 in 10 µm in the center, 5-6 in 10 µm at the margin. Marginal
rimoportulae are barely visible with LM. Two macrorimoportulae are rather small, located about 120° apart.
Chloroplasts are numerous, plate-like, more densely arranged along the girdle.
Distribution:
Coscinodiscus asteromphalus is a cosmopolitan species widely distributed from temperate to tropical waters
worldwide. It has been previously reported from the Arabian Gulf (Simonsen, 1974), from the Shatt
Al-Arab Estuary, Iraq (Tynni, 1983), and from Kuwait’s shores (Hendey, 1970) and coastal waters (Al-
Kandari et al., 2009).
Local occurrence:
Coscinodiscus asteromphalus was occasionally recorded across Kuwait’s waters throughout the year, being
found from 24% of all collected samples but never in large numbers. The species was found to be more
abundant (0.3-1.6∙103 cells/l) in the northern waters.
COSCINODISCOPHYCEAE
Plate 16. Coscinodiscus asteromphalus.

a – live cell in valve view; b, c – detail of valve; d – acid cleaned frustule in valve view; e – detail of cleaned
valve showing areolation; f – central rosette.
LM, BF illumination (a, b, d, f), DIC optic (c), and phase contrast (e); scale bars: a-d – 50 μm; e, f – 25 μm.
Coscinodiscus gigas Ehrenberg
Plate 17
Hustedt, 1930: Figs. 254-256; Allen & Cupp, 1935: Fig. 16; Simonsen, 1974: p. 16; Tynni, 1983: Pl. 4: 31.
Synonymized names:
Coscinodiscus praetextus Janish 1889
Coscinodiscus gigas var. praetexta (Janisch) Hustedt 1891
Coscinodiscus giganteus Péragallo 1908
Cells are very large, solitary, circular in valve view, relatively weakly silicified, 480-610 µm in diameter.
Valves are almost flat with gently sloping mantle. Valve surface is areolated with the exception of hyaline
central area. Areolae are arranged in radial rows, somewhat unequal is size and structure, smaller (3-3.5 in
10 µm), rounded and less silicified in the center and increase in size towards the periphery (2-2.5 in 10 µm),
where they are distinctly hexagonal and dark, forming a broad definite band, but again become smaller near
the valve margin. Marginal rimoportulae are barely observable with LM. Two macrorimoportulae are rather
small, located about 130° apart. Chloroplasts are numerous, small, plate-like, more or less elongated.
Distribution:
Coscinodiscus gigas is a warm water species reported from subtropical and tropical areas of all oceans
(Simonsen, 1974). The species has been previously reported from the Arabian Gulf as principally neritic
and more abundant in the northern part (Simonsen, 1974) as well as from the sediments of Shatt Al-
Arab Estuary, Iraq (Tynni, 1983).
Local occurrence:
Coscinodiscus gigas is recorded for the first time in Kuwait’s waters. This species was rarely observed in
Kuwait’s waters, being recorded from less than 1% of all collected samples and never in large numbers. Its
occurrence seems to be restricted to Kuwait Bay, where this species was observed in the summer, however,
it could be easily misidentified with Coscinodiscus nobilis during phytoplankton sample examinations.
Coscinodiscus granii Gough

Plate 18
Hustedt, 1930: Fig. 237; Cupp, 1943: Fig. 21; Crosby & Wood, 1958: Pl. 36: 17; Hendey, 1964: p. 79; Simonsen, 1974:
p. 16; Hasle & Syvertsen, 1997: Pl. 17; Throndsen et al., 2007: p. 150; Hoppenrath et al., 2009: Fig. 11 h-t.
Cells are large, solitary, characteristically wedge-shaped in girdle view and circular in valve view, 85-120 µm
in diameter. Valves are convex, areolated, with a distinct central rosette composed of larger areolae. Areolae
are polygonal, arranged in radial rows, about 8 in 10 µm near the center and 10-11 in 10 µm at the margin.
One ring of marginal rimoportulae is clearly visible with LM, 8-11 areolae apart, with distinct hyaline lines
radiating towards the valve center. Two macrorimoportulae are located about 130° apart. Chloroplasts are
numerous, plate-like, slightly elongated.
Distribution:
Coscinodiscus granii is a cosmopolitan neritic species widely distributed worldwide. It has been previously
reported from the Indian Ocean (Simonsen, 1974) and from Kuwait’s waters (Al-Yamani et al., 2004;
Al-Kandari et al., 2009).
Local occurrence:
Coscinodiscus granii occurred frequently across Kuwait’s waters, being recorded from 34% of all collected
samples. The species was present throughout the year in low to moderate abundance with higher
concentrations (1.0-3.5∙103 cells/l) in the inshore waters of Kuwait Bay and around Bubiyan Island
during the period from October to February.
COSCINODISCOPHYCEAE
Plate 17. Coscinodiscus gigas.

a – live cell in valve view; b – detail of valve; c – dividing cell in girdle view; d – detail of valve center; e – detail
of valve margin; f – detail of cleaned valve showing areolation; g – central rosette; h – detail of valve margin.
LM, BF illumination; scale bars: a-c – 100 μm; d, e – 25 μm; f – 50 μm; g, h – 20 μm.
Plate 18. Coscinodiscus granii.
a-c – live cells in oblique (a), valve (b) and girdle (c) view; d – acid cleaned valve; e – detail of cleaned
valve.
LM, BF illumination (a, c, d), DIC optic (b), and phase contrast (e); scale bars: a-c – 25 μm; d, e – 10
μm.
COSCINODISCOPHYCEAE
Coscinodiscus janischii Schmidt var. arafurensis Grunow
Plate 19
Hustedt, 1930: Fig. 257; Allen & Cupp, 1935: Fig. 17.
Cells are large, solitary, 220-245 µm in diameter. Valves are almost flat, areolated with the exception of small
hyaline central area. Areolae are arranged in complete and incomplete radial rows, the latter originating
towards the midradius. Near the center, areolae are elongated in radial direction, becoming hexagonal and
increasing in size towards the periphery, 2.5-3 in 10 µm at the central area, about 2 in 10 µm towards the
periphery, with row of small areoles along the margin, about 4 in 10 µm. Marginal rimoportulae are barely
observable with LM. Two macrorimoportulae are rather small, located about 120-130° apart. Chloroplasts
are numerous, small, plate-like.
Distribution:
Coscinodiscus janischii var. arafurensis is a warm water species distributed in subtropical and tropical areas.
The species was reported from the Arafura Sea, Java Sea, Mozambique Channel, Madagascar, Gulf of
Thailand, Malay Archipelago, and Gulf of Bengal (Ostenfeld, 1902; Allen & Cupp, 1935).
Local occurrence:
Coscinodiscus janischii var. arafurensis is recorded for the first time in Kuwait’s waters. This species was
occasionally found mainly in Kuwait Bay and around Bubiyan Island, being present in 7% of all collected
samples and never in large numbers, fewer than 100 cells/l.
Coscinodiscus marginatus Ehrenberg

Plate 20: a-f
Hustedt, 1930: Fig. 223; Allen & Cupp, 1935: Fig. 7; Cupp, 1943: Fig. 19, Pl. 1: 3; Hendey, 1964: Pl. 22: 2; Hendey,
1970: p. 110; Tynni, 1983: Pl. 3: 30; Hasle & Syvertsen, 1997: Pl. 18; Al-Handal, 2009: Fig. 13.
Synonymized names:
Coscinodiscus limbatus Ehrenberg 1840
Coscinodiscus fimbriatus-limbatus Ehrenberg 1854
Cells are small, solitary, strongly silicified, 42-58 µm in diameter. Valves are almost flat, with steeply sloping
mantle, areolated, without central area. Areolae are coarse, polygonal, arranged in irregular radial rows,
decrease in size towards the periphery, 3-4 in 10 µm. In valve view, the walls of mantle areolae appear as
coarse radial striae on a broad marginal band. Chloroplasts are small, plate-like.
Distribution:
Coscinodiscus marginatus is a cosmopolitan principally oceanic species widely distributed worldwide
(Cupp, 1943; Hendey, 1964; Guiry & Guiry, 2017). The species has been previously reported from the
northern Arabian Gulf, from the bottom sediments of Shatt Al-Arab Estuary, Iraq (Tynni, 1983; Al-
Handal, 2009), and from Kuwait’s shores (Hendey, 1970) and coastal waters (Al-Kandari et al., 2009).
Local occurrence:
Coscinodiscus marginatus was sporadically recorded in Kuwait’s waters. Its occurrence could not be defined
due to possible misidentification with small-sized specimens of Coscinodiscus radiatus that were more
frequent and abundant (see below). The illustrated specimens were isolated from the inshore waters of
Kuwait Bay in the spring.
Plate 19. Coscinodiscus janischii var. arafurensis.
a, b – live cells in valve view; c – detail of valve; d – detail of valve center; e – detail of valve margin; f – central
part of cleaned valve showing areolation and central rosette; g – detail of cleaned valve showing valve margin.
LM, BF illumination (a), DIC optic (b-e), and phase contrast (f, g); scale bars: a, b – 50 μm; c – 25 μm; d,
e – 10 μm; f, g – 20 μm.
COSCINODISCOPHYCEAE
Plate 20. Coscinodiscus spp.

a-f – Coscinodiscus marginatus: a, b – live cells in valve view; c-e – acid cleaned valves; f – detail
of cleaned valve; g-k – Coscinodiscus perforatus: g – live cell in valve view; h – acid cleaned
valve; i – detail of cleaned valve showing scattered small labiate processes (arrowheads); j –
central rosette; k – detail of valve margin showing ring of marginal processes (arrowheads).
LM, BF illumination (a, b, g), DIC optic (c, e, h, j, k), and phase contrast (d, f, i); scale bars:
10 μm.
Coscinodiscus nobilis Grunow
Plate 21
Allen & Cupp, 1935: Fig. 13; Simonsen, 1974: p. 15, Pl. 14.
Cells are large, solitary, drum-shaped in girdle view and circular in valve view, relatively weakly silicified, 220-
420 µm in diameter. Valves are weakly convex with almost flat central part. Valve surface is areolated with the
exception of large hyaline central area, and furnished with complete and incomplete alternating radial rows,
sometimes slightly undulated near central area. Areolae are somewhat unequal is size and structure, smaller
(9.5-10 in 10 µm) and roundish to irregularly polygonal near the center, becoming hexagonal and increasing
in size (6-7.5 in 10 µm) towards the periphery. Distinct small rimoportulae are irregularly scattered on valve
surface, from which hyaline ribs radiate towards the center. The ring of small marginal rimoportulae is
barely observable with LM. Two macrorimoportulae are rather small, located about 120° apart. The girdle
is composed of intercalary bands. Chloroplasts are numerous, small, plate-like, more or less elongated.
Distribution:
Coscinodiscus nobilis is a scarcely reported species known from temperate to tropical waters mainly from
Pacific and Indian Ocean (Allen & Cupp, 1935; Sournia, 1968; Guiry & Guiry, 2017).
Local occurrence:
Coscinodiscus nobilis is reported for the first time from Kuwait’s waters. The species occurred sporadically,
and its distribution and seasonality could not be defined due to possible misidentification with
morphologically similar large-sized Coscinodiscus gigas during routine phytoplankton sample analysis.
The illustrated specimens were isolated from the southern offshore waters in the vicinity of Kubbar
Island in the summer.
Coscinodiscus oculus-iridis Ehrenberg

Plate 22
Hustedt, 1930: Figs. 252, 253; Allen & Cupp, 1935: Fig. 15; Cupp, 1943: p. 63, Fig. 26, Pl. 3: 2; Hendey, 1964: p. 78,
Pl. XXIV: 1; Hendey, 1970: p. 111; Simonsen, 1974: p. 17; Tynni, 1983: Pls. 4: 36, 5: 37; Priddle & Fryxell, 1985: p.
138-139; Al-Handal, 2009: Fig. 3.
Synonymized names:
Coscinodiscus radiatus var. oculus-iridis Ehrenberg 1840
Coscinodiscus radiatus var. oculus-iridis (Ehrenberg; Ehrenberg) Van Heurck 1896
Coscinodiscus radiatus var. oculus-iridis (Ehrenberg) Jørgensen 1905
Coscinodiscus oculus-iridis var. genuina Grunow 1884
Coscinodiscus oculus-iridis var. typicus Cleve 1942
Cells are large, solitary, and circular in valve view, 110-140 µm in diameter. Valves are weakly convex. Valve
surface is strongly areolated, with a distinct central rosette composed of enlarged areolae. Areolae are large,
polygonal, with cribra, arranged in radial rows with distinct secondary spiral pattern, increasing in size away
from the central rosette towards the periphery, 3.5-4.5 in 10 µm near the center, 2.5-3 in 10 µm at the margin.
Marginal rimoportulae are barely visible with LM. Two macrorimoportulae are rather small, located about
130° apart. Chloroplasts are numerous, plate-like, rounded.
Distribution:
Coscinodiscus oculus-iridis is a cosmopolitan species widely distributed worldwide (Simonsen, 1974; Guiry
& Guiry, 2017). It has been previously reported from the Arabian Gulf (Simonsen, 1974), from the
bottom sediments of Shatt Al-Arab Estuary, Iraq (Tynni, 1983; Al-Handal, 2009), and from Kuwait’s
shores (Hendey, 1970) and coastal waters (Al-Kandari et al., 2009).
Local occurrence:
Coscinodiscus oculus-iridis is the commonest species of the genus occurring across Kuwait’s waters
throughout the year, being found in 53% of all collected samples, generally fewer than 1000 cells/l. The
species was more abundant (1.1-2.2∙103 cells/l) in the inshore waters of Kuwait Bay and around Bubiyan
Island during the warm season from July to September.
COSCINODISCOPHYCEAE
Plate 21. Coscinodiscus nobilis.

a, b – live cells in valve (a) and girdle (b) view; c, e – detail of valve; d – detail of acid cleaned valve; f – detail
of valve center showing central hyaline area and scattered small processes (arrowheads).
LM, BF illumination; scale bars: a-c – 50 μm; d – 25 μm; e – 20 μm; f – 10 μm.
COSCINODISCOPHYCEAE
Coscinodiscus perforatus Ehrenberg
Plate 20: g-k
Cupp, 1943: Fig. 25A, Pl. 3: 1; Hendey, 1964, p. 77; Simonsen, 1974: p. 17; Tynni, 1983: Pl. 4: 32; Sar et al., 2010a:
Figs. 2-32.
Cells are large, solitary, and circular in valve view, 72-112 µm in diameter. Valves are almost flat, with abruptly
sloped and shallow mantle. Valve surface is areolated, with a distinct central rosette composed of enlarged
areolae, sometimes with small hyaline space inside. Areolae are large, polygonal, with cribra, arranged in
complete and incomplete alternating radial rows, with secondary spiral pattern, increasing in size away
from the central rosette towards the midradius, then smaller again, 4-5 in 10 µm. Marginal rimoportulae
are small, barely visible with LM, arranged in the valve mantle near the margin, 2-6 areolae apart. Small
rimoportulae (interstitial mesh) are scattered on the valve face at the point of origin of some incomplete
striae. Chloroplasts are numerous, plate-like, rounded to irregularly shaped.
Distribution:
Coscinodiscus perforatus is a cosmopolitan species widely distributed worldwide (Hendey, 1964; Sar et al.,
2010a; Guiry & Guiry, 2017). It has been previously reported from the Arabian Gulf (Simonsen, 1974)
and from the bottom sediments of Shatt Al-Arab Estuary, Iraq (Tynni, 1983).
Local occurrence:
Coscinodiscus perforatus was rarely recorded in Kuwait’s waters, being found in less than 2% of all collected
samples. Its distribution was restricted to the southern inshore waters off Khiran, where the species was
found in low concentrations (fewer than 100 cells/l) during the summer season from June to August and
in December.
Coscinodiscus radiatus Ehrenberg

Plate 23
Pavillard, 1925: Fig. 12; Hustedt, 1930: Fig. 225; Allen & Cupp, 1935: Fig. 8; Cupp, 1943: Fig. 20; Pl. 1: 4; Crosby &
Wood, 1958: Pl. 31: 12; Hendey, 1964: Pl. XXII: 7; Simonsen, 1974: p. 17; Tynni, 1983: Pl. 4: 33; Ricard, 1987: Figs.
1-3; Throndsen et al., 2007: p. 151; Hoppenrath et al., 2009: Fig. 12 d-g.
Cells are large to medium-sized, solitary, and circular in valve view, 55-84 µm in diameter. Valves are almost
flat, with abruptly sloped and shallow mantle. Valve surface is areolated, without a distinct central rosette or
with a cluster of somewhat enlarged areolae in large specimens. Areolae are large, polygonal, with cribra,
irregularly arranged, nearly same size on whole valve, 3-4 in 10 µm, except two rows at margin where they
are smaller, 6-7 in 10 µm. Marginal rimoportulae are small, barely visible with LM, arranged in the valve
mantle near the margin. Chloroplasts are numerous, plate-like, rounded, more densely arranged along the
girdle.
Distribution:
Coscinodiscus radiatus is a cosmopolitan species from oceanic to neritic waters from polar to tropical
regions (Cupp, 1943; Hendey, 1964; Simonsen, 1974; Guiry & Guiry, 2017). The species has been
previously reported from the Arabian Gulf (Simonsen, 1974), from the bottom sediments of the Shatt
Al-Arab Estuary, Iraq (Tynni, 1983), and from Kuwait’s waters (Al-Kandari et al., 2009).
Plate 22. Coscinodiscus oculus-iridis.

a, b – live cells in valve view; c – detail of valve center; d, e – acid cleaned valves; f – detail of cleaned valve
showing central rosette and valve margin; g – detail of central rosette in external valve view; h – detail of valve
margin showing morphology of cribra and marginal ring of rimoportulae (arrowheads) in external view.
LM, BF illumination (a-e) and DIC optic (f); SEM (g, h); scale bars: a, b, d-f – 25 μm; c, g, h – 10 μm.
COSCINODISCOPHYCEAE
Local occurrence:
Coscinodiscus radiatus frequently occurred across Kuwait’s waters throughout the year, being found in
43% of all collected samples, generally in low concentrations. The species was more abundant (400-920
cells/l) in the inshore waters of Kuwait Bay and around Bubiyan Island during the period from March
to September.
Genus Coscinodiscopsis Sar et Sunesen
Coscinodiscopsis jonesiana (Greville) Sar et Sunesen

Plates 24, 25
Allen & Cupp, 1935: Figs. 10, 11 (as Coscinodiscus jonesianus); Crosby & Wood, 1958: p. 495 (as Coscinodiscus concinnus
var. jonesianus); Hendey, 1964: p. 79 (as Coscinodiscus jonesianus); Ricard, 1987: Fig. 4 (as Coscinodiscus jonesianus); Hasle
& Syvertsen, 1997: Pl. 17 (as Coscinodiscus jonesianus); Sar et al., 2008: Figs. 1-14.
Synonymized names:
Eupodiscus jonesianus Greville 1862
Coscinodiscus jonesianus (Greville) Ostenfeld 1915
Cells are large, solitary, double convex in girdle view and circular in valve view, 120-165 µm in diameter.
Valves are convex, weakly flattened at the center, areolated, with a distinct central rosette composed of
enlarged radially elongated areolae. Areolae are roundish to hexagonal, arranged in fasciculate pattern,
forming spiral decussate arcs towards the center, with alternating complete and incomplete rows, 5-6 in
10 µm near the central area, about 7-9 in 10 µm towards the periphery. At about 2/3 of radius, there is an
irregular ring of small interstitial rimoportulae. Marginal rimoportulae are small, barely observable with
LM, arranged in ring along the valve margin, and rather irregularly distributed, 6-8 areolae apart. Two
macrorimoportulae are large, distinct, conical in shape and opening with a pore at the top, located about
120° apart. Chloroplasts are numerous, small, plate-like and roundish.
Remarks:
Close examination of Kuwait’s material with emphasis on morphology of rimoportulae on the
valve surface with electron microscopy (Plate 25) allowed to verify the identification and to avoid
misidentification with morphologically similar Coscinodiscopsis commutata (syn. Coscinodiscus commutatus).
Distribution:
Coscinodiscopsis jonesiana is a neritic species widely distributed in warm temperate to tropical regions
(Hendey, 1964; Simonsen, 1974; Guiry & Guiry, 2017). The species has been previously reported (as
Coscinodiscus jonesianus) from the Arabian Gulf (Simonsen, 1974) and Kuwait’s waters (Al-Kandari et
al., 2009).
Local occurrence:
Coscinodiscopsis jonesiana rarely occurred in Kuwait’s waters, being found in 3% of all collected samples,
and never in high concentrations. Its distribution was restricted to the northern inshore waters around
Bubiyan Island, where species was more abundant (50-100 cells/l) during the early spring in February
and March.
Plate 23. Coscinodiscus radiatus.

a, b – live cells in valve view; c, d – acid cleaned valves; e – frustule in external valve view showing external holes
of two macrorimoportulae (arrowheads); f – detail of valve margin showing morphology of cribra, external
hole of macrorimoportula (black arrowhead), marginal ring of microrimoportulae (white arrowheads), and
valve rimoportula (arrow).
LM, BF illumination (a, b) and phase contrast (c, d); SEM (e, f); scale bars: b, d – 20 μm; a, c, e – 10 μm; f – 5
μm.
Plate 24. Coscinodiscopsis jonesiana.
a – live cells in valve view showing two macrorimoportula (arrowheads); c – dividing cell in girdle view; b,
d-g – detail of valve showing central rosette, marginal area, and macrorimoportula (arrowheads).
LM, BF illumination (c) and DIC optic (a, b, d-g); scale bars: a-e – 25 μm; f, g – 10 μm.
COSCINODISCOPHYCEAE
Plate 25. Coscinodiscopsis jonesiana.

a, b – frustule in external valve (a) and oblique (b) view showing large macrorimoportulae
(arrowheads); c – detail of frustule showing macrorimoportulae (arrowheads); d –
detail of valve margin showing macrorimoportula (black arrowhead) within a ring
of marginal microrimoportulae (white arrowheads); e – detail of valve margin with
macrorimoportula (black arrowhead), marginal microrimoportula (white arrowhead),
and valve mesorimoportula (arrow); f – detail of valve surface showing morphology of
cribra and valve mesorimoportula (arrow).
SEM; scale bars: a, b – 20 μm; c-e – 10 μm; f – 5 μm.
Genus Palmerina Hasle
Palmerina hardmaniana Hasle

Plate 26
Allen & Cupp, 1935: Fig. 91 (as Hemidiscus hardmanianus); Simonsen, 1974: p. 19; Hasle & Syvertsen, 1997: Pl. 18 (as
Palmeria hardmaniana).
Synonymized names:
Palmeria hardmaniana Greville 1865
Hemidiscus hardmannianus (Greville) Kuntze 1898
Cells are large, solitary, similar in shape to a sector of a sphere, cuneate in girdle view, 420-450 µm long,
190-230 µm wide, and semilunate in valve view, with straight ventral and convex dorsal margin. Valve face is
flat, with narrow mantle along the ventral margin and apices, and very wide mantle along the convex dorsal
margin. Valve surface is finely areolated, with hyaline central area. Areolae are small, arranged in radial rows
alternating with hyaline lines associated with small marginal rimoportulae, among them two rimoportulae
are larger than the others. Nucleus is centrally located close to the ventral margin, lying in plasma mass
suspended by cytoplasmic strands extending to the cell walls. The cytoplasm contains numerous small disk-
shaped chloroplasts.
Distribution:
Palmerina hardmaniana is a warm water neritic species. It has been previously reported (as Palmeria
hardmaniana) from the Arabian Gulf (Simonsen, 1974) and from Kuwait’s waters (Al-Kandari et al.,
2009).
Local occurrence:
Palmerina hardmaniana is one of the most important species with respect of its contribution to the
phytoplankton standing crop (Polikarpov et al., 2009). This species was occasionally observed across
Kuwait’s waters throughout the year, being recorded from 19% of all collected samples. The species
was most abundant (100-440 cells/l) in the northern inshore waters in Kuwait Bay and around Bubiyan
Island during the warm season from April to June.
Family Heliopeltaceae Smith

Genus Actinoptychus Ehrenberg
The species of the genus Actinoptychus are characterized by valves divided into sectors which are alternately
raised and depressed. The species within the genus are distinguished by valve diameter, the number of
sectors, and the type and distribution of the submarginal processes (Cupp, 1943; Hendey, 1964).
Actinoptychus senarius (Ehrenberg) Ehrenberg

Plate 27: a
Crosby & Wood, 1958: Pl. 32: 19; Hendey, 1964: Pl. XXIII: 1, 2; Hendey, 1970: p. 114; Simonsen, 1974: p. 23; Priddle
& Fryxell, 1985: p. 110-111; Witkowski et al., 2000: Pl. 3: 4, 5; Throndsen et al., 2007: p. 152; Hoppenrath et al., 2009:
Fig. 14 a-m.
Synonymized names:
Actinocyclus senarius Ehrenberg 1838
Actinocyclus undulatus Kützing 1844
Actinoptychus undulatus (Kützing) Ralfs 1861
Cells are small, solitary, undulated in girdle view and circular in valve view, 35-44 µm in diameter. Valve
is divided into six sectors, which are alternately raised and depressed, with hexagonal hyaline central area.
Valve surface is strongly irregularly areolated, areolae are 7 in 10 µm. Each elevated sector possesses a single
rimoportula near the valve margin.
COSCINODISCOPHYCEAE
Plate 26. Palmerina hardmaniana.

a-d – live cells in oblique valve (a-c) and girdle (d) view; e-i – detail of acid cleaned valves showing central
hyaline area (e, g) and marginal area (f, h, i).
LM, BF illumination (a-d), DIC optic (g-i), and phase contrast (e, f); scale bars: a-d – 100 μm; e-i – 10 μm.
Distribution:
Actinoptychus senarius is a cosmopolitan species, commonly occurs in neritic plankton. It has been
previously reported from the Arabian Gulf (Simonsen, 1974) and from Kuwait’s shores (Hendey, 1970).
Local occurrence:
Actinoptychus senarius is a very rare species in Kuwait’s waters. Several frustules were isolated from
plankton net hauls in the inshore waters of Kuwait Bay.
Family Hemidiscaceae Hendey ex Hasle

Genus Actinocyclus Ehrenberg
The species of the genus Actinocyclus are distinguished by the presence of a pseudonodulus, distinct fasciculated
areolation, and more or less distinct marginal zone different from the rest of the valve. Morphological
characters used for the identification of species in the genus include the size and shape, the number of striae
in 10 μm, the position of the pseudonodulus, and the number of labiate processes (rimoportulae) present at
the valve margin (Cupp, 1943; Hasle, 1977; Hasle & Syvertsen, 1997; Throndsen et al., 2007).
Actinocyclus curvatulus Janisch

Plate 27: b-h
Pavillard, 1925: Fig. 19 (as Coscinodiscus curvatulus); Hustedt, 1930: Fig. 307; Cupp, 1943: Fig. 17, Pl. 1: 2 (as Coscinodiscus
curvatulus); Hendey, 1964: p. 81 (as Coscinodiscus curvatulus); Simonsen, 1974: p. 20; Priddle & Fryxell, 1985: p. 106-
107; Witkowski et al., 2000: Pl. 6: 2; Throndsen et al., 2007: p. 147; Hoppenrath et al., 2009: Fig. 13 j-l; Al-Yamani &
Saburova, 2011: Pl. 5: a-d.
Synonymized names:
Coscinodiscus curvatulus var. subocellata Grunow 1884
Actinocyclus subocellatus (Grunow) Rattray 1890
Cells are large, solitary, circular in valve view, 80-95 µm in diameter. Valves are almost flat, covered with
coarse hexagonal areolae, with irregularly outlined central annulus. Areolation is fasciculated, areolae are
arranged in slightly curved rows that are parallel to the side row of the sector, with rimoportulae at the end
of side rows. Areolae are decreasing in size towards the margin, 5-6 in 10 μm in the center of valve and 8-9
in 10 μm near the margin. Rimoportulae are arranged in regular marginal ring. A distinct pseudonodulus is
located near the valve margin. Numerous small discoid golden-brown chloroplasts are arranged around the
cell wall.
Distribution:
Actinocyclus curvatulus is a cosmopolitan species widely distributed from temperate to tropical waters
worldwide. The species has been previously reported from the Arabian Gulf (Simonsen, 1974) and from
Kuwait’s shores (Al-Yamani & Saburova, 2011).
Local occurrence:
Actinocyclus curvatulus is a typically benthic species (Al-Yamani & Saburova, 2011), which occasionally
becomes stirred up from the bottom sediments and carried into the water column. Frustules and live cells
of this species were sporadically found in plankton net hauls in Kuwait’s inshore waters.
COSCINODISCOPHYCEAE
Plate 27. Actinoptychus senarius and Actinocyclus curvatulus.

a – Actinoptychus senarius: acid cleaned valve, arrowheads point to rimoportulae; b-h – Actinocyclus curvatulus:
b, c – live cells in valve view; d-f – acid cleaned valves showing areolation and marginal pseudonodulus
(arrows); g, h – detail valve margin with pseudonodulus (arrow) and marginal rimoportulae (arrowheads).
LM, BF illumination (a), DIC optic (b, c), and phase contrast (d, e, g); SEM (f, h); scale bars: c – 20 μm; a,
b, d-g – 10 μm; h – 5 μm.
Actinocyclus exiguus Fryxell et Semina
Plate 29: a-d
Fryxell & Semina, 1981: Figs. 1-10; Scott & Thomas, 2005: Fig. 2.24.
Cells are small, solitary, circular in valve view, 12-16 μm in diameter. The frustules are cylindrical, with
pervalvar axis less than the diameter. Valve is heavily silicified, almost flat, with high mantle. Areolae are
hexagonal, arranged in irregular radial rows, decreasing in size towards the margin, 13-14 in 10 μm in the
center of valve and 21-23 in 10 μm near the margin. There are five to six rimoportulae per valve, arranged
in marginal ring, with slightly expanded external openings. Small pseudonodulus is located close to valve
mantle. Valvocopula is hyaline. Numerous small discoid golden-brown chloroplasts are arranged around the
cell wall.
Distribution:
Actinocyclus exiguus is a marine species reported from the southern cold water regions, including Southern
Ocean, Southern Indian Ocean (type locality), and sub-Antarctic and Antarctic waters (Fryxell &
Semina, 1981; Scott & Thomas, 2005).
Local occurrence:
Actinocyclus exiguus sporadically occurred in Kuwait’s inshore waters, and was observed in low
concentrations. The photographed specimens were isolated from the northern waters in December 2013.
The species is reported for the first time from Kuwait, however, subtropical occurrence compared to
the known distribution of Actinocyclus exiguus in the southern cold water region needs to be verified
and is awaiting additional material from Kuwait’s waters for detailed morphological and molecular
examination.
Actinocyclus octonarius Ehrenberg

Plate 28
Péragallo & Péragallo, 1897-1908: Pl. CXIV: 1, 2 (as Actinocyclus Ehrenbergii); Pavillard, 1925: Fig. 30 (as Actinocyclus
ehrenbergii); Hustedt, 1930: Figs. 298, 299, 301 (as Actinocyclus ehrenbergii); Hendey, 1964: Pl. XXIV: 3 (as Actinocyclus
octonarius var. octonarius); Hendey, 1970: p. 113; Simonsen, 1974: p. 21; Foged, 1984: Pl. XX: 1 (as Actinocyclus ehrenbergii);
Priddle & Fryxell, 1985: p. 108-109; Podzorski & Håkansson, 1987: Pl. 3: 1; Ricard, 1987: Figs. 35-39; Witkowski et
al., 2000: Pl. 4: 2, 3; Throndsen et al., 2007: p. 147; Al-Handal, 2009: Fig. 8; Al-Kandari et al., 2009: p. 53: Pl. 15: A;
Hoppenrath et al., 2009: Figs. 5, 13 a-i; Al-Yamani & Saburova, 2011: Pls. 6-8.
Synonymized names:
Actinocyclus ehrenbergii Ralfs 1861
Actinocyclus moniliformis Ralfs ex Pritchard 1861
Cells are large, solitary, and circular in valve view, 79-135 µm in diameter. Valve surface is slightly convex
and furnished with fasciculated areolation. Areolae form distinct fascicles separated by radiate longer striae
with rimoportulae at the ends, 6-7 in 10 µm. Marginal zone is broad, with areolae smaller than elsewhere on
the valve face. Pseudonodulus is large, located near the valve margin. Numerous golden-brown disk-shaped
to oval chloroplasts are arranged along cell wall.
Plate 28. Actinocyclus spp.

a-d – Actinocyclus octonarius var. ralfsii: a, b – live cells in valve view; c – acid cleaned valve; d – detail of
cleaned valve with marginal pseudonodulus (arrow); e, f – Actinocyclus octonarius var. crassus: e – live cell
in valve view; f – acid cleaned valve, arrows point to marginal pseudonodulus; g-j – Actinocyclus octonarius
var. tenellus: g, h – live cell in valve view at different focal planes; i – acid cleaned valve, arrow points to
pseudonodulus; j – detail of cleaned valve showing marginal rimoportulae (arrowheads) and pseudonodulus
(arrow); k-m – Actinocyclus subtilis: k – live cell in valve view; l – acid cleaned valve; m – detail of cleaned
valve, arrows point to marginal pseudonodulus.
LM, BF illumination (a, b, e, g, h, k) and phase contrast (c, d, e, i, j, l, m); scale bars: a-c, k, l – 20 μm; e-j,
m – 10 μm.
COSCINODISCOPHYCEAE
Remarks:
Several intraspecific varieties have been described within this taxon (e.g. Hustedt, 1930). Actinocyclus
octonarius var. crassus, var. ralfsii, and var. tenellus are distinguished from Kuwait’s material (see below).
Distribution:
Actinocyclus octonarius is a cosmopolitan species, commonly occurs in neritic plankton. The species has
been previously reported from the Arabian Gulf (Simonsen, 1974), bottom sediments in the Shatt Al-
Arab Estuary, Iraq (Al-Handal, 2009), and from Kuwait’s shores (Hendey, 1970; Al-Yamani & Saburova,
2011) and coastal waters (Al-Kandari et al., 2009).
Local occurrence:
Actinocyclus octonarius is a typically benthic species commonly observed in epiphytic assemblages on
Padina sp. and Sargassum spp. (Al-Yamani & Saburova, 2011), which occasionally becomes stirred up
from the bottom sediments and carried into the water column. This species occurred occasionally across
Kuwait’s waters, being recorded from 16% of all collected samples, more abundantly (0.4-1.4∙103 cells/l)
in Kuwait Bay all the year round.
Actinocyclus octonarius var. crassus (Smith) Hendey

Plate 28: e, f
Péragallo & Péragallo, 1897-1908: Pl. CXIV: 3, 4 (as Actinocyclus crassus); Hustedt, 1930: Fig. 301 (as Actinocyclus
Ehrenbergii var. crassa); Hendey, 1964: p. 83; Tynni, 1983: Pl. 5: 42; Hein et al., 2008: Fig. 1: 2 (as Actinocyclus ehrenbergii
var. crassa); Al-Kandari et al., 2009: p. 53, Pl. 15: B, C; Al-Yamani & Saburova, 2011: Pl. 6: a-d.
Synonymized names:
Eupodiscus crassus Smith 1853
Actinocyclus crassus (Smith) Ralfs ex Pritchard 1861
Actinocyclus ehrenbergii Ralfs 1861 var. crassus (Smith) Hustedt 1927-1930
The variety differs from the nominate species principally in size and valve areolation. Valves are 56-75 μm in
diameter. Radial structure of areolation on the valve is less evident than in the nominate species.
Distribution:
Variety has been previously recorded from the bottom sediments in the Shatt Al-Arab Estuary, Iraq
(Tynni, 1983), and from Kuwait’s intertidal sediments (Al-Yamani & Saburova, 2011) and coastal waters
(Al-Kandari et al., 2009).
Local occurrence:
Actinocyclus octonarius var. crassus is an epiphytic and neritic form associated with the nominate species
and rarely occurred in Kuwait’s waters.
Actinocyclus octonarius var. ralfsii (Smith) Hendey

Plate 28: a-d
Péragallo & Péragallo, 1897-1908: Pl. CXIII: 1, 2 (as Actinocyclus Ralfsii); Hustedt, 1930: Fig. 299 (as Actinocyclus
Ehrenbergii var. Ralfsii); Hendey, 1964: Pl. XXIV: 4; Al-Yamani & Saburova, 2011: Pl. 7: a-d.
Synonymized names:
Eupodiscus ralfsii Smith 1856
Actinocyclus ralfsii (Smith) Ralfs 1861
Coscinodiscus fuscus Norman 1861
Actinocyclus ehrenbergii var. ralfsii (Smith) Hustedt 1927-1930
The variety differs from the nominate species principally in valve areolation. Valves are 100-110 μm in
diameter. Interfascicular space is well-marked. Areolae are arranged radially, but each shorter areolae row
decreases almost parallel to the longer row. Marginal rim is distinct, narrow.
COSCINODISCOPHYCEAE
Distribution:
Variety has been previously recorded from Kuwait’s intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Actinocyclus octonarius var. ralfsii is an epiphytic and neritic form associated with the nominate species and
rarely occurred in Kuwait’s waters.
Actinocyclus octonarius var. tenellus (Brébisson) Hendey

Plate 28: g-j
Péragallo & Péragallo, 1897-1908: Pl. CXIII: 7, 8 (as Actinocyclus tenellus); Hustedt, 1930: Fig. 302 (as Actinocyclus
Ehrenbergii var. tenella); Hendey, 1964: p. 84; Hendey, 1970: p. 113; Al-Kandari et al., 2009: p. 54, Pl. 15: D, E; Al-
Yamani & Saburova, 2011: Pl. 8: a-d.
Synonymized names:
Eupodiscus tenellus Brébisson 1854
Actinocyclus tenellus (Brébisson) Grunow 1867
Actinocyclus ehrenbergii var. tenella (Brébisson) Hustedt 1927-1930
The variety differs from the nominate species principally in size and valve areolation. Valves are 34-41 μm
in diameter. Valve surface is divided into six to seven distinct sectors of loosely fasciculated striae. Each
sectorial stria has a small rimoportula where it terminates on the valve margin.
Distribution:
Variety has been previously reported from Kuwait’s Mina Al-Ahmadi shore (Hendey, 1970), intertidal
sediments (Al-Yamani & Saburova, 2011), and coastal waters (Al-Kandari et al., 2009).
Local occurrence:
Actinocyclus octonarius var. tenellus is an epiphytic and neritic form associated with the nominate species
and rarely occurred in Kuwait’s waters.
Actinocyclus subtilis (Gregory) Ralfs

Plate 28: k-m
Péragallo & Péragallo, 1897-1908: Pl. CXIV: 5; Hustedt, 1930: Fig. 304; Hendey, 1964: p. 84; Hendey, 1970: p. 113;
Simonsen, 1974: p. 21; Foged, 1984: Pl. XVII: 8; Witkowski et al., 2000: Pl. 4: 1; Al-Handal, 2009: Fig. 15; Al-Yamani
& Saburova, 2011: Pl. 9: a-e.
Synonymized names:
Eupodiscus subtilis Gregory 1857
Eupodiscus gregoryanus Brébisson 1870
Actinocyclus subtilis var. disjuncta Rattray 1890
Actinocyclus falsus Smith
Cells are large, solitary, and circular in valve view, 75-90 μm in diameter. Valves are almost flat, finely
areolated, with small, distinct, irregularly areolated central area. Areolae are small, 18-20 in 10 μm, arranged
in closely packed fascicules, with rimoportulae at the end of side rows. Pseudonodulus is large, distinct,
located near the valve margin and surrounded by hyaline ring. Numerous discoid golden-brown chloroplasts
are arranged along cell wall.
Distribution:
Actinocyclus subtilis is a cosmopolitan neritic principally warm water species widely distributed from
temperate to tropical areas worldwide (Hendey, 1964; Simonsen, 1974). The species has been previously
reported from the Arabian Gulf, most frequently in the northern part (Simonsen, 1974), from the bottom
sediments in the Shatt Al-Arab Estuary, Iraq (Al-Handal, 2009), and from Kuwait’s Mina Al-Ahmadi
shore (Hendey, 1970) and intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Actinocyclus subtilis is a typically benthic species commonly observed in epiphytic assemblages mainly on
Sargassum spp. (Al-Yamani & Saburova, 2011), which occasionally becomes stirred up from the bottom
sediments or macroalgae and carried into the water column. This species occurred occasionally across
Kuwait’s waters, being recorded from 8% of all collected samples, more abundantly (160-700 cells/l)
around Bubiyan Island during the period from February to May.
Actinocyclus sp.
Plate 29: e-g
Valves are weakly silicified, almost flat, circular, 13-19 μm in diameter. Areolae are small, arranged in
irregular rows with a tendency toward fasciculation, slightly decreasing in size close to the margin, 24-25
in 10 μm in the center of valve and 27-28 in 10 μm near the margin. The fascicles are separated by shorter
rows ending by hyaline strips with a rimoportula at the end. There are four to five marginal rimoportulae per
valve. Externally, their apertures are round rimmed pores, internally these are fan-shaped projections. Small
pseudonodulus is located slightly away from the marginal ring of rimoportulae toward the valve center, close
to one rimoportula.
Local occurrence:
Actinocyclus sp. is too small for accurate determination during routine phytoplankton examination. The
small size makes this species easy to be overlooked or misidentified with Thalassiosira species. Valves
of Actinocyclus sp. could rarely be observed in electron microscopy preparations from Kuwait’s coastal
waters. The photographed valves were isolated from Kuwait Bay in May 2017.
Genus Roperia Grunow
Roperia tesselata (Roper) Grunow

Plate 29: h, i
Hendey, 1964: Pl. XXII: 3; Hendey, 1970: p. 157; Podzorski & Håkansson, 1987: Pl. 2: 3; Ricard, 1987: Figs. 40-46;
Lee & Lee, 1990: Figs. 1-19; Al-Handal, 2009: Figs. 182, 183; Hoppenrath et al., 2009: Figs. 5, 13 n-r; Al-Yamani &
Saburova, 2011: Pl. 10: a-d.
Synonymized names:
Eupodiscus tesselatus Roper 1858
Actinocyclus tessellatus (Roper) Ralfs 1861
Actinocyclus coscinodiscoides A. Mann 1937
Roperia tesselata var. coscinodiscoides (A. Mann) Kolbe 1955
Cells are small, solitary, subcircular in valve view, 35-41 μm in diameter. Valves are almost circular to slightly
elliptical or ovate, with flat face, distinctly areolated. Areolae are rather small, hexagonal, arranged in
abruptly linear pattern over the major part of valve and tend to be radiate and smaller forwards the margin,
9-12 in 10 μm. The ring of small rimoportulae is located around the junction of the valve face and mantle,
5-7 areolae apart. A single prominent pseudonodulus is located on the valve surface near the margin. The
cytoplasm contains numerous small discoid chloroplasts.
Distribution:
Roperia tesselata is a cosmopolitan species with wide distribution from temperate to tropical waters
worldwide. The species has been previously reported from the bottom sediments of the Shatt Al-Arab
Estuary, Iraq (Al-Handal, 2009) and from Kuwait’s shores (Hendey, 1970; Al-Yamani & Saburova, 2011).
Local occurrence:
Roperia tesselata is of low occurrence in Kuwait’s waters. Live cells of this species were sporadically
observed in the inshore habitats but never in high concentrations.
Plate 29. Actinocyclus spp. and Roperia tesselata. COSCINODISCOPHYCEAE
a-d – Actinocyclus exiguus: a, b – live cell in valve view at different focal planes; c – acid cleaned frustule
in valve view showing marginal rimoportulae (arrowheads) and pseudonodulus (arrow); d – detailed valve
margin with marginal rimoportulae (arrowheads) and pseudonodulus (arrow); e-g – Actinocyclus sp.: e –
external valve view showing four marginal rimoportulae (arrowheads) and pseudonodulus (arrow); f, g –
valves in internal view with four (f) and five (g) rimoportulae, arrows point to pseudonodulus and arrowheads
point to rimoportulae; h, i – Roperia tesselata: h – live cell in valve view; i – acid cleaned valve, arrow points
to pseudonodulus and arrowhead to marginal rimoportula.
LM, DIC optic (a, b, h); SEM (c-g, i); scale bars: h, i – 10 μm; a-c, e-g – 5 μm; d – 2 μm.
Genus Hemidiscus Wallich
Hemidiscus cuneiformis Wallich

Plate 30: a
Pavillard, 1925: Fig. 10 (as Euodia cuneiformis); Hustedt, 1930: Fig. 542; Cupp, 1943: Fig. 121; Crosby & Wood, 1958:
Pl. 34: 53; Hendey, 1964: Pl. XXII: 9; Simonsen, 1974: p. 21; Ricard, 1987: Figs. 30-34; Throndsen et al., 2007: p. 148.
Synonymized names:
Euodia gibba (Bailey) Ralfs 1861
Euodia cuneiformis (Wallich) Schütt 1896
Euodia inornata Castracane 1886
Euodia radiata Castracane 1886
Cells are large, solitary, cuneiform in girdle view and semicircular in valve view, 110 µm long (apical axis)
and 65 µm wide (transapical axis). Valves are almost flat, with almost straight ventral margin with a weak
median inflation, and strongly convex dorsal margin. Valve surface is irregularly fasciculated. Areolae are
small, 7-9 in 10 µm in the center and 12-15 in 10 µm at the valve margin. Rimoportulae are arranged in line
along the valve margin and evenly distributed. Pseudonodule is located at center of side having the least
curvature. Chloroplasts are small, numerous, disk-shaped.
Distribution:
Hemidiscus cuneiformis is an oceanic warm water species with a wide distribution predominantly in
tropical and subtropical waters. The species has been previously reported from Indian Ocean (Simonsen,
1974) and from Kuwait’s waters (Al-Yamani et al., 2004; Al-Kandari et al., 2009).
Local occurrence:
Hemidiscus cuneiformis is a very rare species in Kuwait’s waters. Live cells and frustules of this species
were found in a plankton hauls from the offshore southern waters, but it has never been recorded during
routine phytoplankton monitoring.
Order Stellarimales Nikolaev et Harwood

Family Stellarimaceae Nikolaev ex Sims et Hasle
Genus Stellarima Hasle et Sims
The species of the genus Stellarima are characterized by presence of several labiate processes in valve center
and non along the margin (Throndsen et al., 2007).
Stellarima stellaris (Roper) Hasle et Sims

Plate 30: b
Pavillard, 1925: Fig. 20 (as Coscinodiscus stellaris); Hustedt, 1930: Fig. 207 (as Coscinodiscus stellaris); Cupp, 1943: Fig. 16
(as Coscinodiscus stellaris); Hendey, 1964: p. 81 (as Coscinodiscus stellaris); Simonsen, 1974: p. 18 (as Coscinodiscus stellaris);
Throndsen et al., 2007: p. 152; Hoppenrath et al., 2009: Fig. 12 m, n.
Synonymized names:
Coscinodiscus stellaris Roper 1858
Cells are large, solitary, thin-walled, and circular in valve view, 84-95 µm in diameter. Valves are strongly
convex, areolated, with three to six irregular dark thickenings making a star-like formation at the center.
Areolae are arranged in more or less broad radial sectors within which the middle rows are nearly parallel,
and tangential secondary rows are concave toward the outside, decreasing in size towards the margin, about
12-13 in 10 µm at the center, 15-16 midway, and 17-20 near the margin.
Distribution:
Stellarima stellaris is an oceanic cosmopolitan species widely distributed worldwide (Hendey, 1964). It has
been previously reported from the Arabian Gulf (Simonsen, 1974).
COSCINODISCOPHYCEAE
Plate 30. Hemidiscus cuneiformis, Stellarima stellaris, Podosira stelligera, and Paralia sulcata.
a – Hemidiscus cuneiformis: acid cleaned frustule in valve view; b – Stellarima stellaris: acid cleaned valve; c-e
– Podosira stelligera: acid cleaned frustules in oblique (d) and valve (c, e) view at different focal planes; f-j –
Paralia sulcata: f-h – live cells united to chains in girdle view at different focal planes; i, j – acid cleaned valves.
LM, BF illumination (f-h, j) and phase contrast (a-e, i); SEM (j); scale bars: a-e – 20 μm; f-j – 10 μm.
Local occurrence:
Stellarima stellaris is a very rare species in Kuwait’s waters. Frustules of this species were found in plankton
hauls from the southern offshore waters and were isolated from the bottom sediments. The species is
recorded for the first time in Kuwait.
Subclass Melosirophycidae Cox

Order Melosirales Crawford
Family Hyalodiscaceae Crawford
Genus Podosira Ehrenberg
The species of the genus Podosira are characterized by convex and strongly silicified valves with scattered
rimoportulae and girdle composed of numerous intercalary bands. Cells usually fixed by means of a mucous
pad to the substratum, but frequently found in plankton (Hendey, 1964).
Podosira stelligera (Bailey) Mann

Plate 30: c-e
Hustedt, 1930: Fig. 128; Crosby & Wood, 1958: Pls. 31: 5, 35: 1 (as Hyalodiscus stelliger); Hendey, 1964: Pl. XXII: 6;
Simonsen, 1974: p. 8; Tynni, 1983: Pl. 1: 5; Ricard, 1987: Figs. 191, 192; Hoppenrath et al., 2009: Fig. 16 a-h.
Synonymized names:
Hyalodiscus stelliger Bailey 1854
Podosira maculata Smith 1856
Melosira maculata Lagerstedt 1876
Cells are solitary or in pairs connected by their girdles, strongly silicified, circular in valve view, 55-68 µm in
diameter, 15-30 µm broad (pervalvar axis). Valves are strongly convex, with well differentiated, irregularly
punctated bright central area and with dark finely striated peripheral zone. Striation is fasciculated, giving
the appearance of dividing the valve into sectors. Marginal zone is strong, finely striated. Girdle is composed
of numerous narrow intercalary bands. Chloroplasts are small, numerous, plate-like.
Distribution:
Podosira stelligera is a cosmopolitan tychopelagic species commonly recorded in the phytoplankton
composition worldwide. The species has been previously reported from the Arabian Gulf (Simonsen,
1974) and from the bottom sediments in the Shatt Al-Arab Estuary, Iraq (Tynni, 1983).
Local occurrence:
Podosira stelligera is reported for the first time from Kuwait’s waters. This is a typically benthic species. Its
cells live either singly or in short chains, attached to the substratum by a small pad of mucilage. Podosira
stelligera occasionally become stirred up from the bottom sediments and carried into the water column.
Small numbers of frustules of this species were found in plankton hauls from Kuwait’s inshore waters
and were isolated from the bottom sediments.
Family Paraliaceae Crawford

Genus Paralia Heiberg
The species of the genus Paralia are characterized by short cylindrical heterovalvate cells united to form
chains, with two types of valves, including separation valves at the ends of chains without linking spines, and
linking intercalary valves possessing linking spines. Valve mantle is strongly loculate and ornamented with a
coarse network of sub-hexagonal cellulation (Hendey, 1964; Hoppenrath et al., 2009).
COSCINODISCOPHYCEAE
Paralia sulcata (Ehrenberg) Cleve
Plate 30: f-j
Pavillard, 1925: Fig. 1; Hustedt, 1930: Fig. 118; Allen & Cupp, 1935: Fig. 1; Cupp, 1943: Fig. 2 (as Melosira sulcata);
Crosby & Wood, 1958: Pl. 31: 2 (as Melosira sulcata); Hendey, 1964: Pl. XXIII: 5; Hendey, 1970: p. 109; Simonsen,
1974: p. 8; Tynni, 1983: Pl. 1: 4; Ricard, 1987: Figs. 174-182; Witkowski et al., 2000: Pl. 8: 10, 11; Throndsen et al.,
2007: p. 145; Al-Handal, 2009: Fig. 22; Hoppenrath et al., 2009: Fig. 16 n-r.
Synonymized names:
Gaillonella sulcata Ehrenberg 1838
Melosira sulcata (Ehrenberg) Kützing 1844
Orthoseira marina Smith 1856
Melosira marina (Smith) Janisch 1862
Paralia marina (Smith) Heiberg 1863
Paralia sulcata var. genuina Grunow 1884
Cells are low cylindrical in girdle view, wider than high, and circular in valve view, closely united in more
or less short, straight, dense chains, dark-brownish in color, 15-22 µm in diameter, 8-12 µm broad (pervalvar
axis). Cell wall is coarsely silicified. Valve face is almost flat, with wide hyaline central area surrounded by
a ring of short coarse radiating ribs, and coarse granular marginal markings. Sibling valves are connected
by interlocking ridges and marginal linking spines like a zip. Terminal valves in chains lack spines and
elevations. The cytoplasm contains several small disc-shaped chloroplasts.
Distribution:
Paralia sulcata is commonly recorded in coastal plankton worldwide. The species has been previously
reported from the northern part of the Arabian Gulf (Simonsen, 1974), bottom sediments in the Shatt
Al-Arab Estuary, Iraq (Tynni, 1983; Al-Handal, 2009), and from Kuwait’s shores (Hendey, 1970) and
coastal waters (Al-Yamani et al., 2004; Al-Kandari et al., 2009).
Local occurrence:
Paralia sulcata occasionally occurred across Kuwait’s waters all the year round, being found in 24% of all
collected samples. The species was more abundant (300-520 cells/l) in Kuwait Bay and adjacent waters
during the period from March to October.
Subphylum Bacillariophytina Medlin et Kaczmarska
Class Mediophyceae (Jousé et Proshkina-Lavrenko) Medlin et Kaczmarska
Subclass Biddulphiophycidae Round et Crawford
Order Biddulphiales Krieger
Family Attheyaceae Crawford et Round
Genus Attheya West
Attheya decora West

Plate 32: f
Ricard, 1987: Fig. 340; Al-Yamani & Saburova, 2011: Pl. 18: f.
Cells are small, delicate, weakly silicified, rectangular in girdle view, 35-38 μm long, 28-30 μm wide. Each
angle of frustule is furnished with long thin process. Two large irregularly shaped chloroplasts are lying
along the girdle, one upon either side of central nucleus.
Distribution:
Attheya decora is predominantly neritic form scarcely reported worldwide. The species has been previously
reported from the Iraqi waters (Maulood et al., 2013) and from Kuwait’s intertidal sediments most
probably due to sedimentation from plankton (Al-Yamani & Saburova, 2011).
Local occurrence:
Attheya decora sporadically occurred in Kuwait’s waters, and was never observed in large numbers.
Family Bellerocheaceae Crawford

Genus Bellerochea Van Heurck
The species of the genus Bellerochea are characterized by very weakly silicified cells united to form characteristic
ribbon-shaped chains so that the valves are more or less in contact over the whole surface, except for small
foramina near the margins. Cells are biangular to quadrangular in valve view with concave undulated sides
and roughly rectangular in girdle view, with slightly produced apices and bilabiate process with long external
tube. The species within the genus are distinguished by the shape of colonies, shape of the intercellular
spaces, and the location of bilabiate process (Hendey, 1964; Hasle & Syvertsen, 1997).
Bellerochea horologicalis von Stosch

Plate 31: a-d
Ricard, 1987: Figs. 545, 546; Hasle & Syvertsen, 1997: Pl. 48; Yahia-Kéfi et al., 2005: Figs. 2-17.
Cells are united to form ribbon-shaped curved chains. Cells are biangular, lanceolate in valve view and
roughly rectangular in girdle view. Length of pervalvar axis is 33-38 µm, apical axis is 28-47 µm, transapical
axis is 20-22 µm. An intercellular space between the intercalary cells is dumbbell-shaped, terminal cells in
chain possess a convex external valve. Chloroplasts are small, numerous, elongated.
Distribution:
Bellerochea horologicalis is a warm water species with pantropical distribution (Yahia-Kéfi et al., 2005).
The species has been previously reported from Kuwait’s waters (Al-Yamani et al., 2004; Al-Kandari et
al., 2009).
Local occurrence:
Bellerochea horologicalis occasionally occurred across Kuwait’s waters all the year round, being recorded
from 14% of all collected samples. The species was most abundant (0.9-2.4∙104 cells/l) in Kuwait Bay
and the adjacent waters in September.
Bellerochea malleus (Brightwell) Van Heurck
Plate 31: e-g
Hustedt, 1930: Fig. 456d; Hendey, 1964: Pl. VI: 5; Hendey, 1970, p. 117; Hasle & Syvertsen, 1997: Pl. 48; Hoppenrath
et al., 2009: Fig. 38 t-z.
Synonymized names:
Triceratium malleus Brightwell 1858
Cells occur in straight, long and inseparable ribbon-shaped chains. Cells are very weakly silicified, elliptical
in valve view and roughly rectangular in girdle view, pervalvar axis is 44-52 µm, apical axis is 38-41 µm.
Valves are biangulate, with low elevations at both apices. Sibling cells are tightly joined, connected by apical
elevations. An intercellular space between the intercalary cells is drop-shaped and located close to elevations.
Chloroplasts are small, numerous, round to elongate.
Distribution:
Bellerochea malleus is a commonly reported species from temperate to tropical waters. Hendey (1970)
noted that the triangular form is more frequent in temperate areas, whereas the biangular form appears in
warm waters and common in the Indian Ocean. The species has been previously reported from Kuwait’s
shores (Hendey, 1970) and coastal waters (Al-Kandari et al., 2009).
MEDIOPHYCEAE
Local occurrence:
Bellerochea malleus is a rare species in Kuwait’s waters, being recorded from less than 2% of all collected
samples, and never in large number. The species was more abundant (160-320 cells/l) in the inshore
waters off Salmiya during the warm season from August to October.
Bellerochea cf. yucatanensis von Stosch

Plate 31: h-k
Ricard, 1987: Figs. 547, 548; Hasle & Syvertsen, 1997: p. 229.
Cells are solitary, weakly silicified, elliptical in valve view and roughly rectangular in girdle view, pervalvar
axis is 47-50 µm, apical axis is 35-41 µm. Valves are biangulate, with short rounded elevations at both apices
and an inflated central area with two labiate processes. Chloroplasts are small, numerous, elongate, radiating
from the centrally located nucleus.
Remarks:
Rare occurrence of solitary cells in Kuwait’s material requires further careful examination of this taxon
to confirm the species identification. Additionally, the gross morphology of the observed cells is quite
similar to recently described from temperate waters weakly silicified chain-forming diatom Mediopyxis
helysia Kühn, Hargreaves et Halliger 2006 (Kühn et al., 2006: Figs. 1-43). However, the observed cells
from Kuwait’s subtropical waters were much smaller. The illustrated cells from Kuwait’s material were
provisionally attributed to Bellerochea yucatanensis due to their smaller size (35-41 µm in apical axis
compared to 85-125 µm in Mediopyxis helysia) and subtropical distribution compared to cold temperate
occurrence of Mediopyxis helysia described from the North Sea and the Gulf of Maine (Kühn et al.,
2006). Additional specimens from Kuwait’s waters are needed for detailed morphological examination.
Distribution:
Bellerochea yucatanensis is a warm water species known from its type locality (Porto Progreso, Yucatan)
and Australia (Hasle & Syvertsen, 1997).
Local occurrence:
Small number of cells of Bellerochea cf. yucatanensis was found once from Kuwait’s offshore waters in the
spring of 2009.
Family Biddulphiaceae Kützing
Genus Biddulphia Gray
Biddulphia pulchella Gray

Plate 32: a
Péragallo & Péragallo, 1897-1908: Pl. XCIII: 1, 2; Hustedt, 1930: Fig. 490; Cupp, 1943: Fig. 109; Crosby & Wood,
1958: Pl. 33: 32; Hendey, 1964: Pl. XXV: 1; Simonsen, 1974: p. 34; Foged, 1984: Pls. XXI: 1, 4, XXII: 1, 4; Ricard,
1987: Figs. 254-264; Witkowski et al., 2000: Pl. 8: 8, 9; Hein et al., 2008: Fig. 2: 3, 6; Al-Yamani & Saburova, 2011: Pl.
17: g.
Synonymized names:
Conferva biddulphiana Smith 1807
Diatoma biddulphiana Agardh 1824
Biddulphia australis Montagne 1842
Denticella biddulphiana Ehrenberg 1843
Biddulphia trilocularis Küzing 1844
Biddulphia quinquelocularis Küzing 1844
Biddulphia septemlocularis Küzing 1844
Biddulphia transversa Wigand 1860
Biddulphia unifasciata Wigand 1860
Biddulphia biddulphiana (Smith) Boyer 1900
Cells are solitary or united in straight short chains, cylindrical in girdle view and elliptical in valve view,
20-45 µm along apical axis. Valves are convex, biangulate, with short rounded elevations at both apices,
somewhat constricted at the base, and with two centrally located short spine-like processes. Valve surface is
finely areolated, 6-8 areolae in 10 µm, smaller near the valve center. Girdle is more finely areolated with 10-
12 areolae in 10 µm arranged in nearly parallel lines. Chloroplasts are small, numerous, plate-like.
Distribution:
Biddulphia pulchella is a widely distributed marine epiphytic and neritic species. The species has been
previously reported from the Indian Ocean (Simonsen, 1974), and from Kuwait’s waters (Al-Yamani et
al., 2004; Al-Kandari et al., 2009) and intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Biddulphia pulchella is a typically benthic species (Al-Yamani & Saburova, 2011), which sporadically
occurred in Kuwait’s inshore waters, and was never observed in large numbers.
Biddulphia tuomeyi (Bailey) Roper

Plate 32: b-e
Péragallo & Péragallo, 1897-1908: Pl. XCIII: 3, 4; Pl. XCIV: 2; Hustedt, 1930: Fig. 491; Hendey, 1970: p. 117; Simonsen,
1974: p. 34; Foged, 1984: Pls. XX: 6, XXI: 3; Ricard, 1987: Fig. 269; Al-Yamani & Saburova, 2011: Pl. 18: a-e.
Synonymized names:
Denticella tridens Ehrenberg 1838
Biddulphia tridentata Ehrenberg 1844
Denticella polymera Ehrenberg 1844
Zygoceros tuomeyi Bailey 1844
Denticella margaritifera Shatbolt 1854
Denticella siplex Shadbolt 1854
Biddulphia tridens Boyer 1927
Cells are solitary or united by their valve horns to form short straight chains, rectangular in girdle view and
elliptical in valve view, 51-65 µm along apical axis and 38-46 µm along pervalvar axis. Valves are narrowly
elongated, with three convexities, middle one is largest. Top part of each convexity is furnished with short
spikes. Valve apices are furnished with large prominent horn-like processes with capitate ends. Valve surface
is irregularly areolated, about 4-8 areolae in 10 μm. Girdle surface is finely punctated, puncta are arranged in
slightly undulated longitudinal rows, 10-12 rows in 10 μm. Chloroplasts are numerous, elongated, radiating
from the centrally located nucleus.
Distribution:
Biddulphia tuomeyi is a principally benthic warm water species. It has been previously reported from the
Arabian Gulf (Simonsen, 1974), and from Kuwait’s shores (Hendey, 1970) and intertidal sediments (Al-
Yamani & Saburova, 2011).
Local occurrence:
Biddulphia tuomeyi is a typically benthic species (Al-Yamani & Saburova, 2011), which sporadically
occurred in Kuwait’s inshore waters but never in large numbers.
Genus Eucampia Ehrenberg
The species belonging to the genus Eucampia are distinguished by bipolar frustules, elliptical valves with
elevations bearing ocellus located on each polar, and usually curved chains with large apertures between the
cells (Hustedt, 1930; Cupp, 1943; Hasle & Syvertsen, 1997).
MEDIOPHYCEAE
Eucampia cornuta (Cleve) Grunow
Plate 32: g-l
Hustedt, 1930: Fig. 452; Allen & Cupp, 1935: Fig. 75; Cupp, 1943: Fig. 104; Crosby & Wood, 1958: Pl. 36: 29; Simonsen,
1974: p. 34; Ricard, 1987: Figs. 344-347; Hasle & Syvertsen, 1997: Pl. 34.
Synonymized names:
Moelleria cornuta Cleve 1873
Cells are united by their valve horns to form curved chains, slightly curved in girdle view and concave in the
valve face, 18-24 μm along apical axis and 22-45 μm along pervalvar axis. Valves are elliptical, with narrow
and long cylindrical elevation at the poles. A single labiate process is located on the depressed central part of
valve. Valve surface is finely areolated, 22-24 areolae in 10 μm. Apertures between the cells in chain are large
and elliptical. Girdle is composed of numerous narrow intercalary bands. Chloroplasts are small, numerous,
elongated, radiating from the centrally located nucleus.
Distribution:
Eucampia cornuta is a cosmopolitan neritic warm water species widely distributed in subtropical and tropical
areas worldwide. The species has been previously reported from the Arabian Gulf (Simonsen, 1974).
Local occurrence:
Eucampia cornuta occurred occasionally across Kuwait’s waters all the year round, being recorded from
12% of all collected samples, more abundantly (0.1-4.0∙104 cells/l) during the warm season from July to
September. The species is reported for the first time in Kuwait.
Plate 31. Bellerochea spp.
a-d – Bellerochea hologicalis: detail of chains at different focal planes showing connection of sibling cells; e-g
– Bellerochea malleus: detail of chains showing connection of sibling cells; h-k – Bellerochea cf. yucatanensis:
single cells in valve (h) and girdle (i-k) view.
LM, BF illumination; scale bars: a-d, f, i-k – 10 μm; e, g, h – 20 μm.
Plate 32. Biddulphia spp., Atteya decora, and Eucampia cornuta.

a – Biddulphia pulchella: acid cleaned frustule in girdle view; b-e – Biddulphia tuomey: b – live cells united to
chain; c – acid cleaned frustule in girdle view; d – acid cleaned valve; e – cleaned frustules united to chain;
f – Atteya decora: live cell in girdle view; g-l – Eucampia cornuta: g-i – cells united to arc-shaped chains; j, k –
single live cells in girdle view; l – acid cleaned valve.
LM, BF illumination (a, b, f, j, k), DF illumination (h), and phase contrast (c-e, g, i, l); scale bars: a-i – 20 μm;
j-l – 10 μm.
MEDIOPHYCEAE
Eucampia zodiacus Ehrenberg
Plate 33
1958: Pl. 36: 28; Hendey, 1964: Pl. VII: 1; Simonsen, 1974: p. 34; Ricard, 1987: Figs. 342, 343; Hasle & Syvertsen,
1997: Pl. 35; Throndsen et al., 2007: p. 178; Hoppenrath et al., 2009: Fig. 42 j-q.
Synonymized names:
Eucampia britannica Smith 1853
Eucampia virginica Grunow ex Van Heurck 1882
Eucampia nodosa Schmidt 1888
Cells are united by their valve horns to form spirally curved chains due to unequal length of both side of cell,
trapezoid in girdle view and concave in the valve face, 40-65 μm along apical axis. Valves are elliptical, with
broad, flattened and ocellate elevation at the poles. Valve surface is finely areolated. Areolae are arranged
in radial rows running outward from center towards processes, 16-20 in 10 µm. Apertures between the cells
in chain are narrow lanceolate to elliptical. Girdle is composed of numerous narrow intercalary bands.
Chloroplasts are small, numerous, elongated, radiating from the centrally located nucleus.
Distribution:
Eucampia zodiacus is a cosmopolitan neritic species widely distributed from temperate to tropical waters.
The species has been previously reported from the Arabian Gulf (Simonsen, 1974) and from Kuwait’s
waters (Al-Yamani et al., 2004; Al-Kandari et al., 2009).
Local occurrence:
Eucampia zodiacus is one of the most important species with respect to its contribution to the phytoplankton
standing crop (Polikarpov et al., 2009). This species was frequently observed across Kuwait’s waters all
year round, being recorded from 39% of all collected samples. The highest abundances of this species
(2.3-6.5∙105 cells/l) were restricted to Kuwait Bay and adjacent waters during the warm season in May
and June. Due to its capacity to form blooms, Eucampia zodiacus has been included in the list of potentially
harmful species for Kuwait’s marine environment (Al-Yamani et al., 2012).
Order Briggerales Nikolaev et Harwood

Family Streptothecaceae Crawford
Genus Helicotheca Ricard
Helicotheca tamesis (Shrubsole) Ricard

Plate 34: a-f
Pavillard, 1925: Fig. 90 (as Streptotheca tamesis); Hustedt, 1930: Fig. 455 (as Streptotheca tamesis); Cupp, 1943: Fig. 106 (as
Streptotheca tamesis); Crosby & Wood, 1958: Pl. 36: 32 (as Streptotheca tamesis); Hendey, 1964: Pl. VII: 2 (as Streptotheca
tamesis); Ricard, 1987: Figs. 549, 550; Hasle & Syvertsen, 1997: Pl. 48 (as Streptotheca tamesis); Throndsen et al., 2007:
p. 182; Hoppenrath et al., 2009: Fig. 37 d-g; Hernández-Becerril et al., 2013: Figs. 14-27.
Synonymized names:
Streptotheca tamesis (thamensis) Shrubsole 1891
Cells are either solitary or more often united to form long twisted ribbon-shaped chains with no apertures
between sibling cells, with torsion in relation to pervalvar axis. Cells are weakly silicified, square to
rectangular in girdle view, transapically flattened, 140-160 µm along apical axis and 120-140 µm along
pervalvar axis. Valves are linear oblong with structure not resolvable with LM. The valvar ornamentation
is manifested in SEM as small scattered pores with rimmed edges and tiny randomly distributed silicious
granules. A single process with short outer tube is located close to valve margin. Numerous intercalary bands
are finely perforated but without silicious granules. Nucleus is lying in a central plasma mass, suspended by
cytoplasmic strands extending to the cell walls. Chloroplasts are small, numerous, elongated.
MEDIOPHYCEAE
Plate 33. Eucampia zodiacus.

a, b – live cells united to helically coiled chains; c-e – detail of chains; f – detail of valve view showing ocellus.
LM, BF illumination (a, b) and DIC optic (c); SEM (d, f); scale bars: a, b – 100 μm; c, d – 25 μm; e, f – 10 μm.
Distribution:
Helicotheca tamesis is a neritic predominantly warm water species. The species has been previously
reported from Kuwait’s waters (Al-Yamani et al., 2004; Al-Kandari et al., 2009).
Local occurrence:
Helicotheca tamesis was occasionally observed across Kuwait’s waters all year round, being recorded from
25% of all collected samples, more abundantly (550-840 cells/l) in Kuwait Bay and adjacent waters
during the fall season from September to November.
Genus Streptotheca Shrubsole
Streptotheca indica Karsten

Plate 34: g-i
Karsten, 1907: Tf. XLVI/XII: 8; Allen & Cupp, 1935: Fig. 77; Crosby & Wood, 1958: Pl. 36: 33.
Cells are either solitary or more often united to form short chains with no apertures between sibling cells,
with torsion in relation to pervalvar axis. Cells are weakly silicified, square to rectangular in girdle view,
tripartite in valve view, 100-110 µm along apical axis and 70-85 µm along pervalvar axis. Valves are linear
oblong with structure not resolvable with LM. Nucleus is lying in a central plasma mass, suspended by
cytoplasmic strands extending to the cell walls. Chloroplasts are small, numerous, elongated.
Distribution:
Streptotheca indica is a scarcely reported species known from the tropical waters of the Indian Ocean
(Allen & Cupp, 1935), from Australia and New Zealand (Crosby & Wood, 1958), and Asia (Taiwan,
China) (Guiry & Guiry, 2017).
Local occurrence:
Streptotheca indica is reported for the first time from Kuwait’s waters. The species sporadically occurred
in the southern inshore waters, and was observed in low concentrations.
MEDIOPHYCEAE
Plate 34. Helicotheca tamesis and Streptotheca indica.

a-f – Helicotheca tamesis: a – live cells united to twisted ribbon-like chain; b, c – detail of chain; d – single
cell in broad girdle view; e – detailed marginal area at the cell center showing marginal process; f – detailed
marginal process; g-i – Streptotheca indica: single live cells in girdle view.
LM, BF illumination (a-d, g-i); SEM (e, f); scale bars: a, b – 100 μm; c – 50 μm; d – 25 μm; g-i – 20 μm; e,
f – 2 μm.
Subclass Chaetocerotophycidae Round et Crawford
Order Chaetocerotales Round et Crawford
Family Chaetocerotaceae Ralfs
Genus Bacteriastrum Shadbolt
The species belonging to the genus Bacteriastrum are distinguished by cylindrical cells with delicate wall and
without clearly visible with LM structure bound into loose chains by the fusion of more or less numerous
setae. Setae of two adjacent cells are fused for a certain distance beyond the base, farther out divided again.
Terminal setae are morphologically distinct from the fused bristles, often curved, not fused and not branched.
The genus Bacteriastrum differs from morphologically similar Chaetoceros in radial symmetry and fenestration
of setae (Cupp, 1943; Hasle & Syvertsen, 1997). The length of the fused part of sibling setae between
intercalary cells is an important species identification criterion in Bacteriastrum (Ikari, 1927; Cupp, 1943;
Hasle & Syvertsen, 1997).
Bacteriastrum comosum var. hispidum (Castracane) Ikari

Plate 35
Allen & Cupp, 1935: Fig. 49 (as Bacteriastrum varians var. hispida).
Synonymized names:
Bacteriastrum wallichii var. hispida Castracane 1886
Bacteriastrum varians var. hispidum (Castracane) Schröder 1906
Bacteriastrum elegans var. hispida (Castracane) Pavillard 1924
Bacteriastrum furcatum var. hispida (Castracane) Simonsen 1974
The cells are cylindrical, slightly longer than broad, circular in cross section, 12-16 μm in diameter, united
into chains by the fusion of regularly arranged 10-12 radial setae. Chains are short, straight, and heteropolar,
with small apertures. Inner setae possess a short basal part with transverse bifurcation plane. Forked part is
long and already at the point of division curving toward the posterior end of chain. Anterior valve is with
the setae curved umbrella-like toward the inner part of the chain. Posterior setae are stronger than the rest,
bell-shaped, with spirally arranged little spines. Chloroplasts are small, numerous, disk-shaped.
Distribution:
Bacteriastrum comosum var. hispidum is an oceanic warm water species reported occasionally from the
subtropical and tropical areas including Java Sea (Allen & Cupp, 1935), Japanese and Chinese waters
(Guiry & Guiry, 2017). The species has been previously reported from Kuwait’s waters (Al-Kandari et
al., 2009, as Bacteriastrum varians var. hispida).
Local occurrence:
Bacteriastrum comosum var. hispidum is of low occurrence in Kuwait’s waters, being recorded from less
than 4% of all collected samples. The highest abundances of this species (0.6-1.4∙104 cells/l) were
restricted to the southern localities.
Bacteriastrum delicatulum Cleve

Plate 36
1958: Pl. 38: 61; Hendey, 1964: Pl. VI: 2; Ricard, 1987: Figs. 535, 536.
The cells are cylindrical, much longer than broad, circular in cross section, 10-15 μm in diameter, united into
chains by the fusion of regularly arranged 8-12 radial setae. Chains are long, straight, with relatively large
apertures. Inner setae possess a long basal part with bifurcated ends lie in a plane parallel to the chain axis.
Forked parts are slightly curved, smooth or somewhat wavy. Terminal setae are slightly spirally undulate,
curved towards the chain proper, forming an umbrella-like structure. Chloroplasts are small, numerous,
plate-shaped, and distributed along cell wall.
Distribution:
Bacteriastrum delicatulum is a cosmopolitan oceanic species. It has been previously reported from Kuwait’s
Local occurrence:
Bacteriastrum delicatulum occurred frequently across Kuwait’s waters, being recorded from 34% of all
collected samples. The species was present throughout the year in low to moderate abundance with
higher concentrations (0.3-1.2∙105 cells/l) recorded in the inshore waters from July to January.
MEDIOPHYCEAE
Plate 35. Bacteriastrum comosum.

a – chain in girdle view; b – anterior part of chain; c, d – posterior part of chain; e – detail of chain; f – valve
view showing posterior terminal and inner setae; g – valve view showing anterior terminal setae.
LM, BF illumination; scale bars: a – 50 μm; b-d, f, g – 25 μm; e – 20 μm.
Plate 36. Bacteriastrum delicatulum.
a, c-e – part of chains in girdle view; b, f – valve views showing inner and terminal setae; g – acid cleaned
valve with inner setae; h – acid cleaned valve with terminal setae.
LM, BF illumination (a-f) and phase contrast (g, h); scale bars: a, b, f-h – 25 μm; c, d – 20 μm; e – 10 μm.
Bacteriastrum furcatum Shadbolt
Plate 37
Allen & Cupp, 1935: Fig. 48 (as Bacteriastrum varians); Crosby & Wood, 1958: Pl. 38: 60 (as Bacteriastrum varians);
Hendey, 1970: p. 121 (as Bacteriastrum varians); Ricard, 1987: Figs. 538, 539 (as Bacteriastrum varians); Hasle & Syvertsen,
1997: Pl. 37; Sarno et al., 1997: Figs. 19-34; Bosak et al., 2015: Figs. 1-17.
Synonymized names:
Bacteriastrum curvatum Shadbolt 1854
Bacteriastrum nodulosum Shadbolt 1854
Bacteriastrum varians Lauder 1864
The cells are cylindrical, slightly longer than broad, circular in cross section, 9-14 μm in diameter, united
into chains by the fusion of regularly arranged 6-10 radial setae. Chains are short, straight, and heteropolar.
The apertures between cells are distinct due to a relatively long basal part of the setae. Forked parts of the
inner setae are lying in the valvar plane. The anterior setae are oriented almost parallel with the valve with
the proximal part bent slightly towards the inside of the chain, forming a wide umbrella-like shape. The
posterior setae form a wide simple umbrella-like shape, with the base slightly arched towards the inside of
the chain and then curved away from it. Chloroplasts are small, numerous, disk-shaped. Biconvex smooth
resting spores develop endogenously in the middle of the mother cells.
Distribution:
Bacteriastrum furcatum is a widely distributed species, reported from warm temperate to tropical waters
MEDIOPHYCEAE
(Guiry & Guiry, 2017). The species has been previously reported from Kuwait’s shores (Hendey, 1970).
Local occurrence:
Bacteriastrum furcatum occurred occasionally across Kuwait’s waters all year round, being recorded from
about 11% of all collected samples, more abundantly (0.2-5.8∙104 cells/l) during the warm season from
May to September.
Bacteriastrum hyalinum Lauder

Plate 38
Pavillard, 1925: Fig. 58; Hustedt, 1930: Fig. 354; Allen & Cupp, 1935: Fig. 47; Cupp, 1943: Fig. 56; Hendey, 1964: Pl.
VI: 1; Ricard, 1987: Fig. 537; Throndsen et al., 2007: p. 163; Hoppenrath et al., 2009: Fig. 30 i-q.
Synonymized names:
Bacteriastrum varians f. hyalina (Lauder) Frenguelli 1928
Chaetoceros spirillum (Castracane) De Toni 1894
Actiniscus varians (Lauder) Grunow 1882
Bacteriastrum spirillum Castracane degli Antelminelli 1886
Bacteriastrum varians var. princeps Castracane 1886
Bacteriastrum varians var. borealis Ostenfeld 1901
Bacteriastrum solitarium Mangin 1912
Bacteriastrum hyalinum var. princeps (Castrachane) Ikari 1927
The cells are cylindrical, often shorter than broad, circular in cross section, 22-26 μm in diameter, united into
chains by the fusion of regularly arranged numerous (up to 30) radial setae. Chains are long, straight, and
isopolar. The apertures between sibling cells are narrow due to a short basal part of the setae. Forked parts
of the inner setae are lying in the pervalvar plane giving cells a hairy appearance. The terminal setae are
shorter than the inner ones, umbrella-shaped, gradually curved and bent downwards, with spiral undulation.
Chloroplasts are small, numerous, disk-shaped. In Bacteriastrum hyalinum var. princeps, the inner setae possess
strongly spirally undulated bifurcated ends (Plate 38: e-j).
Distribution:
Bacteriastrum hyalinum is a widely distributed species in temperate to tropical waters. It has been previously
Local occurrence:
Bacteriastrum hyalinum occurred occasionally across Kuwait’s waters, being recorded from 8% of all
collected samples. The species was present throughout the year in low to moderate abundance with higher
concentrations (0.4-2.4∙104 cells/l) recorded in the southern inshore waters from April to September.
Plate 37. Bacteriastrum furcatum.
a, b, d – chains in girdle view; c, f, h – anterior part of chain; e – detail of chain with resting spores; g –
posterior part of chain; i – acid cleaned valve with inner setae; j – acid cleaned valve with terminal setae.
LM, BF illumination (a-h) and phase contrast (i, j); scale bars: a-d – 25 μm; i, j – 20 μm; e-h – 10 μm.
MEDIOPHYCEAE
Plate 38. Bacteriastrum hyalinum and Bacteriastrum hyalinum var. princeps.

a-d – Bacteriastrum hyalinum: a – detail of chain in girdle view; b-d – cells in valve view showing inner setae;
e-j – Bacteriastrum hyalinum var. princeps: e, f – detail of chain in girdle view; g, h – cells in valve view showing
inner setae; i - acid cleaned valve with inner setae; j – acid cleaned valve with terminal setae.
LM, BF illumination (a, e-h) and phase contrast (b-d, i, j); scale bars: a, e – 50 μm; b, g – 25 μm; h-j – 20 μm;
c, d – 10 μm.
Genus Chaetoceros Ehrenberg
The genus Chaetoceros is an important component of the marine plankton worldwide in terms of diversity
and biomass. This is one of the largest genus of marine planktonic diatoms with approximately 200 species
described (Guiry & Guiry, 2017). The genus comprises diatoms with bipolar valves and setae of different
structure to the valves. The species of Chaetoceros are mainly chain-forming and only a few are solitary.
Traditionally, the identification at species level has been based on morphological features observed by light
microscopy. The species within the genus can be distinguished by morphology of the chains (straight, curved
or twisted), shape and dimensions of cells, thickness and direction of the setae, ways of connections in the
chains (fusion of setae, fusion of edge valves and setae, holding of setae), morphology of the terminal setae
(shape, coarseness, and direction), difference of some inner setae from the others, shape and size of aperture
determined by point of origin of setae on valve surface and by point of fusion of sibling setae, number and
shape of chloroplasts, and presence and morphology of resting spores (e.g., Cupp, 1943; Hendey, 1964;
Hasle & Syvertsen, 1997).
Harmful Effect:
Blooms produced by several Chaetoceros species are considered harmful. Large concentrations of
Chaetoceros spp. may clog the gills of farmed fish, and also cause bleeding (Tangen, 1987; Hallegraeff,
1993). The spiny Chaetoceros setae actually penetrate the gill tissue (Bell, 1961), the gills start to produce
mucus, inducing hypoxia (or deficiency of oxygen to the tissues of the body) and hypercapnia (or excessive
amount of carbon dioxide to the blood) (Rensel, 1993). In Kuwait’s waters, blooms of some Chaetoceros
species have been related to local fish mortality (Al-Yamani et al., 2012; see also Sect. 2 in Vol. I).
Chaetoceros affinis Lauder

Plates 39: c-g; 40: a-c
Allen & Cupp, 1935: Fig. 66; Cupp, 1943: Fig. 78; Hendey, 1964: Pl. XVIII: 3; Hasle & Syvertsen, 1997: Pl. 46; Jensen
& Moestrup, 1998: Figs. 30-43; Shevchenko et al., 2006: Figs. 20-24; Throndsen et al., 2007: p. 171; Sunesen et al.,
2008: Fig. 4 A-C; Hoppenrath et al., 2009: Fig. 32 j-l; Kooistra et al., 2010: Figs. 3-9; Doan-Nhu et al., 2014: Figs. 100,
101.
Synonymized names:
Chaetoceros schuttii Cleve 1873
Chaetoceros javanicus Cleve 1873
Chaetoceros ralfsii Cleve 1873
Chaetoceros angulatus Schütt 1895
Frustules are rectangular in girdle view. Valves are elliptical in outline, with slightly concave surface and high
mantle. The cells are with an apical axis of 9-14 µm, united into straight and short chains. Apertures between
sibling cells are narrow and elliptical to lanceolate. The inner setae are long and thin, diverging at an angle
of 10-30° from the chain axis. The terminal setae are thicker and bend in a broad U- or V-shaped curve, with
spines arranged in spirals. Cell possesses a single large chloroplast. The resting spores are biconvex, with
strongly arched primary valve covered with small spines and with rather flat secondary valve.
Distribution:
Chaetoceros affinis is a primarily neritic temperate to warm water species widely distributed worldwide.
The species has been previously reported from Kuwait’s waters (Al-Kandari et al., 2009).
Local occurrence:
Chaetoceros affinis occurred occasionally across Kuwait’s waters, being recorded from 11% of all collected
concentrations (0.1-3.3∙105 cells/l) recorded during the warm season from August to October.
MEDIOPHYCEAE
Plate 39. Chaetoceros spp.

a, b – Chaetoceros subtilis var. abnormis: part of chain, division in progress resulting in formation of new
terminal setae; c-g – Chaetoceros affinis: parts of chains showing inner and terminal setae.
LM, BF illumination (a-c, f, g) and DIC optic (d, e); scale bars: a – 50 μm; b, f, g – 25 μm; c-d – 20 μm.
Chaetoceros brevis Schütt
Plates 40: d-f; 41
Hustedt, 1930: Fig. 403 a, b; Allen & Cupp, 1935: Fig. 70; Cupp, 1943: Fig. 82; Hendey, 1964: Pl. IX: 5; Ricard, 1987:
Figs. 465, 516; Jensen & Moestrup, 1998: Figs. 49-62; Throndsen et al., 2007: p. 171; Sunesen et al., 2008: Fig. 4 D-I;
Doan-Nhu et al., 2014: Figs. 119-122.
Synonymized names:
Chaetoceros hiemalis Cleve 1900
Frustules are rectangular in girdle view, with rounded apices. The girdle occupies about 1/3 of the pervalvar
axis, with suture marked by a notch. Valves are elliptical in outline, with slightly inflated surface. The cells
are with an apical axis of 18-22 µm, united into straight, more or less long chains. Apertures between sibling
cells are wide and slightly constricted in the middle. The inner setae are long and thin, arising inside from the
margins of the valves, with long basal part, almost straight, not so far from apical plane. The terminal setae
are diverging in broad girdle view at a wide angle. Cell possesses a single large chloroplast. The resting spores
are unequally convex, with strongly arched primary valve covered with small spines and with centrally
vaulted secondary valve ornamented with a few longer spines.
Distribution:
Chaetoceros brevis is a neritic warm water species widely distributed in south temperate to tropical waters.
Local occurrence:
Chaetoceros brevis occurred occasionally across Kuwait’s waters, being recorded from 18.5% of all
higher concentrations (0.6-1.6∙105 cells/l) recorded in the inshore waters during the warm season from
March to July. The species is reported for the first time from Kuwait’s waters.
Chaetoceros coarctatus Lauder

Plate 42
Hustedt, 1930: Fig. 370; Allen & Cupp, 1935: Fig. 52; Cupp, 1943: Fig. 62; Crosby & Wood, 1958: Pl. 37: 38; Hendey,
1964: Pl. XII: 1; Simonsen, 1974: p. 30; Hasle & Syvertsen, 1997: Pl. 40; Doan-Nhu et al., 2014: Figs. 35, 36.
Synonymized names:
Chaetoceros rudis Cleve 1901
Frustules are rectangular in girdle view, with rounded apices and strong constrictions at girdle level. The
girdle occupies about 1/3 of the pervalvar axis, with well visible suture. Valves are broadly elliptical in
outline. The cells are with an apical axis of 28-33 µm, united into straight, short, and robust in appearance
chains. Apertures between sibling cells are very narrow. The setae are thick, robust and ornamented with
spines. The inner setae are curved smoothly toward the posterior end of the chain. The posterior terminal
setae are short, thick, strongly curved, and heavily spined. The anterior terminal setae are longer, less robust,
curved toward posterior end, and spined less heavily. Cell possesses numerous small round to elongated
chloroplasts. Some small chloroplasts occur in the basal part of the inner setae. Chains are often found with
more or less abundant epiphytic peritrich ciliate Vorticella oceanica Zacharias attached.
Distribution:
Chaetoceros coarctatus is an oceanic warm water species distributed in subtropical and tropical regions.
Local occurrence:
Chaetoceros coarctatus occurred occasionally across Kuwait’s waters, being recorded from 5.5% of
all collected samples. The species was present throughout the year in low abundance with higher
concentrations (360-480 cells/l) recorded in the inshore waters of Kuwait Bay during the warm season
from May to October.
MEDIOPHYCEAE

a-c – Chaetoceros affinis: a, b – complete short chains, c – part of chain with resting spores; d-f – Chaetoceros
brevis: parts of chains.
LM, BF illumination (a-d, f) and DIC optic (e); scale bars: a, b – 100 μm; d, e – 50 μm; c – 20 μm; f – 10 μm.
Plate 41. Chaetoceros brevis.
a, b – complete short chains; c-f – detail of chains showing inner and terminal setae and single
chloroplast; g – two sibling cells; h – two sibling cells showing resting spores formation; i –
complete resting spore, primary valve.
LM, BF illumination (d) and DIC optic (a-c, e, f); SEM (g-i); scale bars: a – 50 μm; b – 25 μm;
c – 20 μm; d-g – 10 μm; h – 5 μm; i – 2 μm.
MEDIOPHYCEAE
Plate 42. Chaetoceros coarctatus.

a-c – posterior part of chains in girdle view showing inner and posterior terminal setae; d, e – detail of
chain with attached peritrich ciliates Vorticella oceanica (arrowheads); f – chain in valve view; g – acid
cleaned valve with inner setae; h – acid cleaned valve with posterior terminal seta.
LM, BF illumination (a-f) and phase contrast (g, h); scale bars: a – 100 μm; b, d – 50 μm; c, e-h – 20 μm.
Chaetoceros compressus Lauder
Plate 43
Pavillard, 1925: Fig. 72; Hustedt, 1930: Figs. 388, 389; Allen & Cupp, 1935: Fig. 60; Cupp, 1943: Fig. 74; Crosby &
Wood, 1958: Pl. 37: 39; Hendey, 1964: Pl. XVI: 5; Simonsen, 1974: p. 30; Ricard, 1987: Figs. 456, 506-509; Hasle &
Syvertsen, 1997: Pl. 42; Shevchenko et al., 2006: Figs. 31-39; Sunesen et al., 2008: Fig. 5 A-F; Lee & Lee, 2011: Fig. 2
A-F; Doan-Nhu et al., 2014: Figs. 67, 68.
Frustules are rectangular or almost quadrate in girdle view, with rounded apices not touching those of
sibling cells. Valves are elliptical in outline, with slightly convex or flat surface. The cells are with an apical
axis of 17-21 µm, united into straight, long, more or less twisted about the axis chains. Apertures between
sibling cells are wide, four- or six-sided, and slightly constricted in the middle. The setae are arising within
the margin of the valve, with distinct basal part. There are two types of the inner setae termed common and
special. The common inner setae are delicate and diverging almost perpendicular to the chain axis. The
special inner setae are thick, coarse, curved, with spirally twisted row of spines, directed to the chain end.
Terminal setae are thin and delicate, directed almost parallel to the chain axis. Cell possesses a few small
round chloroplasts.
Distribution:
Chaetoceros compressus is a neritic, boreal to south temperate species. It has been previously reported from
the Arabian Gulf (Simonsen, 1974) and from Kuwait’s waters (Al-Kandari et al., 2009).
Local occurrence:
Chaetoceros compressus occurred commonly across Kuwait’s waters, being recorded from 52% of all
higher concentrations (1.8-3.6∙105 cells/l) recorded during the period from March to September.
Chaetoceros costatus Pavillard

Plate 44
Cupp, 1943: Fig. 79; Hendey, 1964: Pl. XIX: 3; Ricard, 1987: Fig. 460; Hasle & Syvertsen, 1997: Pl. 46; Jensen &
Moestrup, 1998: Figs. 109-111; Throndsen et al., 2007: p. 172; Hoppenrath et al., 2009: Fig. 33 g-i; Kooistra et al., 2010:
Figs. 30-37; Lee & Lee, 2011: Fig. 1 A-I; Doan-Nhu et al., 2014: Figs. 110, 111.
Synonymized names:
Chaetoceros adhaerens Mangin 1913
Frustules are rectangular in girdle view. Valves are narrowly elliptical in outline, with numerous intercalary
bands. The cells are with an apical axis of 16-23 µm, united into straight and long chains. Apertures between
sibling cells are narrow, elliptical or spindle-shaped, and shorter than the apical axis due to the presence of
symmetrical submarginal projections close to each pole of valve, joining with those of the valve belonging to
the sibling cell. The terminal and inner setae are similar and delicate. The setae are arising from the rounded
angles on the inner sides of the valves, emerge perpendicular to the chain axis, cross without a basal part and
diverge strongly from the apical plane. Cell possesses a single plate-shaped chloroplast lying on the girdle.
Distribution:
Chaetoceros costatus is a neritic warm water species. It has been previously reported from Kuwait’s waters
Local occurrence:
Chaetoceros costatus rarely occurred across Kuwait’s waters, being recorded from less than 4% of all
collected samples. The species was present in low to moderate abundance with higher concentrations
(0.4-1.3∙104 cells/l) recorded during the warm months of June and July.
MEDIOPHYCEAE
Plate 43. Chaetoceros compressus.

a-d – parts of chains in broad (a, b, d) and narrow (c) girdle view showing common and special
inner setae; e, f – detail of chains in girdle view showing chloroplasts; g, h – two sibling valves
with special setae.
LM, BF illumination (a-f); SEM (g, h); scale bars: a, d – 50 μm; b, c – 25 μm; e-h – 10 μm.
Plate 44. Chaetoceros costatus.
a-f – parts of chains in girdle view showing inner setae and chloroplasts.
LM, BF illumination; scale bars: a, b, d, f – 25 μm; c, e – 10 μm.

Chaetoceros curvisetus Cleve
Plate 45
Pavillard, 1925: Fig. 82; Hustedt, 1930: Fig. 426; Cupp, 1943: Fig. 93; Hendey, 1964: Pl. XVII: 6; Simonsen, 1974: p.
30; Priddle & Fryxell, 1985: p. 44-45; Ricard, 1987: Figs. 475, 476, 526, 527; Hasle & Syvertsen, 1997: Pl. 44; Jensen
& Moestrup, 1998: Figs. 114-116; Shevchenko et al., 2006: Figs. 43-46; Throndsen et al., 2007: p. 172; Sunesen et al.,
2008: Fig. 6 A-F; Kooistra et al., 2010: Figs. 38-47; Doan-Nhu et al., 2014: Figs. 155, 156, 158.
Frustules are rectangular in girdle view. The girdle occupies about 1/2 of the pervalvar axis, with suture
marked by a notch. Valves are elliptical in outline, with low mantle. The cells are with an apical axis of
9-12 µm, united into long, spirally curved chains. The sibling cells are connected by conspicuous corners.
Apertures between sibling cells are wide, oval to almost circular. The setae are thin, arising from corners of
cells, with characteristic curvature, all directed toward the outside of the spiral. Cell possesses a single plate-
shaped chloroplast lying on the girdle. The resting spores are smooth, without spines, originating in pairs
united by valves of sibling cells. The primary valve is broadly convex and covered by a siliceous sheath. The
secondary valve is slightly convex or almost flat.
Distribution:
Chaetoceros curvisetus is a cosmopolitan neritic widely distributed species. It has been previously reported
from the Indian Ocean (Simonsen, 1974) and from Kuwait’s waters (Al-Yamani et al., 2004; Al-Kandari
et al., 2009).
MEDIOPHYCEAE
Local occurrence:
Chaetoceros curvisetus occurred frequently across Kuwait’s waters, being recorded from 49% of all
higher concentrations (0.5-2.2∙106 cells/l) recorded during the period from March to September.
Harmful Effect:
Blooms produced by Chaetoceros curvisetus are potentially harmful. Large concentrations of Chaetoceros
curvisetus were associated with local fish mortality in June-July, 2011 in Kuwait Bay and in small
semienclosed bays at Ras Salmiya, Marina Crescent, and the Rescue Dock. The extremely high water
temperature coupled with high concentration of organic carbon supplied by the die-off phase of
phytoplankton bloom of this species, in addition to water stratification in the shallow semienclosed
water bodies along Kuwait’s shores resulted in strong oxygen depletion, with negative impact on the
marine biota (for details see also Sect. 2 in Vol. I). This species has been included in the list of potentially
harmful species for Kuwait’s marine environment (Al-Yamani et al., 2012) due to its capacity to form
high biomass blooms in Kuwait’s waters.
Chaetoceros danicus Cleve

Plate 46
Cupp, 1943: Fig. 65; Hendey, 1964: Pl. X: 5; Hernández-Becerril, 1996: Pls. 13: 1-7, 14: 1-8; Jensen & Moestrup, 1998:
Figs. 10-15; Throndsen et al., 2007: p. 166; Sunesen et al., 2008: Fig. 2 A-G; Hoppenrath et al., 2009: Fig. 31 d, e;
Kooistra et al., 2010: Figs. 48-53; Doan-Nhu et al., 2014: Figs. 17, 18.
Frustules are rectangular in girdle view. Valves are elliptical to nearly circular in outline, with high mantle.
The cells are with an apical axis of 11-18 µm, solitary or united into straight short 2-6-celled chains. Apertures
between sibling cells are very narrow. The setae are long and thick, arising from the corners of the valves,
extending perpendicular to pervalvar axis. The setae are not on a plane and reach out all directions so
that they form a cross in valve view. Cell possesses numerous small round chloroplasts. Small elongated
chloroplasts occur within the setae.
Distribution:
Chaetoceros danicus is a cosmopolitan neritic widely distributed species.
Local occurrence:
Chaetoceros danicus occurred occasionally across Kuwait’s waters, being recorded from 24% of all
higher concentrations (0.3-1.3∙104 cells/l) recorded during the period from March to October. The
species is reported for the first time from Kuwait’s waters.
Plate 45. Chaetoceros curvisetus.
a-e – curved chains in narrow girdle view; f – part of chain in broad girdle view; g – part of chain showing
resting spore formation; h – part of chain with complete resting spores.
LM, BF illumination (a, b, f-h) and DIC optic (c, d); scale bars: a-d – 20 μm; e-h – 10 μm.

MEDIOPHYCEAE
Plate 46. Chaetoceros danicus.

a, d, h, i – short chains in valve view; b, c – single cells in girdle view; e-g – single cells in valve view.
LM, BF illumination (a-d), DF illumination (f, h, i) and DIC optic (e, g); scale bars: a, b, e, f, h, i – 50 μm; c,
d, g – 25 μm.
Harmful Effect:
In the western coast of USA, Chaetoceros danicus has been involved in mass mortality of fishes by anoxia,
due to gill clogging caused by mucus production generated in response to the damage of spines on the
gills (Horner et al., 1997).
Chaetoceros decipiens Cleve

Plate 47
Pavillard, 1925: Fig. 70A; Hustedt, 1930: Fig. 383; Cupp, 1943: Figs. 70A, 70B-a, b; Crosby & Wood, 1958: Pl. 37: 44;
Hendey, 1964: Pl. XII: 2; Simonsen, 1974: p. 30; Ricard, 1987: Figs. 450, 495-499; Hasle & Syvertsen, 1997: Pl. 42;
Jensen & Moestrup, 1998: Figs. 122-131; Shevchenko et al., 2006: Figs. 53-58; Throndsen et al., 2007: p. 169; Sunesen
et al., 2008: Fig. 7 A-F; Hoppenrath et al., 2009: Figs. 6, 34 a-c; Doan-Nhu et al., 2014: Figs. 48-51; Li et al., 2017:
Figs. 1-3, 20 E, F.
Synonymized names:
Chaetoceros decipiens var. grunowii (Schütt) Cleve 1897
Chaetoceros grunowii Schütt 1895
Frustules are rectangular in girdle view, with sharp corners touching those of the sibling cell. The girdle
occupies about 1/2 of the pervalvar axis, with suture marked by a notch. Valves are narrowly elliptical in
outline, with high mantle. The cells are with an apical axis of 21-35 µm, united into straight, stiff, flattened,
and more or less long chains. Apertures between sibling cells are wide and elliptical. The setae are long,
thick, straight, with small spines arranged in longitudinal rows. The sibling inner setae are fused for a long
distance, before diverging at an angle of 10-25° from the chain axis. The terminal setae are thicker and
diverging from cell corners and bend in a broad U- or V-shaped curve, nearly parallel to the chain axis at the
ends. Cell possesses a few small round chloroplasts.
Distribution:
Chaetoceros decipiens is an oceanic species. It has been previously reported from the Indian Ocean
Local occurrence:
Chaetoceros decipiens occurred occasionally across Kuwait’s waters, being recorded from 19% of all
higher concentrations (0.8-4.3∙105 cells/l) recorded during the period from January to May.
Chaetoceros denticulatus Lauder

Plate 48, 49: a-c
Allen & Cupp, 1935: Fig. 53; Crosby & Wood, 1958: Pl. 37: 45; Simonsen, 1974: p. 31; Doan-Nhu et al., 2014: Figs.
20, 22.
Synonymized names:
Chaetoceros denticulatus f. angusta Hustedt ex Simonsen 1987
Chaetoceros denticulatus f. lata Hustedt ex Schmidt 1920
Chaetoceros nanodenticulatus Okamura 1907
Frustules are rectangular in girdle view. The girdle occupies more than 1/2 of the pervalvar axis, with suture
marked by a notch. Valves are elliptical to nearly circular in outline, with raised corners and small spine in
the center. The cells are with an apical axis of 28-33 µm, united into straight short chains. Apertures between
sibling cells are elliptical to hexagonal. The setae are long, straight and thick, ornamented with small spines,
arising close to valve center, at first almost parallel with chain axis, then with an elbow, turning at right
angles. There is a small tooth-like process on the inner side of setae at the crossover of the setae of the sibling
cells. Cell possesses numerous small rod-shaped chloroplasts. Small elongated chloroplasts occur within the
setae.

MEDIOPHYCEAE
Plate 47. Chaetoceros decipiens.

a – part of chain in girdle view; b – complete short chain showing inner and terminal setae; c – chain in
division resulting in formation of new terminal setae in the middle; d – part of chain in girdle view showing
fusion of inner setae (arrowheads); e – acid cleaned sibling valves with fused setae (arrowheads).
LM, BF illumination (a, d), DF illumination (b), and phase contrast (c, e); scale bars: a-c – 50 μm; d, e – 20
μm.
Plate 48. Chaetoceros denticulatus.
a, b – complete short chains in girdle view; c-f – parts of chains in girdle view showing chloroplasts inside
setae and tooth-like processes on the inner side of setae (arrowheads).
LM, BF illumination; scale bars: a, b – 100 μm; c, d – 25 μm; e, f – 20 μm.

MEDIOPHYCEAE

a-c – Chaetoceros denticulatus: a – complete short chain; b – detailed sibling cells; c – acid cleaned sibling
frustules showing tooth-like processes on the inner side of setae (arrowheads); d-g – Chaetoceros diadema:
parts of chains in broad girdle view.
LM, BF illumination (d-g), DF illumination (a), and phase contrast (b, c); scale bars: d – 100 μm; a – 50 μm;
b, e – 25 μm; f, g – 20 μm; c – 10 μm.
Distribution:
Chaetoceros denticulatus is a typical species in tropical areas of the Indian Ocean with neritic preference.
It has been previously reported from the Arabian Gulf (Simonsen, 1974) and from Kuwait’s waters (Al-
Yamani et al., 2004; Al-Kandari et al., 2009).
Local occurrence:
Chaetoceros denticulatus occurred frequently across Kuwait’s waters, being recorded from 41% of all
higher concentrations (1.4-3.1∙104 cells/l) recorded in Kuwait Bay during the period from November to
January.
Chaetoceros diadema (Ehrenberg) Gran

Plate 49: d-g
Hendey, 1964: Pl. X: 1; Hernández-Becerril, 1996: Pls. 34: 1-6, 35: 1-8; Hasle & Syvertsen, 1997: Pl. 46; Jensen &
Moestrup, 1998: Figs. 132 -141; Shevchenko et al., 2006: Figs. 59-63; Throndsen et al., 2007: p. 172; Sunesen et al.,
2008: Fig. 7 G-K; Hoppenrath et al., 2009: Fig. 34 d-k; Kooistra et al., 2010: Figs. 59-63; Doan-Nhu et al., 2014: Figs.
136-138.
Synonymized names:
Syndendrium diadema Ehrenberg 1854
Chaetoceros groenlandicus Cleve 1896
Chaetoceros ralfsii Schütt 1896
Chaetoceros paradoxus Péragallo 1897
Frustules are rectangular in girdle view. Valves are elliptical in outline, with slightly convex surface and low
mantle. The cells are with an apical axis longer than pervalvar one, of 12-16 µm, united into straight or
slightly twisted long chains. Apertures between sibling cells are wide, with a slight central constriction. The
inner setae are thin and delicate, with short basal part, cross over at the colony margin. The terminal setae
are slightly thicker. Cell possesses a single plate-shaped chloroplast lying on the girdle.
Distribution:
Chaetoceros diadema is a cosmopolitan neritic species widely distributed in temperate waters.
Local occurrence:
Chaetoceros diadema is of low occurrence in Kuwait’s waters, and was never observed in large numbers.
The present observation constitutes the first record of this species for Kuwait.
Chaetoceros didymus Ehrenberg

Plate 50: a-e
Allen & Cupp, 1935: Fig. 61; Cupp, 1943: Fig. 75A; Crosby & Wood, 1958: Fig. 47; Hendey, 1964: Pl. XVII: 2; Ricard,
1987: Fig. 457; Hasle & Syvertsen, 1997: Pl. 43; Jensen & Moestrup, 1998: Figs. 142-144, 146; Shevchenko et al., 2006:
Figs. 64-70; Throndsen et al., 2007: p. 170; Sunesen et al., 2008: Fig. 8 A-J; Hoppenrath et al., 2009: Fig. 34 l, m; Doan-
Nhu et al., 2014: Figs. 86-89.
Frustules are rectangular in girdle view. Valves are elliptical in outline, with raised corners, low mantle, and
a dome-shaped centrally located protuberance. The cells are with an apical axis of 16-22 µm, united into
straight long chains. Apertures between sibling cells are wide, panduriform, with a central constriction. The
setae are coarse, long and straight. The inner setae are fusing at the margin or near the margin of the chain,
and diverging in acute angle in valvar plane. The terminal setae are directed towards the chain axis. Cell
possesses two plate-shaped chloroplasts. The resting spores are paired, united by thick setae, smooth, with
protuberance on the primary valve and flat secondary valve.
Distribution:
Chaetoceros didymus is a neritic species widely distributed in temperate and warm waters (Hasle &
Syvertsen, 1997; Guiry & Guiry, 2017). The species has been previously reported from the Arabian Gulf
(Simonsen, 1974) and from Kuwait’s waters (Al-Kandari et al., 2009).

MEDIOPHYCEAE

a-e – Chaetoceros didymus: a – complete chain in broad girdle view; b-d – detail of chains showing
chloroplasts and centrally located protuberances (arrowheads); e – sibling valves showing two pairs
of reduced setae and two pairs of well-developed resting spores; f-i – Chaetoceros protuberans: f –
complete short chain in broad girdle view with attached flagellates; g – detail of chain showing
chloroplasts and centrally located protuberances (arrowheads); h – two sibling valves; i - valve.
LM, BF illumination (a-g); SEM (h, i); scale bars: a – 50 μm; b, f – 25 μm; c-d, g – 20 μm; i – 5 μm.
Local occurrence:
Chaetoceros didymus occurred occasionally across Kuwait’s waters, being recorded from 12% of all
higher concentrations (1.2-6.6∙104 cells/l) recorded in July.
Chaetoceros distans Cleve

Plate 51
Allen & Cupp, 1935: Fig. 68; Hernández-Becerril, 1996: Pls. 30, 31; Doan-Nhu et al., 2014: Figs. 124-128.
marked by a notch. Valves are elliptical in outline, with slightly inflated surface. The cells are with an apical
axis of 17-24 µm, united into straight, more or less long chains. Apertures between sibling cells are wide
and rectangular. The inner setae are long, arising parallel to the pervalvar axis before crossing over and rise
out perpendicular to the chain axis. The terminal setae are diverging at a wide angle. Cell possesses a single
plate-shaped chloroplast.
Remarks:
Chaetoceros distans is regarded as a synonym of Chaetoceros dichaeta Ehrenberg (Guiry & Guiry, 2017)
or Chaetoceros laciniosus Schütt (Shevchenko et al., 2006), but presented here as a separate taxon due to
its morphological distinction in size, robustness, orientation of the terminal setae, and resting spores
observed in Kuwait’s waters.
Distribution:
Chaetoceros distans is a neritic warm water species.
Local occurrence:
Chaetoceros distans is of low occurrence in Kuwait’s waters, and was never observed in large numbers.
The species is reported for the first time from Kuwait’s waters.
Chaetoceros diversus Cleve

Plate 52
Hustedt, 1930: Fig. 409; Allen & Cupp, 1935: Fig. 71; Cupp, 1943: Fig. 87; Hendey, 1964: Pl. XVII: 4; Simonsen, 1974:
p. 31; Shevchenko et al., 2006: Figs. 71-73; Lee & Lee, 2011: Fig. 4 A-F; Doan-Nhu et al., 2014: Figs. 102-104, 141-144.
marked by a notch. Valves are elliptical in outline, with flat or slightly inflated surface and high mantle. The
cells are with an apical axis of 11-14 µm, united into straight short chains. Apertures between sibling cells
are very narrow, slit-like. There are two types of the inner setae termed common and special. The common
inner setae are thin, more or less curved, and usually slightly turned toward the chain ends. The special inner
setae are thick, increasing in thickness from the base out, then becoming thinner again near the ends. The
first two-thirds of setae is almost straight and at a sharp angle from the chain axis, then turning gradually
and at obtuse angle toward one or the other end of the chain, and running almost parallel to the chain axis
in the outer part. The thicker setae are ornamented with spirally arranged small spines. The terminal setae
are thin and differ from others in position – the basal parts are more or less U-shaped, and the outer parts are
nearly parallel to chain axis. Cell possesses a single plate-shaped chloroplast.
Distribution:
Chaetoceros diversus is a neritic, tropical and subtropical species. It has been previously reported from the
Arabian Gulf (Simonsen, 1974) and from Kuwait’s waters (Al-Yamani et al., 2004; Al-Kandari et al.,
2009).
Local occurrence:
Chaetoceros diversus occurred frequently across Kuwait’s waters, being recorded from 30% of all collected
concentrations (0.5-1.1∙105 cells/l) recorded in the southern inshore waters in July.

MEDIOPHYCEAE
Plate 51. Chaetoceros distans.
a – complete chain in broad girdle view; b – detail of chain showing chloroplasts, inner and terminal setae;
c – detail of chain; d-f – formation of resting spores.
LM, BF illumination (a, b, d); SEM (c, e, f); scale bars: a – 50 μm; b-d – 20 μm; e, f – 10 μm.
Plate 52. Chaetoceros diversus.
a – complete short chain in broad girdle view; b-e – parts of chains showing common and special inner
setae, and chloroplasts; f – detail of chain; g – two sibling valves.
LM, BF illumination (a-e); SEM (f, g); scale bars: a-d – 20 μm; e-g – 10 μm.

Chaetoceros laciniosus Schütt
Plate 53
Allen & Cupp, 1935: Fig. 69; Cupp, 1943: Fig. 80; Crosby & Wood, 1958: Fig. 50; Hendey, 1964: Pl. XIII: 2; Ricard,
1987: Figs. 463, 464; Hernández-Becerril, 1996: Pls. 30: 1-6, 31: 1-8; Hasle & Syvertsen, 1997: Pl. 43; Jensen &
Moestrup, 1998: Figs. 153-158; Shevchenko et al., 2006: Figs. 80-83; Throndsen et al., 2007: p. 170; Sunesen et al.,
2008: Fig. 9 C, D; Doan-Nhu et al., 2014: Figs. 129-131.
Frustules are rectangular in girdle view, with raised corners. The girdle occupies about 1/3 of the pervalvar
axis, with suture marked by a notch. Valves are elliptical in outline, with flat or slightly inflated surface and
low mantle. The cells are with an apical axis of 10-14 µm, united into long, straight or slightly curved chains.
Apertures between sibling cells are wide and rectangular. The inner setae are long, with long basal part,
arising parallel to the pervalvar axis before crossing over and diverging perpendicular to the chain axis, with
far outer part bent toward one chain end. The terminal setae are curving smoothly in direction to the chain
axis, often converging or crossing at the ends. Cell possesses two plate-shaped chloroplasts.
Distribution:
Chaetoceros laciniosus is a neritic tropical and temperate species. The species has been previously reported
from Kuwait’s waters (Al-Kandari et al., 2009).
Local occurrence:
MEDIOPHYCEAE
Chaetoceros laciniosus occurred occasionally across Kuwait’s waters, being recorded from 21.5% of all
higher concentrations (1.5-3.7∙105 cells/l) observed in the inshore waters during the warm season from
May to July.
Chaetoceros lauderi Ralfs

Plate 54: a-g
1958: Pl. 37: 51; Hendey, 1964: Pl. XIII: 3; Ricard, 1987: Fig. 455; Jensen & Moestrup, 1998: Fig. 159; Sunesen et al.,
2008: Fig. 10 A-F; Hoppenrath et al., 2009: Fig. 34 n, o; Kooistra et al., 2010: Figs. 64-70; Doan-Nhu et al., 2014: Fig.
66.
Synonymized names:
Chaetoceros weissflogii Schütt 1895
Frustules are rectangular in girdle view, with pervalvar axis much longer than apical one. Valves are broadly
elliptical to almost circular in outline, with flat surface and low mantle. The cells are with an apical axis
of 19-22 µm, united into long chains, twisted about the chain axis. Apertures between sibling cells are
narrow, linear-elliptic. The setae are long, arising from the corners of the valves at right angles to the chain
axis. The inner setae are diverging in acute angles to the valvar plane. The terminal setae bend in a broad
U-shaped curve, directed to the chain axis. Cell possesses numerous round chloroplasts. The resting spores
are with unequal valves. The primary valve is capitate, covered by conspicuous spines and surrounded by a
marginal ring of spines. The secondary valve is less convex or almost flat, with a marginal ring of long spines
becoming filament-like distally.
Distribution:
Chaetoceros lauderi is a neritic south temperate species.
Local occurrence:
Chaetoceros lauderi occurred occasionally across Kuwait’s waters, being recorded from 17% of all
July to September. The present observation constitutes the first record of this species for Kuwait.
Plate 53. Chaetoceros laciniosus.
a, b – complete short chains in broad girdle view; c-f – parts of chains showing inner setae and chloroplasts.
LM, BF illumination (a, b, d-f) and DIC optic (c); scale bars: a-c – 50 μm; d – 20 μm; e, f – 10 μm.

MEDIOPHYCEAE

a-g – Chaetoceros lauderi: a-c – parts of chains in girdle view showing inner setae and torsion;
d – part of chain with resting spore; e-g – resting spores; h-k – Chaetoceros vanheurckii: h – part
of chain showing inner setae and chloroplasts; i – detailed sibling cells; j, k – resting spores.
LM, BF illumination (a-f, h-k); SEM (g); scale bars: a, b, h – 25 μm; c, d – 20 μm; e, f, i – 10
μm; g, j, k – 5 μm.
Chaetoceros lorenzianus Grunow
Plate 55, 56
Pavillard, 1925: Fig. 70B; Hustedt, 1930: Fig. 385; Allen & Cupp, 1935: Fig. 58; Cupp, 1943: Fig. 71; Crosby & Wood,
1958: Pl. 38: 52; Hendey, 1964: Pl. XVI: 1; Simonsen, 1974: p. 31; Ricard, 1987: Figs. 452, 453; Hasle & Syvertsen,
1997: Pl. 42; Jensen & Moestrup, 1998: Figs. 160-165; Shevchenko et al., 2006: Figs. 84-89; Throndsen et al., 2007: p.
169; Sunesen et al., 2008: Fig. 11 A-F; Hoppenrath et al., 2009: Fig. 35 a-e; Kooistra et al., 2010: Figs. 71-79; Doan-
Nhu et al., 2014: Figs. 53-60.
Synonymized names:
Chaetoceros cellulosum Lauder 1864
Frustules are rectangular in girdle view, with sharp corners touching those of the sibling cell. The girdle
occupies about 1/2 of the pervalvar axis, with suture marked by a notch. Valves are narrowly elliptical
in outline, with high mantle. The cells are with an apical axis of 12-33 µm, united into straight or slightly
arched, stiff, flattened, and more or less long chains. Apertures between sibling cells are wide and elliptical.
The setae are long, thick, straight, with small spines arranged in longitudinal rows. The inner setae are
arising from the corners of valves, fusing near the chain margin, perpendicular or diverging in an angle of
25-45° to the chain axis. The terminal setae are thicker, diverging for their entire length or bend in a broad
U-shaped curve. Cell possesses a few small plate-shaped chloroplasts.
Distribution:
Chaetoceros lorenzianus is a neritic tropical and temperate species. It has been previously reported from
the Arabian Gulf (Simonsen, 1974) and from Kuwait’s waters (Al-Yamani et al., 2004; Al-Kandari et
al., 2009).
Local occurrence:
Chaetoceros lorenzianus occurred commonly across Kuwait’s waters, being recorded from 62% of all
higher concentrations (0.2-1.3∙105 cells/l) recorded in the offshore waters during the spring in May.
Harmful Effect:
Chaetoceros lorenzianus can be considered as a harmful bloom-forming species. Its blooms have been
associated with mass mortality of anchovies in the coastal Costa Rican waters (Vargas-Montero & Freer,
2004).
Chaetoceros paradoxus Cleve

Plate 57: a-c
Allen & Cupp, 1935: Fig. 67; Crosby & Wood, 1958: Fig. 54; Doan-Nhu et al., 2014: Figs. 112-116.
Frustules are rectangular in girdle view, with pervalvar axis usually much longer than apical one. The girdle
is narrow and constricted. Valves are broadly elliptical in outline. The cells are with an apical axis of 18-23
µm, united into short chains, slightly twisted about the chain axis. Apertures between sibling cells are wide,
elliptic. The setae are robust, long, arising from the corners of the valves at right angles to the chain axis.
The terminal and inner setae are not in the same plane. Cell possesses two large plate-shaped chloroplasts.
Distribution:
Chaetoceros paradoxus is a neritic warm water species.
Local occurrence:
Chaetoceros paradoxus rarely occurred in Kuwait’s waters, and was never observed in large numbers. The

MEDIOPHYCEAE
Plate 55. Chaetoceros lorenzianus.

a – long chain in division resulting in formation of new terminal setae in the middle; b-e – short complete
chains in broad girdle view showing inner and terminal setae and chloroplasts.
LM, BF illumination (a, c), DF illumination (b), and DIC optic (d, e); scale bars: 50 μm.
Plate 56. Chaetoceros lorenzianus.
a-e – parts of chains in broad girdle view showing inner and terminal setae, connection between sibling cells,
and chloroplasts; f – part of two acid cleaned sibling valves with inner setae; g – two sibling valves.
LM, BF illumination (a), DIC optic (b-d), and phase contrast (e, f); SEM (g); scale bars: a-g – 20 μm; f, g – 10
μm.

MEDIOPHYCEAE
a-c – Chaetoceros paradoxus: a, c – short complete chain in girdle view; b – part of chain; d, e – Chaetoceros
pseudobrevis: d – short complete chain in broad girdle view; e – detailed sibling cells showing chloroplasts and
pigmentation inside inner setae (arrowheads).
LM, BF illumination; scale bars: a-d – 50 μm; e – 20 μm.
Chaetoceros peruvianus Brightwell
Plate 58
Pavillard, 1925: Fig. 67; Hustedt, 1930: Figs. 379-381; Allen & Cupp, 1935: Figs. 56, 57; Cupp, 1943: Fig. 68 a-c;
Crosby & Wood, 1958: Pl. 34: 49; Hendey, 1964: Pl. IX: 3; Simonsen, 1974: p. 31; Priddle & Fryxell, 1985: p. 40-41;
Ricard, 1987: Fig. 448; Hasle & Syvertsen, 1997: Pl. 38; Shevchenko et al., 2006: Figs. 13-16; Sunesen et al., 2008: Fig.
3 A-D; Kooistra et al., 2010: Figs. 80-86; Doan-Nhu et al., 2014: Figs. 31-34.
Synonymized names:
Chaetoceros peruvianum var. currens Péragallo 1897
Chaetoceros peruvio-atlanticum Karsten 1907
Chaetoceros convexicornis Margin 1919
Chaetoceros chilensis Krasske 1941
Frustules are rectangular in girdle view, heterovalvar. The girdle is narrow. The cells are with an apical axis
of 18-25 µm, usually solitary or seldom united into short 2-3-celled chains. Apertures between sibling cells
are very narrow. The anterior valve is convex, the posterior valve is flat to slightly concave. The setae are
robust, long, transversely striated at the base, and ornamented with longitudinal rows of spines. All setae
directed towards the same end of the cell. The setae of anterior valve are fused together after a short base in
the valve centre, turning sharply and running backward in more or less wide, outwardly convex curves, and
more or less divergent to convergent at the end. The setae of posterior valve are arising from the corners,
slightly convex toward outside, more nearly parallel to chain axis than those of the anterior valve, at the end,
more or less divergent or even convergent. Cell possesses numerous small round to elongate chloroplasts.
Small rod-shaped chloroplasts occur within the setae.
Distribution:
Chaetoceros peruvianus is an oceanic species distributed from south temperate to tropical waters. It has
been previously reported from the Arabian Gulf (Simonsen, 1974) and from Kuwait’s waters (Al-Yamani
et al., 2004; Al-Kandari et al., 2009).
Local occurrence:
Chaetoceros peruvianus occurred frequently across Kuwait’s waters, being recorded from 31% of all
June to September.
Harmful Effect:
Chaetoceros peruvianus is a nontoxic bloom-forming species. Although no harmful incidents have been
reported, the species may be regarded as potentially harmful due to its morphology (Hansen et al., 2001).
Chaetoceros peruvianus Brightwell f. gracilis (Schröder) Hustedt

Plate 59: a-c
Hustedt, 1930: Fig. 381 b; Cupp, 1943: Fig. 68 d-f.
Synonymized names:
Chaetoceros peruvianus var. gracilis Schröder 1900
Compared to the type, cells are more slender and delicate, with an apical axis of 10-15 µm, and higher valve
mantle. The setae are long and thin, without visible ornamentation.

MEDIOPHYCEAE
Plate 58. Chaetoceros peruvianus.

a – 2-celled chain in girdle view; b – part of chain; c – part of seta with spines and chloroplasts inside; d –
single cell in girdle view; e – detailed single cell; f – acid cleaned frustule in girdle view; g – detail of frustule;
h – detailed upper valve; i – detail of seta.
LM, BF illumination (a-e) and phase contrast (f); SEM (g-i); scale bars: a, d – 100 μm; b, e – 50 μm; f – 25
μm; c, g-i – 10 μm.
a-c – Chaetoceros peruvianus f. gracilis: a, b – single cells in girdle view; c – detailed single cell; d – Chaetoceros
tennuissimus: single cells in girdle view; e-g – Chaetoceros seiracanthus: e – part of chain; f, g – resting spores;
h - Chaetoceros sp.: single cells in girdle view.
LM, BF illumination (a, e, h) and DIC optic (b-d); SEM (f, g); scale bars: a, b – 50 μm; c – 25 μm; e, h – 20
μm; d – 10 μm; f, g – 5 μm.

Chaetoceros protuberans Lauder
Plate 50: f-i
Allen & Cupp, 1935: Fig. 62; Cupp, 1943: Fig. 75B-a; Jensen & Moestrup, 1998: Fig. 145 (as Chaetoceros didymus var.
protuberans); Shevchenko et al., 2006: Figs. 97-100; Hoppenrath et al., 2009: Fig. 35f; Kooistra et al., 2010: Figs. 87-95;
Doan-Nhu et al., 2014: Figs. 90, 91.
Synonymized names:
Chaetoceros didymus var. protuberans (Lauder) Gran et Yendo 1914
Frustules are rectangular in girdle view. Valves are elliptical in outline, with raised corners, low mantle,
and a prominent dome-shaped centrally located protuberance. The cells are with an apical axis of 13-18
µm, united into straight short chains. Apertures between sibling cells are very wide, often more than cells
pervalvar axis length. The inner setae are thin and straight, with long basal part. Each seta joins with the
seta of the sibling cell outside the margin of the chain, and diverging in acute angle in valvar plane. The
terminal setae bend in a broad V-shaped curve and are directed towards the chain axis. Cell possesses two
plate-shaped chloroplasts.
Distribution:
Chaetoceros protuberans is a neritic species widely distributed in temperate and warm waters.
MEDIOPHYCEAE
Local occurrence:
Chaetoceros protuberans rarely occurred in Kuwait’s waters, and was never observed in large numbers. The
Chaetoceros pseudobrevis Pavillard

Plate 57: d, e
Jensen & Moestrup, 1998: Figs. 167-175; Doan-Nhu et al., 2014: Figs. 123, 132, 133.
Frustules are rectangular in girdle view. Valves are elliptical in outline, with slightly inflated surface. The
cells are with an apical axis of 20-26 µm, united into straight short chains. Apertures between sibling cells
are wide and constricted in the middle. The inner setae are long and thick, arising inside from the margins
of the valves, with a short basal part, almost straight, perpendicular to the chain axis. The terminal setae
are diverging in broad girdle view at a wide angle. Cell possesses a single large chloroplast. Numerous small
elongate pigmented granules are distributed inside the setae.
Remarks:
Chaetoceros pseudobrevis is currently regarded as a synonym of Chaetoceros brevis Schütt (Guiry & Guiry,
2017), but presented here as a separate taxon due to its morphological distinction from C. brevis in size,
robustness, and pigmentation of the setae in Kuwait’s specimens.
Distribution:
Chaetoceros pseudobrevis is a neritic temperate to warm water species.
Local occurrence:
Chaetoceros pseudobrevis rarely occurred in Kuwait’s waters, and never in large numbers. The species is
reported for the first time from Kuwait’s waters.
Chaetoceros pseudocurvisetus Mangin
Plate 60
Pavillard, 1925: Fig. 83; Hustedt, 1930: Fig. 427; Allen & Cupp, 1935: Fig. 73; Cupp, 1943: Fig. 94; Hendey, 1964: Pl.
XVIII: 1; Hasle & Syvertsen, 1997: Pl. 44; Shevchenko et al., 2006: Figs. 107-109; Kooistra et al., 2010: Figs. 96, 97;
Doan-Nhu et al., 2014: Figs. 159-164.
Frustules are rectangular in girdle view. Valves are broadly elliptical in outline, with high mantle. The cells
are with an apical axis of 10-14 µm, united into long curved to helical chains. Apertures between sibling
cells are narrow, lenticular. There are two short projections along each valve margin, medial to the point of
emergence of a seta, merging with the corresponding pair on the valve of the sibling cell to form four small
lateral apertures. The setae are thin and delicate, arising from the apices of the valve and cross at their point
of origin. All setae point towards the convex side of the chain. Cell possesses a single large lobed chloroplast.
The resting spores are smooth, with equally convex valves, and a collar around both valves.
Distribution:
Chaetoceros pseudocurvisetus is a neritic, tropical to subtropical species. It has been previously reported
from Kuwait’s waters (Al-Yamani et al., 2004; Al-Kandari et al., 2009).
Local occurrence:
Chaetoceros pseudocurvisetus occurred frequently across Kuwait’s waters, being recorded from 42% of all
higher concentrations (0.8-2.8∙106 cells/l) recorded in the inshore waters of Kuwait Bay during the fall
months of September and October.
Harmful Effect:
Blooms produced by Chaetoceros pseudocurvisetus could be harmful in Kuwait’s marine environment. Large
concentrations of Chaetoceros pseudocurvisetus were associated with local fish mortality in June-July, 2011
in Kuwait Bay and in small semienclosed bays at Ras Salmiya, Marina Crescent, and Rescue Dock (for
details see also Sect. 2, Vol. I). This species has been included in the list of potentially harmful species for
Kuwait’s marine environment (Al-Yamani et al., 2012) due to its capacity to form high biomass blooms
in Kuwait’s waters.
Chaetoceros seiracanthus Gran

Plate 59: e-g
Hustedt, 1930: Fig. 405; Cupp, 1943: Fig. 84; Hendey, 1964: Pl. XV: 1; Hasle & Syvertsen, 1997: Pl. 46; Jensen &
Moestrup, 1998: Figs. 191-195; Doan-Nhu et al., 2014: Figs. 139, 140.
Frustules are rectangular in girdle view, with rounded apices. The girdle occupies about 1/2 of the pervalvar
axis, with suture marked by a notch. Valves are elliptical in outline, with slightly inflated surface. The cells
have an apical axis of 16-20 μm, united into straight, more or less long chains. Apertures between sibling cells
are wide, oval. The inner setae are long and thin, arising inside from the margins of the valves perpendicular
to the chain axis. The terminal setae are diverging in broad girdle view at a wide angle. Cell possesses a
single large chloroplast. The resting spores are with strongly arched in center and flatter at the margin valves
covered with long and thin spines at the central part and with long parallel spines at the margin of primary
valve.
Distribution:
Chaetoceros seiracanthus is a warm water species distributed from south temperate to tropical waters (Hasle
& Syvertsen, 1997; Guiry & Guiry, 2017).
Local occurrence:
Chaetoceros seiracanthus was found occasionally across Kuwait’s waters, however, the species could
easily be misidentified with Chaetoceros brevis/laciniosus complex during routine phytoplankton analysis.
Reliable identification is based on resting spores. The species is reported for the first time from Kuwait’s
waters.

MEDIOPHYCEAE
Plate 60. Chaetoceros pseudocurvisetus.

a, b, d – parts of curved chains in narrow girdle view; c, e, f – parts of curved chains in broad girdle view;
g – part of chain with formation of resting spores; h – acid cleaned sibling frustules in broad girdle view; i –
detailed sibling cells in broad girdle view; j, k – detail of chains showing the insertion of setae.
LM, BF illumination (a-g, i) and phase contrast (h); SEM (j, k); scale bars: a, b – 50 μm; c-g – 20 μm; h-k – 10
μm.
Chaetoceros socialis Lauder
Plate 61
Hustedt, 1930: Fig. 435; Cupp, 1943: Fig. 100; Crosby & Wood, 1958: Pl. 38: 57; Hendey, 1964: Pl. XV: 3; Priddle &
Fryxell, 1985: p. 52-53; Ricard, 1987: Figs. 484, 485, 534; Hasle & Syvertsen, 1997: Pl. 47; Jensen & Moestrup, 1998:
Figs. 207-215; Shevchenko et al., 2006: Figs. 118-123; Throndsen et al., 2007: p. 173; Sunesen et al., 2008: Fig. 13 A-H;
Hoppenrath et al., 2009: Fig. 35 k-m; Kooistra et al., 2010: Figs. 98-102; Doan-Nhu et al., 2014: Figs. 174-176.
Synonymized names:
Chaetoceros radians Schütt 1895
Chaetoceros socialis f. autumnalis Proshkina-Lavrenko 1953
Chaetoceros socialis f. vernalis Proshkina-Lavrenko 1953
Chaetoceros socialis f. radians (Schütt) Proshkina-Lavrenko 1963
Chaetoceros socialis var. radians (Schütt) Tsarenko 2009
Frustules are rectangular in girdle view, corners are not touching those of adjacent cells. Valves are broadly
elliptical to almost circular in outline. The cells are small, with an apical axis of 3-5 µm, united into short
curved chains often aggregated in irregular globose colonies embedded in mucilage. Apertures between
sibling cells are rather wide, elliptical. The setae are hair-like, arising from corners of the cell, with short
basal part. Typically, the cell possesses three short curved setae and one longer and straight seta directed
towards the center of the globose colony. The cells in the colony are connected by crossing long setae. Cell
possesses a single chloroplast.
Distribution:
Chaetoceros socialis is a neritic temperate species. It has been previously reported from Kuwait’s waters
Local occurrence:
Chaetoceros socialis occurred occasionally across Kuwait’s waters, being recorded from 8% of all collected
concentrations (1.3-1.6∙106 cells/l) recorded in the inshore waters around Bubiyan Island during the
winter season in January.
Harmful Effect:
Chaetoceros socialis has been cited as a harmful species, associated with the production of mucilage,
causing clogging of gills and anoxia in fishes and benthic organisms during its blooms (Hallegraeff et al.,
1995). Blooms of this species occasionally affected cultured fish in Chile (Clément & Lembeye, 1993).
This species has been included in the list of potentially harmful species for Kuwait’s marine environment
(Al-Yamani et al., 2012) due to its capacity to form high biomass blooms in Kuwait’s waters.
Chaetoceros subtilis var. abnormis Prosckina-Lavrenko

Plate 39: a, b
Aké-Castillo et al., 2004: Figs. 2b, 4; Sunesen et al., 2008: Fig. 14 A-N; Doan-Nhu et al., 2014: Figs. 150-152 (as
Chaetoceros abnormis).
Synonymized names:
Chaetoceros abnormis Proschkina-Lavrenko 1953
Chaetoceros apendiculatus Müller-Melchers 1953
Chaetoceros atlantidae Müller-Melchers 1953
Frustules are rectangular in girdle view. Valves are narrowly elliptical in outline, heteropolar. The anterior
valve is convex, and the posterior one is concave. The cells are with an apical axis of 17-22 µm, united into
straight short chains. Valves of the sibling cells fit one in the other in the chain without apertures. The inner
setae are thin, almost straight, and perpendicular to the chain axis or slightly directed towards the posterior
end of the chain. The posterior terminal valves possess a single terminal thicker seta with spirally arranged
spines, directed towards the posterior end of the chain, almost parallel to the chain axis. Cell possesses a
single plate-shaped chloroplast.

MEDIOPHYCEAE
Plate 61. Chaetoceros socialis.

a – globular colony; b-d – part of colony with numerous chains lying together; e – part of single chain; f – acid
cleaned frustules.
LM, BF illumination (a-e) and phase contrast (f); scale bars: a – 50 μm; b – 25 μm; c – 20 μm; d-f – 10 μm.
Distribution:
Chaetoceros subtilis var. abnormis is a neritic temperate to subtropical species scarcely reported in literature.
Local occurrence:
Chaetoceros subtilis var. abnormis occurred occasionally across Kuwait’s waters, being recorded from
5% of all collected samples. The occurrence of this species was restricted mainly to the coastal waters
around Bubiyan Island, where it was observed during the period from October to February. The higher
concentrations (0.4-1.6∙104 cells/l) were recorded during the winter season in 2004-2006. The species is
Chaetoceros tenuissimus Meunier

Plate 59: d
Hasle & Syvertsen, 1997: Pl. 47; Throndsen et al., 2007: p. 174; Hoppenrath et al., 2009: Fig. 36 a, b; Kooistra et al.,
2010: Figs. 103-106.
Frustules are square in girdle view. Valves are elliptical in outline, with flat surface. The cells are small, with
an apical axis of 4-6 µm, solitary. The setae are thin and straight, arising from the poles of the valve and
forming a 45° angle to both the apical and pervalvar axes. Cell possesses a single plate-shaped chloroplast.
Distribution:
Chaetoceros tenuissimus is a neritic brackish water to marine species distributed mainly in temperate areas.
Local occurrence:
Chaetoceros tenuissimus is of low occurrence in Kuwait’s waters, and was never observed in large numbers.
However, its abundance could be underestimated during routine phytoplankton surveys due to its small
size and fragility. The species is reported for the first time from Kuwait’s waters.
Chaetoceros teres Gran

Plate 62
Cupp, 1943: Fig. 72; Crosby & Wood, 1958: Fig. 58; Hendey, 1964: Pl. X: 3; Ricard, 1987: Fig. 454; Hasle & Syvertsen,
1997: Pl. 42; Jensen & Moestrup, 1998: Figs. 218, 219; Shevchenko et al., 2006: Fig. 134; Throndsen et al., 2007: p. 168;
Hoppenrath et al., 2009: Fig. 36 c-l; Doan-Nhu et al., 2014: Figs. 61-65.
Frustules are rectangular in girdle view. Valves are broadly elliptical in outline, with flat surface and low
mantle. The cells are with an apical axis of 31-35 µm, united into long straight chains. Apertures between
sibling cells are narrow, linear-elliptic. The setae are long, arising from the corners of the valves at right
angles to the chain axis and diverging in acute angles to the valvar plane. The terminal setae are in U-shape,
directed to the chain axis. Cell possesses numerous round chloroplasts. The resting spores are with unequal
valves. The primary valve is convex and smooth. The secondary valve is less convex, with a marginal ring of
long spines becoming filament-like distally.
Remarks:
Chaetoceros teres is closely related in gross morphology to Chaetoceros lauderi but differ by the resting spores
having no spine on vaulted primary valve (e.g., Hasle & Syvertsen, 1997).
Distribution:
Chaetoceros teres is a neritic species distributed in cold to temperate waters.
Local occurrence:
Chaetoceros teres rarely occurred in Kuwait’s waters, and was never observed in large numbers. However,
its abundance may be underestimated during routine phytoplankton examination because it might be
easily confused with Chaetoceros lauderi. The species is reported for the first time from Kuwait’s waters.

MEDIOPHYCEAE
Plate 62. Chaetoceros teres.
a, b – short complete chains with attached cells of Amphora; c-g – detailed parts of chains in girdle view
showing connection of sibling cells, inner setae, and chloroplasts; h – resting spore.
LM, BF illumination (a-g); SEM (h); scale bars: a-c – 50 μm; e, g – 25 μm; d, f – 20 μm; h – 5 μm.
Chaetoceros tortissimus Gran
Plate 63
Hustedt, 1930: Fig. 434; Cupp, 1943: Fig. 99; Hendey, 1964: Pl. XI: 2; Ricard, 1987: Fig. 483; Hasle & Syvertsen, 1997:
Pl. 45; Shevchenko et al., 2006: Figs. 135, 136; Hoppenrath et al., 2009: Fig. 36m; Doan-Nhu et al., 2014: Fig. 173.
Frustules are rounded-rectangular in girdle view, weakly siliceous. Valves are elliptical in outline, with
slightly convex surface. The cells are with an apical axis of 21-25 µm, united into straight long chains,
strongly twisted about the chain axis. Sibling valves are touching in the middle, but not at corners, therefore
apertures are apparent only at corners. The setae are long and thin, arising a little inside corners, about at
right angles to chain axis, oriented in all directions. Cell possesses a single large chloroplast.
Distribution:
Chaetoceros tortissimus is a neritic temperate species. It has been previously reported from Kuwait’s waters
Local occurrence:
Chaetoceros tortissimus occurred occasionally across Kuwait’s waters, being recorded from 29% of all
higher concentrations (4.1-9.2∙104 cells/l) recorded in the inshore waters in Kuwait Bay and around
Bubiyan Island during the period from February to November.
Chaetoceros vanheurckii Gran

Plate 54: h-k
Allen & Cupp, 1935: Fig. 65; Cupp, 1943: Fig. 77; Shevchenko et al., 2006: Figs. 137, 138.
Frustules are rectangular in girdle view. Valves are broadly elliptical in outline, with flat surface and low
mantle. The cells are with an apical axis of 26-30 µm, united into long straight chains. Apertures between
sibling cells are narrow, linear-elliptic. The setae are long, thick, arising from the corners of the valves,
diverging in a angle of 40-60° to the chain axis. Cell possesses two plate-shaped chloroplasts. The resting
spores are spiny with unequal valves. The primary valve is broadly convex, the secondary valve is with
central undulation.
Distribution:
Chaetoceros vanheurckii is a neritic temperate to tropical species known mainly from the Pacific, including
coastal waters of China, Japan, Korea, Taiwan, Russian Far East, Australia, and New Zealand (Guiry
& Guiry, 2017).
Local occurrence:
Chaetoceros vanheurckii rarely occurred in Kuwait’s waters, and was observed in low concentrations. The

MEDIOPHYCEAE
Plate 63. Chaetoceros tortissimus.

a-e – short complete chains in girdle view showing chain torsion, inner setae and chloroplasts.
Family Leptocylindraceae Lebour
Genus Leptocylindrus Cleve
The species belonging to the genus Leptocylindrus are characterized by long, cylindrical cells, united into
straight chains by whole valve surface. Numerous intercalary bands are difficultly resolved with LM in
weakly siliceous species, but distinct in coarse frustules as in Leptocylindrus mediterraneus. The species within
the genus are distinguished by size, frustule ornamentation, and number of chloroplasts (Cupp, 1943; Hasle
& Syvertsen, 1997).
Leptocylindrus danicus Cleve

Plate 64: a-f
Pavillard, 1925: Fig. 35; Hustedt, 1930: Figs. 318, 319; Allen & Cupp, 1935: Fig. 27; Cupp, 1943: Fig. 39; Crosby &
Wood, 1958: Pl. 39: 80; Hendey, 1964: Pl. V: 2; Ricard, 1987: Fig. 250; Throndsen et al., 2007: p. 146; Hoppenrath et
al., 2009: Fig. 29 q, r.
Cells are high cylindrical in girdle view, much longer than broad, and circular in valve view, closely united
in more or less long, straight, dense chains, 9-11 µm in diameter, 32-37 µm broad (pervalvar axis). Cell wall
is weakly silicified. Of the sibling cells in a chain, one valve is slightly concave, the other is convex. The
cytoplasm contains several to numerous small plate-shaped chloroplasts, distributed throughout the cell.
Distribution:
Leptocylindrus danicus is a neritic species with widespread distribution. This species has been previously
Local occurrence:
Leptocylindrus danicus occurred frequently across Kuwait’s waters, being recorded from 48.5% of all
higher concentrations (0.9-2.7∙105 cells/l) recorded in the inshore waters of Kuwait Bay.
Harmful Effect:
Leptocylindrus danicus is a nontoxic bloom-forming species. Blooms of this species may cause fish kill in
mariculture cages (Yan et al., 2002).
Leptocylindrus mediterraneus (Péragallo) Hasle

Plate 65
Pavillard, 1925: Fig. 34 (as Dactyliosolen mediterraneus); Hustedt, 1930: Fig. 317 (as Dactyliosolen mediterraneus); Cupp,
1943: Fig. 38 (as Dactyliosolen mediterraneus); Crosby & Wood, 1958: Pl. 39: 81 (as Dactyliosolen mediterraneus); Priddle
& Fryxell, 1985: p. 62-63; Throndsen et al., 2007: p. 146.
Synonymized names:
Lauderia mediterranea Péragallo 1888
Dactyliosolen mediterraneus (Péragallo) Péragallo 1892
Dactyliosolen mediterraneus var. tenuis Cleve 1897
Dactyliosolen tenuis (Cleve) Gran 1902
Dactyliosolen meleagris Karsten 1906
Cells are high cylindrical in girdle view, much longer than broad, and circular in valve view, closely united
in more or less long, straight, stiff chains, 15-18 µm in diameter, 30-33 µm broad (pervalvar axis). Cell wall
is coarsely structured and heavily silicified. Intercalary bands are numerous, with ends lying in a straight
pervalvar line, distinctly areolated. Living cells are nearly always are found with epiphytic flagellate, Solenicola
setigera Pavillard, attached on the chains in the girdle-band zone.

Distribution:
Leptocylindrus mediterraneus is a neritic, sporadically oceanic widespread species.
Local occurrence:
Leptocylindrus mediterraneus occurred occasionally across Kuwait’s waters, being recorded from 5.5%
of all collected samples. The species was present throughout the year in low to moderate abundance
with higher concentrations (0.3-1.9∙104 cells/l) recorded in the southern inshore waters. The species is
reported for the first time from Kuwait.
Leptocylindrus minimus Gran

Plate 64: g-l
Hustedt, 1930: Fig. 321; Hendey, 1964: Pl. V: 3; Throndsen et al., 2007: p. 146; Hoppenrath et al., 2009: Fig. 29 s, t.
Cells are small, rod-shaped, high cylindrical in girdle view, much longer than broad, and circular in valve
view, closely united in more or less long, straight or slightly undulated dense chains, 3-4 µm in diameter, 7-11
µm broad (pervalvar axis). Cell wall is weakly silicified. The cytoplasm contains one or two small, elongated,
centrally located chloroplasts.
Distribution:
MEDIOPHYCEAE
Leptocylindrus minimus is a neritic species with widespread distribution. It has been previously reported
Local occurrence:
Leptocylindrus minimus occurred occasionally across Kuwait’s waters, being recorded from 29% of all
higher concentrations (0.6-1.8∙105 cells/l) recorded in the inshore waters during the period from January
to July.
MEDIOPHYCEAE
Plate 65. Leptocylindrus mediterranius.

a, c-e – parts of chains in girdle view with attached epiphytic flagellate Solenicola setigera; b – 3-celled complete
chain showing distinct girdle bands and cells with cytoplasm content; f – acid cleaned valve; g – frustule in
girdle view showing the morphology of girdle bands.
LM, BF illumination (a, c-e) and phase contrast (b, f); SEM (g); scale bars: a-e – 20 μm; f, g – 5 μm.
Plate 64. Leptocylindrus spp.

a-f – Leptocylindrus danicus: a, b – parts of long chains in girdle view showing sibling cells connection and
chloroplasts; c-e – short complete chains; f – valve view; g-l – Leptocylindrus minimus: g, h, k, l – parts of
long chains in girdle view showing sibling cells connection and chloroplasts; i, j – complete 2-celled chains.
LM, BF illumination (a-e, g, h) and DIC optic (i-l); SEM (f); scale bars: a-e, g-l – 10 μm; f – 2 μm.
Order Hemiaulales Round et Crawford
Family Hemiaulaceae Heiberg
Genus Cerataulina Péragallo ex Schütt
The species belonging to the genus Cerataulina are characterized by cylindrical weakly silicified cells united in
twisted chains. Valves possess two blunt projections near their margin. Sibling cells are attached by means of
a fine, small, wing-like extension, which fits into the sibling valve. Girdle is composed of numerous annular
intercalary bands. The species within the genus are distinguished by structure of girdle and elevations with
ocellus projected on each polar in valve face with wing-like extensions (Cupp, 1943; Hasle & Syvertsen,
1997).
Cerataulina bicornis (Ehrenberg) Hasle

Plate 66
Hasle & Sims, 1985: Figs. 1-17; Hasle & Syvertsen, 1997: Pl. 32.
Synonymized names:
Syringidium bicorne Ehrenberg 1845
Syringidium daemon Greville 1866
Cerataulina compacta Ostenfeld 1901
Cerataulina daemon (Greville) Hasle 1980
Cells are high cylindrical in girdle view, much longer than broad, closely united in more or less long,
straight, twisted chains, 22-37 µm in diameter, 65-95 µm broad (pervalvar axis). Cell wall is weakly silicified.
Girdle is composed of numerous narrow collar-like intercalary bands, poorly discerned with LM in water
mounts. Valve is circular in outline, slightly arched, with two opposite marginal elevations ornamented with
conspicuous broad wing-like extensions. Wing-like extensions of the elevations are fitting into V-shaped
deep furrows on mantle of the sibling valve in chain. The apertures between the sibling cells in chains are
narrow and slit-like, and cells are twisted about 90° in the pervalvar axis. The cytoplasm contains numerous
rod-shaped chloroplasts distributed throughout the cell. Nucleus is centrally located. The resting spore is
heavily silicified and heterovalvate. The primary valve has two prominent elevations with long wing-shaped
spines at the ends. A single long tubular process is located between the two elevations, tapering slightly
towards its open end. The secondary valve tapers gently towards a single process ending in a short spine.
Distribution:
Cerataulina bicornis is a coastal warm water species. It has been previously reported from Kuwait’s waters
Local occurrence:
Cerataulina bicornis occurred rarely in Kuwait’s waters, being recorded from less than 2% of all collected
samples in low numbers. The resting spores of Cerataulina bicornis occurred occasionally in the bottom
sediments, mainly in Kuwait Bay and around Bubiyan Island.
Cerataulina dentata Hasle

Plate 67
Hasle & Syvertsen, 1997: Pl. 33; Sunesen & Sar, 2007b: Fig. 3.
Cells are high cylindrical in girdle view, much longer than broad, closely united in more or less long, straight,
twisted chains, 20-29 µm in diameter, 49-61 µm broad (pervalvar axis). Cell wall is weakly silicified. Girdle
is composed of numerous narrow collar-like intercalary bands, clearly discerned with LM in water mounts.
Valve is circular in outline, flat, with two short opposite marginal elevations ornamented with small wing-like
extensions. Cells are united into a chain by the opposition of two elevations and twisted about 90° around
the chain axis. The apertures between the sibling cells in chains are narrow and slit-like. The cytoplasm
contains numerous elongate plate-shaped chloroplasts distributed throughout the cell. Nucleus is centrally
located.

Distribution:
Cerataulina dentata is a coastal warm water species. It has been previously reported from Kuwait’s waters
Local occurrence:
Cerataulina dentata occurred occasionally across Kuwait’s waters, being recorded from 8% of all collected
samples. The species was present in low to moderate abundance with higher concentrations (0.4-4.2∙103
cells/l) recorded in the inshore waters around Bubiyan Island during the warm season from May to
October.
Cerataulina pelagica (Cleve) Hendey

Plate 68
Pavillard, 1925: Fig. 99 (as Cerataulina bergonii); Hustedt, 1930: Fig. 517 (as Cerataulina bergonii); Allen & Cupp, 1935:
Fig. 86 (as Cerataulina bergonii); Cupp, 1943: Fig. 117 (as Cerataulina bergonii); Crosby & Wood, 1958: Pl. 36: 18; Hendey,
1964: Pl. IV: 4; Simonsen, 1974: p. 33 (as Cerataulina bergonii); Ricard, 1987: Figs. 331-338; Throndsen et al., 2007: p.
177; Hoppenrath et al., 2009: Fig. 29 a-h.
Synonymized names:
Zygoceros pelagicum Cleve 1889
MEDIOPHYCEAE
Cerataulus bergonii Péragallo 1892
Cerataulina bergonii (Péragallo) Schütt 1896
Cerataulina bergonii var. elongata Schröder 1900
Cerataulina bergonii Ostenfeld 1903
Cells are high cylindrical in girdle view, much longer than broad, closely united in more or less long, straight
or slightly arched, twisted chains, 18-26 µm in diameter, 38-62 µm broad (pervalvar axis). Cell wall is weakly
silicified. Girdle is composed of numerous narrow collar-like intercalary bands, poorly discerned with LM in
water mounts. Valve is circular in outline, convex, with two short opposite marginal elevations ornamented
with small broadly triangular wing-like extensions. Wing-like extensions of the elevations are fitting into
complementary triangular grooved areas on the mantle of the sibling valve in chain. There is a single labiate
process (rimoportula), which is subcentrally located. The areolae are arranged radially, centered on the
rimoportula. The apertures between the sibling cells in chains are narrow and slit-like, and cells are twisted
about 90° in the pervalvar axis. The cytoplasm contains numerous plate-shaped chloroplasts distributed
throughout the cell. Nucleus is centrally located close to cell wall.
Distribution:
Cerataulina pelagica is a neritic globally distributed species. It has been previously reported from the
Arabian Gulf (Simonsen, 1974, as Cerataulina bergonii) and from Kuwait’s waters (Al-Yamani et al.,
Local occurrence:
Cerataulina pelagica occurred occasionally across Kuwait’s waters, being recorded from 28% of all
higher concentrations (3.1-5.2∙103 cells/l) recorded during the period from December to March.
Harmful Effect:
Cerataulina pelagica has been reported as a nontoxic bloom-forming species. Death of benthic shellfish
and bony fish was attributed to anoxia and clogging of the gills during a bloom of Cerataulina pelagica off
the coast of northeastern New Zealand in 1983 (Taylor et al., 1985).
Plate 66. Cerataulina bicornis.
a-c – 2-celled chains in girdle view; d, f – detailed sibling cells, arrowheads point to wing-like extensions; e –
single live cell showing paired marginal wing-like extensions (arrowheads); g – resting spore.
LM, BF illumination (a-e) and phase contrast (f, g); scale bars: b, c – 20 μm; a, d-g – 10 μm.

MEDIOPHYCEAE
Plate 67. Cerataulina dentata.

a – complete 5-celled chain in girdle view; b-d – parts of chains showing chloroplasts and short
marginal elevations (arrowheads); e – single cell in girdle view; f-h – detailed contiguous areas of
sibling cells with connected marginal elevations (arrowheads); i – collapsed part of chain showing
girdle bands and marginal elevations (arrowheads).
LM, BF illumination (h, i) and DIC optic (a-g); scale bars: a – 20 μm; b-i – 10 μm.
Genus Climacodium Grunow
Climacodium frauenfeldianum Grunow

Plate 69: a-d
Hustedt, 1930: Fig. 453; Allen & Cupp, 1935: Fig. 76; Cupp, 1943: Fig. 105; Crosby & Wood, 1958: Pl. 36: 30;
Simonsen, 1974: p. 33; Ricard, 1987: Fig. 341.
Cells are straight and flattened, weakly silicified, united into long ribbon-like chains. In girdle view cells are
I-shaped, with small linear middle part and more or less long, thin extensions on the poles of the apical axis,
80-90 μm along the apical axis and 14-16 µm broad (pervalvar axis) due to the absence of intercalary bands.
Valves are linear-elliptical, with extensions at the poles by which the cells are united into the chain. Valve
surface between the extensions is flat, and apertures between the sibling cells are almost right-angled or very
large oblong, usually wider than the cell. The cytoplasm contains numerous irregularly shaped chloroplasts
radiating from the centrally located nucleus.
Distribution:
Climacodium frauenfeldianum is an oceanic species with distribution from the tropical to temperate
latitudes of all oceans. It has been previously reported from the Arabian Gulf (Simonsen, 1974) and
MEDIOPHYCEAE
Local occurrence:
Climacodium frauenfeldianum occurred occasionally across Kuwait’s waters, being recorded from 7%
of all collected samples. The species was present throughout the year in low abundance with higher
concentrations (0.5-4.1∙103 cells/l) recorded in the southern waters.
Genus Hemiaulus Heiberg
The genus Hemiaulus encompasses marine, planktonic, typically chain-forming species. Cells are usually
rectangular in girdle view, bipolar. Valves are elliptical in outline, with two narrow, more or less long horn-
like elevations with claws or pointed tips at the ends of the apical axis, parallel to pervalvar axis. Apertures
between the sibling cells are usually almost rectangular. Within the genus, the species are distinguished by
morphology of horn-like elevations and frustule areolation (Cupp, 1943; Hendey, 1964; Hasle & Syvertsen,
1997).
Plate 68. Cerataulina pelagica.

a – complete 7-celled chain in girdle view; b – complete 4-celled chain in girdle view; c, d – parts of chains
showing chloroplasts and paired opposite marginal wing-like extensions (arrowheads); e – single cell in
girdle view with wing-like extensions (black arrowheads); f – detailed contiguous area of sibling cells with
connected wing-like extensions (white arrowhead); g, h – collapsed frustule in oblique view (g) and valve
view (h) showing subcentral rimoportula (white arrowhead) and two opposite marginal elevations with wing-
like extensions (black arrowheads); i – detailed valve surface showing rimoportula (white arrowhead) and
marginal elevation with wing-like extension (black arrowhead); j – detailed marginal elevation.
LM, BF illumination (a-d) and DIC optic (e, f); SEM (g-j); scale bars: a-c – 20 μm; d-h – 10 μm; i – 5 μm; j – 2
μm.
Hemiaulus hauckii Grunow
Plate 69: e-g
Pavillard, 1925: Fig. 100; Hustedt, 1930: Fig. 518; Cupp, 1943: Fig. 118; Crosby & Wood, 1958: Pl. 36: 23; Hendey,
1964: p. 106; Simonsen, 1974: p. 33; Lobban et al., 2012: Pl. 8: 4.
Synonymized names:
Hemiaulus delicatulus Lemmermann 1905
Cells are long, H-shaped in girdle view, with straight margins, 12-35 μm along the apical axis and 30-42
µm broad (pervalvar axis). Frustules are weakly siliceous, without visible ornamentation. Valves are linear-
elliptical in outline, with two narrow, long horn-like elevations ending in claws on the poles of the apical
axis, parallel to pervalvar axis. Cells are often united into more or less turned about the long axis chains by
horn-like elevations with clasped claws. Valve surface is concave, apertures between the sibling cells are wide
and rounded rectangular. The cytoplasm contains numerous small round to elongate chloroplasts. Nucleus
is centrally located.
Distribution:
Hemiaulus hauckii is a neritic temperate to warm water species. It has been previously reported from the
Indian Ocean (Simonsen, 1974).
Local occurrence:
Hemiaulus hauckii occurred rarely across Kuwait’s waters, being recorded from 2.3% of all collected
samples. The species was present throughout the year in low abundance with higher concentrations
(0.4-3.8∙103 cells/l) recorded in the southern inshore waters during the spring in March. The species is
Hemiaulus membranaceus Cleve

Plate 69: h-k
Allen & Cupp, 1935: Fig. 90; Cupp, 1943: Fig. 120; Crosby & Wood, 1958: Pl. 36: 25; Simonsen, 1974: p. 33; Ricard,
1987: Fig. 330.
Cells are H-shaped in girdle view, with straight margins, 30-97 μm along the apical axis and 45-60 µm
broad (pervalvar axis). Frustules are weakly siliceous, without visible ornamentation. Valves are elliptical in
outline, with two narrow, short horn-like elevations with sharp points at the ends of the apical axis, parallel
to pervalvar axis. Cells are solitary or united into chains by horn-like elevations. Valve surface is nearly flat
or slightly concave. Apertures between the sibling cells are narrow, nearly rectangular to broadly elliptical.
The cytoplasm contains numerous small round chloroplasts. Nucleus is centrally located.
Distribution:
Hemiaulus membranaceus is a tropical oceanic species widely distributed in warm water regions (Cupp,
1943; Hasle & Syvertsen, 1997). The species have been previously reported from the Arabian Gulf
Local occurrence:
Hemiaulus membranaceus occurred occasionally across Kuwait’s waters, being recorded from 21%
of all collected samples. The species was present throughout the year in low abundance with higher
concentrations (480-680 cells/l) recorded in the inshore waters during the period from March to
November.

MEDIOPHYCEAE
Plate 69. Climacodium frauenfeldianum and Hemiaulus spp.

a-d – Climacodium frauenfeldianum: a, c – parts of chains in girdle view; b – 2-celled chain; d – 3-celled
chain; e-g – Hemiaulus hauckii: e – part of chain in girdle view; f – detailed contiguous area of sibling cells;
g – acid cleaned valve with two long horn-like elevations; h-k – Hemiaulus membranaceae: h, i – 2-celled
chains in girdle view; j, k – single live cell in girdle view at different focal planes.
LM, BF illumination (a-e), DIC optic (f, h-k), and phase contrast (g); scale bars: a-d, h – 25 μm; e-g, i-k
– 10 μm.
Hemiaulus sinensis (chinensis) Greville
Plate 70
1958: Pl. 36: 24; Hendey, 1970: p. 119; Simonsen, 1974: p. 33; Ricard, 1987: Figs. 326-329.
Cells are H-shaped in girdle view, with straight margins, 15-36 μm along the apical axis and 38-55 µm broad
(pervalvar axis). Valves are broadly elliptical to nearly circular in outline, with high mantle, possessing two
narrow, short horn-like elevations ending in a strong claw on the poles of the apical axis, parallel to pervalvar
axis. Cells are usually united into curved chains by horn-like elevations with clasped claws. Valve surface is
nearly flat or slightly convex. Apertures between the sibling cells are nearly rectangular. Frustules are heavily
siliceous, coarsely areolated. Areolae are irregularly polygonal, 7-9 in 10 µm in the center of the valve, 11-13
in 10 µm on the base of the mantle, arranged in radial rows of a characteristic excentric pattern, in that the
center of the areolation does not coincide with the center of the valve, but lies on one of the mantle surfaces.
The cytoplasm contains numerous small plate-shaped chloroplasts. Nucleus is centrally located, close to the
cell wall.
Distribution:
Hemiaulus sinensis is a neritic, south temperate to subtropical species. It has been previously reported
from the Arabian Gulf (Simonsen, 1974), and from Kuwait’s shores (Hendey, 1970) and coastal waters
Local occurrence:
Hemiaulus sinensis occurred commonly across Kuwait’s waters, being recorded from 50% of all collected
concentrations (5.2-7.2∙104 cells/l) recorded in the inshore waters of Kuwait Bay in November and
December.
Subclass Cymatosirophycidae Round et Crawford

Order Cymatosirales Round et Crawford
Family Cymatosiraceae Hasle
Genus Cymatosira Grunow
Cymatosira lorenziana Grunow

Plate 71: a-c
Péragallo & Péragallo, 1897-1908: Pl. LXXXII: 24; Hendey, 1970: Pl. 6: 69; Foged, 1984: Pl. XXVIII: 1-3, 7; Hustedt,
1985: Fig. 648; Ricard, 1987: Figs. 379-381; Throndsen et al., 2007: p. 180; Witkowski et al., 2000: Pl. 11: 12-15; Hein
et al., 2008: Fig. 2: 4; Al-Yamani & Saburova, 2011: Pl. 20: e-g.
Cells are small, usually united in small packets, valve to valve, by means of a system of spines. Frustules are
rectangular in girdle view, with acutely rounded corners slightly incised beneath apices. Valves are rhombic-
lanceolate with protracted, acutely rounded apices, 26-32 µm long, 7-10 µm broad. Transapical striae are
composed of puncta, 8-12 in 10 µm. Raphe is absent, sternum (pseudoraphe) is not visible.
Distribution:
Cymatosira lorenziana is a marine littoral species, occasionally found in plankton. It has been previously
reported from Kuwait’s shores (Hendey, 1970; Al-Yamani & Saburova, 2011) .
Local occurrence:
Cymatosira lorenziana occurred rarely in Kuwait’s waters, and was observed in low numbers.

MEDIOPHYCEAE
Plate 70. Hemiaulus sinensis.

a, b – parts of arc-shaped chains in narrow girdle view; c, d – detailed sibling cells connected by short horn-
like elevations (arrowheads); e – part of chain in broad girdle view; f, g – single cell in broad (f) and narrow
(g) girdle view; h, i – acid cleaned valves.
LM, BF illumination (a, b, e-g), DIC optic (c, d), and phase contrast (h, i); scale bars: a, b – 25 μm; c-i – 10 μm.
Subclass Thalassiosirophycidae Round et Crawford
Order Eupodiscales Nikolaev et Harwood
Family Eupodiscaceae Ralfs
Genus Odontella Agardh
The species belonging to the genus Odontella are characterized by bipolar elliptical or lanceolate valves with
horn-like elevation at each pole and usually two or more labiate processes (rimoportulae) usually with long
external tubes. Horn-like elevations are ending in a top plate perforated by closely packed holes (ocellus)
through which mucus is extruded. Cells are solitary or often form either straight chain when sibling cells are
united by two horns or zigzag chains when sibling cells are united by one horn (Hasle & Syvertsen, 1997;
Throndsen et al., 2007).
Odontella aurita (Lyngbye) Agardh

Plates 71: d-g; 72
Hustedt, 1930: Fig. 501 (as Biddulphia aurita); Cupp, 1943: Fig. 112 (as Biddulphia aurita); Crosby & Wood, 1958: Pl. 33:
33 (as Biddulphia aurita); Hendey, 1964: Pl. XXIV: 6 (as Biddulphia aurita); Hendey, 1970: p. 117 (as Biddulphia aurita);
Simonsen, 1974: p. 27; Foged, 1984: Pl. XIX: 4, 5 (as Biddulphia aurita); Witkowski et al., 2000: Pls. 8: 12, 13; 9: 1-3;
Throndsen et al., 2007: p. 176; Hein et al., 2008: Fig. 4: 7; Al-Kandari et al., 2009: Pls. 16: D-I, 17: A, B; Hoppenrath
et al., 2009: Fig. 39 a-l; Al-Yamani & Saburova, 2011: Pl. 12: a-g; Lobban et al., 2012: Pl. 6: 1, 2.
Synonymized names:
Diatoma auritum Lyngbye 1819
Biddulphia aurita (Lyngbye) Brébisson 1838
Denticella gracilis Ehrenberg 1841
Denticella aurita Ehrenberg 1854
Biddulphia aurita var. minuscule, var. minima Grunow 1881
Biddulphia pumila Castracane 1886
Biddulphia sansibarica Schmidt 1888
Cells are solitary or united in either straight or zigzag chains by horn-like elevations. Frustules are rectangular
in girdle view with strongly convex in the middle margins and four cornuted elevations from each angle.
Girdle zone is sharply differentiated from the valve zone by a clear depression. Valves are bipolar, elliptical in
outline, inflated in the center, radially punctated, 30-48 μm long. Two apices of each valve are prominently
elevated above the valve surface. Central part of valve is convex, more or less flattened at the top from
which long curved external tubes of two rimoportulae are arising. Cell wall is strongly siliceous, areolated-
punctated. Girdle band is punctated. Puncta are arranged in pervalvar rows, 7-10 rows in 10 μm. On the
valve, areolae are arranged in radial rows, 8-10 in 10 μm. Numerous elongate to irregular in shape golden-
brown chloroplasts are arranged around cell wall. Nucleus is centrally located.
Distribution:
Odontella aurita is a widely distributed neritic and littoral species. It has been previously reported from
the Indian Ocean (Simonsen, 1974), and from Kuwait’s shores (Hendey, 1970; Al-Yamani & Saburova,
2011) and coastal waters (Al-Kandari et al., 2009).
Local occurrence:
Odontella aurita occurred frequently across Kuwait’s waters, being recorded from 40% of all collected
concentrations (1.1-2.5∙104 cells/l) recorded in the inshore waters around Bubiyan Island.
Plate 71. Cymatosira lorenziana and Odontella spp.

a-c – Cymatosira lorenziana: a, b – sibling frustules in girdle view; c – acid cleaned valve; d-g – Odontella aurita:
d – live cells in girdle view united corner to corner with pads of mucilage to zigzag chain; e – single cell in
girdle view; f, g – acid cleaned frustules in valve (f) and girdle (g) view; h-j – Odontella weissflogii: single cells
in girdle view.
LM, BF illumination (a, d, e, h-j) and phase contrast (b, c, f, g); scale bars: h-j – 20 μm; a-g – 10 μm.

MEDIOPHYCEAE
Plate 72. Odontella aurita.
a, b – live cells in broad girdle view united corner to corner with pads of mucilage to zigzag chain; c – single
cell in broad girdle view; d – two sibling cells in narrow girdle view; e – acid cleaned frustule in girdle view;
f – two acid cleaned valves, note the size difference; g – external valve view.
LM, BF illumination (a-d) and phase contrast (e, f); SEM (g); scale bars: a – 20 μm; b-g – 10 μm.

Odontella weissflogii (Grunow) Grunow
Plate 71: h-j
Priddle & Fryxell, 1985: p. 70; Scott & Thomas, 2005: Fig. 2.21.
Synonymized names:
Biddulphia weissflogii Grunow 1882
Biddulphia striata Karsten 1905
Zygoceros caballeroi Azpeitia 1911
Cells are weakly silicified, solitary or united in short straight 2-3-celled chains. Frustules are rectangular in
girdle view with slightly raised mammiform corners. Valves are bipolar, elliptical in outline, with flat surface
and two long curved outer processes of rimoportulae divergent in direction, 72-87 μm long. Numerous plate-
shaped chloroplasts are arranged around cell wall. Nucleus is centrally located.
Distribution:
Odontella weissflogii is scarcely reported from temperate to southern cold water regions (Hasle & Syvertsen,
1997; Scott & Thomas, 2005)
Local occurrence:
Odontella weissflogii is of low occurrence in Kuwait’s waters, and was observed in low numbers. The
MEDIOPHYCEAE
Genus Trieres Ashworth et Theriot
The genus Trieres, with Trieres sinensis as the type species, was recently erected for some marine typical
planktonic species previously assigned to Odontella (Ashworth et al., 2013). The species belonging to this
genus are characterized by elliptical valves delimited by a ridge, with slender polar elevations and internally
sessile labiate processes (rimoportulae) with external tubes longer than the elevations (Ashworth et al., 2013).
Trieres mobiliensis (Bailey) Ashworth et Theriot

Plate 73
Pavillard, 1925: Fig. 96 (as Biddulphia mobiliensis); Hustedt, 1930: Fig. 495 (as Biddulphia mobiliensis); Allen & Cupp,
1935: Fig. 80 (as Biddulphia mobiliensis); Cupp, 1943: Fig. 110 (as Biddulphia mobiliensis); Crosby & Wood, 1958: Pl. 36:
19 (as Biddulphia mobiliensis); Hendey, 1964: Pl. XX: 3 (as Biddulphia mobiliensis); Hendey, 1970: p. 116 (as Biddulphia
mobiliensis); Simonsen, 1974: p. 27 (as Odontella mobiliensis); Ricard, 1987: Figs. 432, 433 (as Odontella mobiliensis);
Throndsen et al., 2007: p. 176 (as Odontella mobiliensis); Hoppenrath et al., 2009: Fig. 40 k-m (as Odontella mobiliensis);
Lavigne et al., 2015: Figs. 66-84.
Synonymized names:
Zygoceros mobiliensis Bailey 1851
Biddulphia mobiliensis (Bailey) Grunow ex Van Heurck 1882
Odontella mobiliensis (Bailey) Grunow 1884
Denticella mobiliensis (Bailey) Grunow 1884
Cells are weakly silicified, solitary or united in short straight chains by the elevations. Frustules are roundly
dodecagonal in girdle view, with slender, divergent, polar elevations. Valves are elliptical-lanceolate in
outline, 43-65 μm along the apical axis, convex, with a flat or nearly flat central part and deep, oblique
mantle. Polar elevations are arising inside the margin of the valve, directed diagonally outward, each bearing
a terminal ocellus. Two rimoportulae with long external tubes are placed far apart, but about equally far
from the processes, diagonally oriented with respect to the transapical axis, directed obliquely outward,
straight or often bent abruptly in their outer part, about twice as long as polar elevations. Frustule is finely
areolated. Areolae are small, hexagonal, 14-16 in 10 µm on valve and valve mantle, 17-18 on girdle band.
The cytoplasm contains numerous plate-shaped chloroplasts. Nucleus is centrally located.
Distribution:
Trieres mobiliensis is a neritic, truly planktonic, temperate to south temperate species. It has been previously
reported from the Arabian Gulf (Simonsen, 1974), and from Kuwait’s shores (Hendey, 1970) and coastal
Local occurrence:
Trieres mobiliensis occurred occasionally across Kuwait’s waters, being recorded from 7% of all collected
concentrations (0.6-1.3∙103 cells/l) recorded in the inshore waters during the warm season from May to
September.
Trieres sinensis (chinensis) (Greville) Ashworth et Theriot

Plate 74
Hustedt, 1930: Fig. 493 (as Biddulphia sinensis); Allen & Cupp, 1935: Fig. 81 (as Biddulphia sinensis); Hendey, 1964: Pl.
XX: 1 (as Biddulphia sinensis); Simonsen, 1974: p. 27 (as Odontella sinensis); Throndsen et al., 2007: p. 176 (as Odontella
sinensis); Hoppenrath et al., 2009: Fig. 41 g-k (as Odontella sinensis); Lavigne et al., 2015: Figs. 85-98.
Synonymized names:
Zygoceros sinensis (Greville) Cleve 1901
MEDIOPHYCEAE
Biddulphia sinensis Greville 1866
Odontella sinensis (Greville) Grunow 1884
Cells are large, weakly silicified, solitary or united into straight chains by the elevations. Frustules are nearly
rectangular in girdle view, with short, slender, barely divergent polar elevations. Valves are elliptical-lanceolate
in outline, 80-150 μm along the apical axis, with a nearly flat central part and deep, slightly oblique mantle.
One rimoportula with long external tube and apical spinule is placed at each pole just inside and very close
to the polar elevation, directed obliquely outward, straight or slightly bent in its outer part, about twice
as long as polar elevation or longer. Frustule is finely areolated. The cytoplasm contains numerous plate-
shaped chloroplasts scattered throughout the cell. Nucleus is centrally located.
Distribution:
Trieres sinensis is a truly planktonic species, widely spread in temperate and warm waters. It has been
previously reported from the Arabian Gulf (Simonsen, 1974) and from Kuwait’s waters (Al-Yamani et
al., 2004; Al-Kandari et al., 2009).
Local occurrence:
Trieres sinensis occurred occasionally across Kuwait’s waters, being recorded from 29% of all collected
concentrations (1.1-2.2∙103 cells/l) recorded in the inshore waters of Kuwait Bay during the fall season
from September to November.
Plate 73. Trieres mobiliensis.

a-c – live cells united to short 2-celled chains in broad girdle view; d, e – single cells in girdle view; f, g – acid
cleaned frustules in girdle view; h – complete frustule in girdle view; i – valve view showing paired polar
elevations and two rimoportulae with long external tubes; j – detailed polar elevation.
LM, BF illumination (a-d) and DIC optic (e-g); SEM (h-j); scale bars: a-c, h – 20 μm; d-g, i – 10 μm; j – 2 μm.
Plate 74. Trieres sinensis.
a – live cells united to 6-celled chain in broad girdle view; b – single cell in broad girdle view; c-e – detailed
sibling cells at different focal planes; f, g – detailed short polar elevation and closely located rimoportula with
long external tube and apical spinule; h - detailed contiguous area of sibling cells.
LM, BF illumination; scale bars: a – 100 μm; b-d – 50 μm; e – 25 μm; f-h – 10 μm.

Order Lithodesmiales Round et Crawford
Family Lithodesmiaceae Round
Genus Ditylum Bailey ex Bailey
The genus Ditylum encompasses marine planktonic species with characteristically triangular (seldom four
cornered) valves with a strong central siliceous hollow spine, and a marginal ridge, strengthened by ribs.
Girdle is composed of more or less numerous intercalary bands. Outer valve margin more or less strongly
undulated, giving the appearance of lines running from valve to valve (Cupp, 1943; Hasle & Syvertsen,
1997).
Ditylim brightwellii (West) Grunow ex Van Heurck

Plate 75
Pavillard, 1925: Fig. 91; Hustedt, 1930: Figs. 457-459; Cupp, 1943: Fig. 107-A, 107-B; Crosby & Wood, 1958: Pl. 36:
27; Hendey, 1964: Pl. V: 1; Ricard, 1987: Figs. 552-554; Throndsen et al., 2007: p. 181; Hoppenrath et al., 2009: Figs.
8, 38 j-s.
Synonymized names:
Triceratium brightwellii West 1860
Ditylum trigonum Bailey ex Bailey 1862
Ditylum inaequale Bailey ex Bailey 1862
MEDIOPHYCEAE
Cells are solitary or paired immediately after division, prism-shaped to cylindrical, much longer than broad,
weakly silicified, 28-45 µm in diameter and 85-103 µm long (pervalvar axis). Frustules are rectangular in
girdle view. Girdle is composed of numerous intercalary bands barely visible with LM. Valves are triangular
with more or less rounded corners, undulate, with fimbriate marginal ridge furnished with a corona of short,
but stout spines. The central area of valve is slightly raised, bearing a strong long siliceous hollow spine.
Valve surface is coarsely furnished with large radially arranged areolae starting from a central hyaline area
around a central process. The outer part of the valve surface and mantle are finely areolated. Numerous
small round chloroplasts are grouping toward the cell center around the nucleus.
Distribution:
Ditylim brightwellii is a neritic, south temperate species. It has been previously reported from Kuwait’s
Local occurrence:
Ditylim brightwellii occurred occasionally across Kuwait’s waters, being recorded from 16% of all
higher concentrations (0.6-1.4∙104 cells/l) recorded in the inshore waters of Kuwait Bay during the
spring season from March to May.
Ditylim sol (Grunow ex Van Heurck) de Toni

Plate 76
Allen & Cupp, 1935: Fig. 79.
Cells are solitary or paired immediately after division, prism-shaped, usually longer than broad, weakly
silicified, 73-95 µm in diameter and 150-200 µm long (pervalvar axis). Frustules are rectangular in girdle
view. Valves are triangular with undulated margin, giving the appearance of numerous longitudinal parallel
lines in girdle view. Each valve bears a strong long siliceous hollow spine. Numerous small plate-shaped
chloroplasts are distributed throughout the cytoplasm. Nucleus is centrally located.
Distribution:
Ditylim sol is a scarcely reported warm water species (Allen & Cupp, 1935; Hasle & Syvertsen, 1997).
Local occurrence:
Ditylim sol is of low occurrence in Kuwait’s waters, being recorded from 3% of all collected samples. The
species was observed sporadically in low concentrations from May to November. Ditylim sol is reported
for the first time from Kuwait’s waters.
Plate 75. Ditylim brightwellii.
a – live cells united to 2-celled chains in girdle view; b – end of cell with prominent long hollow spine; c – acid
cleaned valve; d – external valve view; e, f – internal valve view.
LM, BF illumination (a, b) and phase contrast (c); SEM (d-f); scale bars: 20 μm.

MEDIOPHYCEAE
Plate 76. Ditylim sol.

a, b – live dividing cells in girdle view; c – detailed dividing cell; d – girdle in high focus; e, f – single cells in
girdle view; g – cells united to 2-celled chains in girdle view.
LM, BF illumination; scale bars: a, b, e-g – 50 μm; d – 25 μm; c – 20 μm.
Order Stephanodiscales Nikolaev et Harwood
Family Stephanodiscaceae Makarova
Genus Cyclotella (Kützing) Brébisson
The genus Cyclotella is primarily freshwater, and contains just a few brackish water and marine species. The
species of this genus are characterized by circular valves with different ornamentation between central and
marginal areas of the valve face, and a ring of strutted processes (fultoportulae) interrupted by a single labiate
process (rimoportula) on the valve mantle. Marginal area of valve is alveolate-striated, whereas central area
is devoid of striae but either reticulate-rugose or warty or granulated, bearing one to many fultoportulae
(Prasad et al., 1990; Hasle & Syvertsen, 1997).
Due to the morphological similarity among species and high intra-specific variation, taxonomic
discrimination of the Cyclotella species during routine phytoplankton analysis is hard and often unreliable.
Close examination of frustules morphology revealed the presence at least six Cyclotella species in the
composition of the phytoplankton of Kuwait’s waters. The Cyclotella species commonly occurred across
Kuwait’s waters, being recorded from 62% of all collected samples. This species complex was present all year
round, more abundantly (0.5-1.8∙106 cells/l) in the inshore waters of Kuwait Bay during the warm season
from June to November. Due to their capacity to form blooms, Cyclotella species have been included in the
list of potentially harmful species for Kuwait’s marine environment (Al-Yamani et al., 2012).
Cyclotella atomus Husted

Plate 77
Chung et al., 2010: Figs 2-21; Sar et al., 2010b: Figs 41-46; Cavalcante et al., 2013: Figs 4, 5; Park et al., 2013: Fig. 1;
Lee, 2015: Fig. 1.
Cells are very small, solitary or occur in short chain, drum-shaped in girdle view and circular in valve view,
3.1-5.7 μm in diameter. Valve face is flat or slightly undulated with smooth central area. A single valve face
fultoportula may be present subcentrally, opening externally by a rimmed pore and internally surrounded by
two satellite pores. The marginal area is striated with raised striae and depressed interstriae. The marginal
fultoportulae occur regularly on every third to sixth interstriae, with external opening as a rimmed pore
and internally as a tube with two satellite pores. A single marginal rimoportula present on the mantle on an
interstria between two marginal fultaportulae. The cytoplasm contains 2-3 small plate-shaped chloroplasts.
Remarks:
Cyclotella atomus var. marina (Pl. 77: g-i) is distinguished from the other variety C. atomus var. atomus (Pl.
77: d-f) by the absence of valve face fultoportula.
Distribution:
Cyclotella atomus is a cosmopolitan fresh- to marine neritic species (Sar et al., 2010b).
Local occurrence:
Cyclotella atomus is too small for accurate determination during routine phytoplankton examination. The
structure of the valve surface is indistinct under the light microscope. The minute size makes this species
easy to be overlooked. Both varieties of Cyclotella atomus appear occasionally in electron microscopy
preparations from Kuwait’s coastal waters. The species is recorded for the first time from Kuwait.

MEDIOPHYCEAE
Plate 77. Cyclotella atomus.

a – two live cells in valve (top) and girdle (bottom) view united by organic thread; b – live cell in valve
view; c – live cell in girdle view; d-f – Cyclotella atomus var. atomus: d – two acid cleaned frustules
in valve (top) and girdle (bottom) view showing a single valve face fultoportula (black arrowhead)
and marginal fultoportulae (white arrowheads); e – acid cleaned frustule in external valve view
showing a single valve face fultoportula (black arrowhead) and marginal fultoportulae (white
arrowheads); f – internal valve view showing a single valve face fultoportula (black arrowhead),
marginal fultoportulae (white arrowheads) and marginal rimoportula (arrow); g-i – Cyclotella atomus
var. marina: g – acid cleaned frustule in oblique girdle view showing marginal fultoportulae (white
arrowheads); h, i – internal valve view showing marginal fultoportulae (white arrowheads).
LM, DIC optic (a-c); SEM (d-i); scale bars: a-c – 5 μm; d-f – 2 μm; g-i – 1 μm.
Cyclotella choctawhatcheeana Prasad
Plate 78
Prasad et al., 1990: Figs 2-26; Prasad & Nienow, 2006: Figs 47-55; Oliva et al., 2008: Figs 2, 3; Sar et al., 2010b: Figs
47-54.
Synonymized names:
Cyclotella hakanssoniae Wendker 1991
Cells are small, solitary, drum-shaped in girdle view and circular in valve view, 6.4-11.1 μm in diameter.
Valve face is undulated, with circular calliculate central area and striated marginal area restricted to about
one third of the valve diameter. The marginal striae density is 26-29 in 10 μm. The central area bears 1-3
fultoportulae on the internally concave portion of the valve, with 3 satellite pores internally. The marginal
fultoportulae occur regularly on every third to sixth interstriae, opening externally by a rimmed pore and
internally surrounded by two satellite pores. A single marginal rimoportula is present on the mantle on
an interstria between two fultaportulae, internally with radially oriented lip. A ring of stubby papilla-like
granules is irregularly arranged near the valve margin.The cytoplasm contains numerous small plate-shaped
chloroplasts. Nucleus is centrally located. Delicate organic threads radiating from around the edge of the
valve are often observed.
Distribution:
Cyclotella choctawhatcheeana is a marine neritic widely distributed species (Guiry & Guiry, 2017).
Local occurrence:
The occurrence of Cyclotella choctawhatcheeana in Kuwait’s waters could not be defined due to possible
misidentification with other small Cyclotella species during routine phytoplankton analysis. The illustrated
specimens were isolated from Kuwait’s southern inshore waters during spring 2008. The species is
recorded for the first time from Kuwait.
Cyclotella litoralis/cubiculata complex:

Plates 79, 80
Remarks:
Close examination of Kuwait’s material with emphasis on morphology of internal valve view with
electron microscopy revealed variability in morphology of marginal fultoportulae and rimoportula in
Cyclotella litoralis. Some Cyclotella specimens do not coincide with Cyclotella litoralis described by Lange
and Syvertsen (1989), but perfectly fit in morphology with those described by Sar, Sunesen and Lavigne
as Cyclotella cubiculata (Sar et al., 2010b). Examination with SEM, it is obvious that the external view of
Cyclotella cubiculata resembles Cyclotella litoralis and that both species frequently co-exist in the samples.
The Cyclotella litoralis/cubiculata species complex is included below.
Cyclotella litoralis Lange et Syvertsen

Plate 79
Hasle & Syvertsen, 1997: Pl. 1; Prasad & Nienow, 2006: Figs 35-39; Hoppenrath et al., 2009: Fig. 25 o, p; Sar et al.,
2010b: Figs 55-70; Park et al., 2013: Fig. 3.
Cells are solitary or occur in short chain, drum-shaped in girdle view and circular in valve view, 15-25 μm
in diameter. Valve face is strongly undulated, with circular calliculate central area and striated marginal
area restricted to a half of the valve surface. The marginal striae density is 12-16 in 10 μm. The central area
bears 3-6 fultoportulae arranged in semicircle irregular row on the internally concave portion of the valve,
surrounded internally by 3 satellite pores. The marginal fultoportulae occur regularly, mostly on every second
costa, less frequently on each third costa or in pairs, opening externally by rimmed pores and surrounded
by two satellite pores internally. A single marginal rimoportula is present on the mantle on one of the
costae between two fultoportulae, opening externally by slit-like pore with radially oriented lip internally.
Marginal chambers are absent. The external part of costae is covered with stubby papilla-like granules and
spines towards the mantle. The cytoplasm contains numerous small plate-shaped chloroplasts. Nucleus is
centrally located. Delicate organic threads radiating from around the edge of the valve are often observed.
Auxospores are formed as large bladders by separation of the valves and formation of larger cells in these.

MEDIOPHYCEAE
Plate 78. Cyclotella choctawhatcheeana.

a – live cell in valve view with radiating organic threads; b – cell in girdle view; c – acid cleaned frustule
in oblique valve view showing a single fultoportula (black arrowhead) in the central area and marginal
fultoportulae on every fifth costa (white arrowheads); d – internal valve view showing a single valve face
fultoportula (black arrowhead), marginal fultoportulae (white arrowheads) and marginal rimoportula
(arrow); e-g – acid cleaned frustules in valve view possessing one to three valve face fultoportulae (black
arrowheads), marginal rimoportulae are indicated by arrows.
LM, DIC optic (a, b); SEM (c-g); scale bars: a, b – 5 μm; c-g – 2 μm.
Distribution:
Cyclotella litoralis is a marine neritic species distributed primarily in temperate waters (Hasle & Syvertsen,
1997).
Cyclotella cubiculata Sar, Sunesen et Lavigne

Plate 80
Sar et al., 2010b: Figs 71-91.
Cells are solitary or occur in short chain, drum-shaped in girdle view and circular in valve view, 11-29 μm
in diameter. Valve face is strongly undulated, with circular calliculate central area and striated marginal
area restricted to a half of the valve surface. The marginal striae density is 13-16 in 10 μm. The central area
bears 2-6 fultoportulae arranged in semicircle irregular row on the internally concave portion of the valve,
surrounded internally by 3 satellite pores. The marginal fultoportulae occur regularly, mostly on every second
costa, less frequently on every first or third costa, opening externally by rimmed pores. A single marginal
rimoportula is present on the mantle on one of the costae between two fultoportulae slightly above the ring
of mantle fultoportulae. Externally, its aperture is slit-like rimmed pore, internally it is obliquely oriented
lip. Internally, valve bears marginal chambers separated by one or two coarse costae. Internal short tubes of
marginal fultoportulae are associated with every recessed costa and surrounded by two radial satellite pores.
The external part of costae is covered with stubby papilla-like granules and spines towards the mantle. The
cytoplasm contains numerous small plate-shaped chloroplasts. Nucleus is centrally located. Delicate organic
threads radiating from around the edge of the valve are often observed.
MEDIOPHYCEAE
Distribution:
Cyclotella cubiculata is a marine neritic species described from Argentinean coastal waters (Sar et al.,
2010b).
Local occurrence:
Cyclotella litoralis/cubiculata species complex was present throughout the year, more abundantly in the
inshore waters of Kuwait Bay during the warm season from June to November. The observations of
Cyclotella litoralis and Cyclotella cubiculata can be considered as new records for Kuwait.
Cyclotella stylorum Brightwell

Plate 81
Hustedt, 1930: Fig. 179; Hendey, 1970: Pl. 6: 68; Simonsen, 1974: p. 13; Foged, 1984: Pl. XVII: 1; Podzorski &
Håkansson, 1987: p. 20, Pl. 1: 1, 5, 6; Prasad et al., 1990: Fig. 30; Hasle & Syvertsen, 1997: Pl. 1; Al-Handal, 2009: p.
5; Al-Kandari et al., 2009: p. 44, Pl. 3: F-I; Al-Yamani & Saburova, 2011: Pl. 3 a-e.
Cells are solitary, rectangular in girdle view and circular in valve view, 29-56 μm in diameter. Valve face is
undulated, with large circular calliculate central area and striated marginal area, from about 1/3 of radius.
The marginal striae density is 10-13 in 10 μm. Valve edge has a ring of prominent marginal chambers
covering three alveolus openings. The central area bears 8-11 fultoportulae arranged in semicircle irregular
row on the internally concave portion of the valve. Marginal fultoportulae occur irregularly, mostly grouped
Plate 79. Cyclotella litoralis/cubiculata complex.

a-l – Cyclotella litoralis: a – live cell in valve view with radiating organic threads (arrowheads); b – live cell
in valve view; c – dividing cell in girdle view; d – auxospore with attached valves; e – collapsed valve with
row of fultoportulae (arrowheads); f – acid cleaned frustule in valve view showing fultoportulae on the
internally concave portion of valve (arrowheads); g, h – frustules in valve view showing semicircular row
of fultoportulae on the internally concave portion of valve (black arrowheads), marginal fultoportulae on
every second (mostly) costa (white arrowheads), and marginal rimoportula (arrow); i – detailed section of
valve in Fig. h showing marginal fultoportulae (white arrowheads) and a single rimoportula (arrow) between
two fultoportulae; j – acid cleaned frustule in external valve view; k – internal valve view showing valve
face fultoportulae (black arrowheads), marginal fultoportulae (white arrowheads), and marginal rimoportula
(arrow); l – detailed section of valve in Fig. k.
LM, DIC optic (a-e) and phase contrast (f); SEM (g-i); scale bars: a-d – 10 μm; e-l – 5 μm.
MEDIOPHYCEAE
Plate 81. Cyclotella stylorum.
a, b, d, e – live cells in valve view at different focal planes; c – cell in girdle view; f, g – acid cleaned frustules in
valve view showing semicircular row of fultoportulae on the internally concave portion of valve (arrowheads);
h – frustule in valve view showing semicircular row of fultoportulae (black arrowheads) in the central area
and ring of marginal fultoportulae (white arrowheads) grouped in pairs or quadruples; i – detailed section
of valve in Fig. h showing fultoportulae on valve face (black arrowheads) and marginal fultoportulae (white
arrowheads).
LM, BF illumination (c), DIC optic (a, b, d, e), and phase contrast (f, g); SEM (h, i); scale bars: a-g – 10 μm;
h, i – 5 μm.
Plate 80. Cyclotella litoralis/cubiculata complex.

a – live cells united to short chain in girdle view; b – detailed connection of sibling cells by organic threads; c-l
– Cyclotella cubiculata: c – live cell in oblique valve view; d – cell in girdle view; e – cell in oblique valve view
with radiating organic threads; f – acid cleaned frustule in valve view; g – frustule in oblique external valve
view showing marginal fultoportulae on every second or third costa (arrowheads) and marginal rimoportula
(arrow); h – frustule in external valve view; i, l – frustules in girdle view; j, k – internal valve view showing
valve face fultoportulae (black arrowheads), marginal fultoportulae (white arrowheads), and marginal
rimoportula (arrow).
LM, DIC optic (a-e), and phase contrast (f); SEM (g-l); scale bars: a-f – 10 μm; g-l – 5 μm.
in pairs or sometimes in triplets or quadruples. A single marginal rimoportula is present on the mantle on one
of the costae between two fultoportulae. The cytoplasm contains numerous small plate-shaped chloroplasts.
Nucleus is centrally located. Delicate organic threads radiating from around the edge of the valve are often
observed.
Distribution:
Cyclotella stylorum is a benthic littoral species, mainly distributed in warm waters, allochthonous in the
plankton. It has been previously reported from the Arabian Gulf (Simonsen, 1974), and from Kuwait’s
shores (Hendey, 1970; Al-Yamani & Saburova, 2011) and coastal waters (Al-Kandari et al., 2009).
Local occurrence:
The occurrence of Cyclotella stylorum in Kuwait’s waters could not be defined due to possible
misidentification with other Cyclotella species during routine phytoplankton analysis.
Order Thalassiosirales Glezer et Makarova

Family Lauderiaceae (Schütt) Lemmermann
Genus Lauderia Cleve
Lauderia annulata Cleve

Plates 82, 83
Pavillard, 1925: Fig. 32; Hustedt, 1930: Fig. 313; Cupp, 1943: Fig. 35; Hendey, 1964: p. 143; Ricard, 1987: Fig. 147; Sar
et al., 2001: Figs. 62-65; Hoppenrath et al., 2009: Fig. 19 a-i.
Synonymized names:
Lauderia borealis Gran 1900
Cells occur in straight chains, united by mucilage threads from strutted processes (fultoportulae) on valve
face. Cells are cylindrical, rectangular with rounded apices in girdle view. Valves are circular in outline, flat
to slightly concave in the center, rounded at the margins, 38-55 μm in diameter. Numerous fultoportulae are
scattered over valve surface and arranged in a conspicuous marginal ring. There is a ring of long occluded
processes in marginal zone. A single large rimoportula is present in the valve margin. Valve surface is
ornamented with radial ribs. Girdle is composed of numerous narrow collar-like intercalary bands, barely
discerned with LM. Intercalary bands are delicately areolated. The cytoplasm contains numerous small
plate-shaped chloroplasts. Nucleus is centrally located, close to cell wall.
Distribution:
Lauderia annulata is a neritic temperate species. It has been previously reported from Kuwait’s waters
Local occurrence:
Lauderia annulata occurred commonly across Kuwait’s waters, being recorded from 58.5% of all collected
concentrations (0.8-2.6∙105 cells/l) recorded in the inshore waters from June to December.

MEDIOPHYCEAE
Plate 82. Lauderia annulata.

a, b, d, e – live cells united to long chains in girdle view; c – complete short 4-celled chain; f-i – detailed
contiguous area of sibling cells showing girdle bands, radiating delicate organic threads, and marginal
ring of fultoportulae.
LM, BF illumination; scale bars: a-e – 20 μm; f-i – 10 μm.
Plate 83. Lauderia annulata.
a – cell connected to another cell in girdle view; b, c – detailed contiguous area of sibling cells showing
long occluded processes (black arrowheads) and large rimoportula (arrow); d-f – frustules in valve view
showing fultoportulae (white arrowhead), occluded processes (black arrowhead), rimoportula (arrows), and
fine threads from fultoportulae; g – detail of valve interior showing rimoportula (arrow).
SEM; scale bars: a-f – 10 μm; g – 5 μm.

Family Skeletonemataceae Lebour
Genus Skeletonema Greville
The genus Skeletonema encompasses small-sized chain-forming species with nearly cylindrical cells with
a row of fine spines at the edge of the valve parallel to longitudinal pervalvar axis, which join midway
between sibling cells and unite cells into chains. Prior to the year 2005, Skeletonema costatum was regarded
as cosmopolitan species and one of the most commonplace diatoms in the marine coastal environment
(e.g., Hasle & Syvertsen, 1997; Throndsen et al., 2007). Recent fine structural morphological investigations
coupled with molecular analyses have revealed high taxonomic diversity within the genus Skeletonema,
and a range of distinct species have been recognized within the taxon thus far perceived as Skeletonema
costatum (Sarno et al., 2005, 2007; Zingone et al., 2005). The modern-day concept of the genus defines the
basic morphological characters distinguishing Skeletonema species, including the structure of the external
processes of the intercalary fultoportulae and way of their junction, the structure of the external process of
the terminal fultoportula; the morphology of the intercalary rimoportulae, the girdle bands structure, and
number of chloroplasts per cell (Sarno et al., 2005).
The species attributed to Skeletonema costatum has been previously reported from Kuwait’s waters (Al-
Kandari et al., 2009). In the present study, Kuwaiti material was reinvestigated in more details based on
LM and SEM examination, resulted in assignment of Kuwait’s specimens to Skeletonema grevillei Sarno et
Zingone, however, further molecular studies are required to confirm the identification.
Skeletonema grevillei Sarno et Zingone
MEDIOPHYCEAE
Plates 84, 85
Zingone et al., 2005: Figs. 5 A-f, 6 A-I; Sarno et al., 2007: Fig. 2.
Cells are small, lens-shaped to cylindrical with rounded ends in girdle view, united in more or less long
straight chains by a ring of fultoportulae with long external processes. Spaces between cells are usually
longer than cells themselves. Valves are circular in outline, with convex face, 6-8 μm in diameter. Valve
surface is areolated and furnished with small spinulae. Areolae are radiating from the valve center. Valve
bears a marginal ring of fultoportulae with long external processes open along their length. Each process
connects with one process of the sibling valve (1:1 junction), but occasionally, one process may be connected
with two processes of the sibling valve (1:2 junction). The bases of fultoportulae are linked to each other and
to valve mantle by distinct siliceous ridges. Internally, apertures of fultoportulae are small, round rimmed
openings with 3 satellite pores. Valve bears a single rimoportula. The intercalary rimoportula is marginal,
located between two fultoportulae, opens externally with a short tubule and internally as sessile obliquely
oriented lip. The terminal rimoportula lies close to valve margin, with external long, open tube with funnel-
shaped distal end, as long as the terminal fultoportulae. The cytoplasm contains a single irregularly shaped
chloroplast. Nucleus is centrally located.
Distribution:
Based on a limited data set, Skeletonema grevillei is a neritic species reported from the geographically
distant sites in Asia (Hong Kong, China) and the Arabian Sea (Muscat, Oman) (Zingone et al., 2005;
Sarno et al., 2007; Guiry & Guiry, 2017). The records of Skeletonema attributed to S. costatum from the
Arabian Gulf (Simonsen, 1974) and Kuwait’s waters (Al-Kandari et al., 2009) need to be verified.
Local occurrence:
Skeletonema grevillei occurred occasionally across Kuwait’s waters, being recorded from 6.4% of all
collected samples. The species was present throughout the year with higher concentrations (0.8-7.0∙106
cells/l) recorded in the inshore waters of Kuwait Bay from May to November.
Harmful Effect:
Skeletonema can be considered as a harmful nontoxic bloom-forming species. Dense blooms of these
small-sized chain-forming diatoms may lead to loss of appetite and change in behavior among farmed
fish. Blooms of these species produce basically harmless water discolorations, with the result that the
recreational value of the bloom area decreases due to low visibility of the water and eventually, under
exceptionally weather conditions in sheltered bays, the blooms can grow so dense that they cause escape
reactions and indiscriminate fish kills and kills of benthic invertebrates due to oxygen depletion, or
mechanically damage the gills of fish by clogging them (Andersen, 1996). In Kuwait’s waters, high
biomass bloom of Skeletonema grevillei was associated with local fish kill incident, which occurred along
Kuwait Bay shores in June of 2015 (for details see also Sect. 2, Vol. I).
Plate 84. Skeletonema grevillei.
a-e – live cells united to chains in girdle view; f – acid cleaned frustule showing common 1:1 junction between
sibling valves; g-i – sibling intercalary valves showing valve areolation, connection of intercalary fultoportulae
processes and siliceous ridges (arrows) joining the bases of fultoportulae processes.
LM, BF illumination (a-e) and phase contrast (f); SEM (g-i); scale bars: a-f – 5 μm; g-i – 2 μm.

MEDIOPHYCEAE
Plate 85. Skeletonema grevillei.

a –1:1 and 1:2 (arrowheads) junctions between intercalary fultoportulae of sibling valves; b, c
– intercalary valves in external view showing marginal ring of fultoportulae (arrowheads) and
marginal intercalary rimoportula (arrow); d – intercalary valve in internal view showing internal
apertures of fultoportulae (arrowheads) and rimoportula (arrow); e – detailed valve margin
in internal view showing apertures of fultoportulae (arrowheads) and rimoportula (arrow); f,
g – terminal valves in external view showing marginal rim of fultoportulae (arrowheads) and
marginal long tube-shaped terminal rimoportula (arrow);
SEM; scale bars: 2 μm.
Family Thalassiosiraceae Lebour
Genus Cymatotheca Hendey
Cymatotheca minima Voigt

Plate 86
Voigt, 1960: Pl. 1: 6-9; Tynni, 1983: Pl. V: 40, 41 (as Hemidiscus weissflogi var.); Ricard, 1987: Fig. 151; Estrada-Gutiérrez
et al., 2017: Fig. 8.
Cells are small, solitary. Valves vary from subcircular to elliptical in outline, 10.4-11.5 μm long and 8.7-9.8
μm wide. Valve face is undulated along the short axis, divided into two halves lying in different focal planes.
The depressed sector is smaller than raised one and surrounded by a wide semicircular hyaline border along
the valve margin. Valve surface is furnished with small round areolae. Areolae are arranged in irregular radial
rows, decreasing in size towards the margin, 19-24 in 10 μm in the center of valve and 29-36 in 10 μm near
the margin. Valve mantle is irregularly punctate. A single valve face fultoportula is located off center, always
at the raised sector, externally appears as a simple pore, slightly larger than adjacent areolae, internally as
a short tube surrounded with two satellite pores. Six marginal fultoportulae are arranged in the ring along
the junction between the valve face and mantle, with external openings as raised pores, and internally as
short tubes with 3 satellite pores. Fultoportulae are more densely distributed along the depressed part of
valve (four of six). A single rimoportula is located midway between two marginal fultoportulae, next to the
depressed area in front of the valve face fultoportula. It opens externally by raised pore, internally is sessile
and tangentially oriented. The cingulum is composed of four open bands. The valvocopula is areolated with
one row of larger pores along the advalvar edge, 63-65 in 10 μm, and with three rows of finer pores toward
the abvalvar side (75-78 in 10 μm). The next bands are also finely areolated.
Distribution:
Cymatotheca minima is a marine warm water benthic species, allochthonous in the plankton samples
(Voight, 1960) with scarcely known geographic distribution.
Local occurrence:
Cymatotheca minima can be easily overlooked on account of its small size during phytoplankton
examination and appears occasionally in electron microscopy preparations from Kuwait’s coastal
waters. The photographed specimens were isolated from Kuwait’s northern waters in November 2017.
This species is recorded for the first time from Kuwait.
Plate 86. Cymatotheca minima

a-c – acid cleaned frustules in valve view showing variability in shape from circular (a, b) to elliptical (c); d
– external valve view showing valve face fultoportula (white arrowhead) and marginal fultoportulae (black
arrowheads); e – internal valve view showing valve face fultoportula (white arrowhead), ring of marginal
fultoportulae (black arrowheads), and marginal rimoportula (arrow); f-h – detailed valve sectors in external (f,
g) and internal (h) view with valve face fultoportula (white arrowhead), ring of marginal fultoportulae (black
arrowheads), marginal rimoportula (arrow), and cingulum (g); i, j – valve face fultoportula (arrowhead) in
external (i) and internal (j) view; k – detail of collapsed frustule showing cingulum.
SEM; scale bars: a-g – 2 μm; h-k – 1 μm.

MEDIOPHYCEAE
Genus Detonula Schütt ex De Toni
Detonula pumila (Castracane) Gran

Plate 87
Pavillard, 1925: Fig. 33 (as Schroederella delicatula); Hustedt, 1930: Fig. 314 (as Schroederella delicatula); Allen & Cupp,
1935: Fig. 26 (as Schroederella delicatula); Cupp, 1943: Fig. 36 (as Schroederella delicatula); Crosby & Wood, 1958: Pl. 35:
15 (as Schroederella delicatula); Hendey, 1964: Pls. V: 4, VII: 6; Simonsen, 1974: Pl. 6: 3; Ricard, 1987: Fig. 133; Sar et
al., 2001: Figs. 60, 61; Throndsen et al., 2007: p. 128; Hoppenrath et al., 2009: Fig. 18 g-m.
Synonymized names:
Lauderia pumila Castracane 1886
Lauderia delicatula Péragallo 1888
Detonula delicatula (Péragallo) Gran 1900
Lauderia schroderi Bergon 1902
Detonula schroderi Gran 1905
Schroederella delicatula (Péragallo) Pavillard 1913
Schroederella schroderi (Bergon) Pavillard 1925
Schroederella delicatula f. schroederi (Bergon) Sournia 1968
Cells are united in long straight chains by a mucilage thread running through the fultoportula at the center
of the valve. Frustules are cylindrical to drum-shaped in girdle view, usually shorter than broad. Valves are
circular in outline, more or less convex, sometimes almost flat, always with a depression in the middle, 19-
26 µm in diameter. Each valve bears a single central fultoportula and a ring of marginal fultoportulae with
short external tubes, 6-8 in 10 µm. Girdle is composed of numerous collar-like intercalary bands. Frustule is
delicately areolated, 18-20 areolae in 10 µm. The cytoplasm contains several small plate-shaped chloroplasts.
Distribution:
Detonula pumila is a cosmopolitan species, but most frequently encountered in warmer seas. It has been
previously reported from the Arabian Gulf (Simonsen, 1974) and from Kuwait’s waters (Al-Kandari et
al., 2009).
Local occurrence:
Detonula pumila occurred occasionally across Kuwait’s waters, being recorded from 7% of all collected
samples. The species wass present throughout the year in low to moderate abundance with higher
concentrations (0.2-3.0∙104 cells/l) recorded in the inshore waters in Kuwait Bay and around Bubiyan
Island during the warm season from April to September.
Genus Planktoniella Schütt
Planktoniella sol (Wallich) Schütt

Plate 88
1958: Pl. 32: 18; Hendey, 1964: p. 82; Simonsen, 1974: p. 12; Ricard, 1987: Figs. 124-128; Throndsen et al., 2007: p.
130.
Synonymized names:
Coscinodiscus sol Wallich 1860
Planktoniella woltereckii Schimper 1905
Cells are solitary, disk-shaped, with a weakly silicified hyaline wing-like expansion all around consisting
of extracellular chambers strengthened by radial rays. The central disk is 45-70 µm in diameter, and entire
cell is 100-110 µm. Valves are circular in outline, nearly flat, with low mantle, coarsely areolated. Areolae
are large, hexagonal, arranged in radial to tangential rows. The central area is composed of a single central
areola with seven areolae grouped around it. The density of areolae is 5-7 in 10 µm at the valve center, 7-8

midway to margin, and 8-9 near margin. Chloroplasts are numerous, small, and disc-shaped. Nucleus is
centrally located.
Distribution:
Planktoniella sol is an oceanic species, widely distributed but most common in subtropical and tropical
waters (Cupp, 1943). It has been previously reported from the Arabian Gulf (Simonsen, 1974) and from
Kuwait’s waters (Al-Kandari et al., 2009).
Local occurrence:
Planktoniella sol is of low occurrence in Kuwait’s waters, being recorded from less that 1% of all collected
samples, and was observed in low abundance.
MEDIOPHYCEAE
Plate 87. Detonula pumila.

a – live cells united to long chain in girdle view; b – collapsed cells in chain; c, d – acid cleaned valves; e –
collapsed frustules in oblique valve view; f – detailed contiguous area of sibling cells showing marginal ring
of fultoportulae with short external tubes.
LM, BF illumination (a, b) and phase contrast (c, d); SEM (e, f); scale bars: a, b – 20 μm; c-e – 10 μm; f – 5 μm.
Plate 88. Planktoniella sol.
a-e, g – live cells in valve view; h, i – detailed cells in valve view showing valve surface areolation and hyaline
wing-like expansion; f – acid cleaned frustule in valve view; j – acid cleaned collapsed frustule in oblique valve
view with attached valve of Paralia sulcata.
LM, BF illumination (a-e, g-i) and phase contrast (f, j); scale bars: a-f – 25 μm; g-j – 10 μm.

Genus Porosira Jørgensen
Porosira pentaportula Syvertsen et Lange
Plates 89, 90
Hasle & Syvertsen, 1997: Pl. 3; Sar et al., 2001: Figs. 69-71; Lee et al., 2015: Fig. 8g.
Cells are solitary or often united in chains. Frustules are rectangular in girdle view, weakly silicified. Valves
are circular in outline, nearly flat or slightly convex, 27-41 μm in diameter. Valve surface is finely areolated
with small areolae arranged in radially branching rows, 33-45 in 10 μm at the center, much denser toward
the margin, up to 75-83 in 10 μm. Numerous fultoportulae are scattered over the valve face except in the
central area. The marginal fultoportulae are arranged in zig-zag pattern along the valve margin, 2-2.5 μm
apart. There is a single rimoportula with short external tube located close to valve margin and surrounded by
four small fultoportulae (with two ones on each side). The cytoplasm contains numerous small plate-shaped
chloroplasts. Nucleus is centrally located.
Distribution:
Porosira pentaportula is a marine species scarsely reported worldwide from temperate to tropical waters.
It was recorded from the Uruguayan and Brazilian coast, Norvegian waters, Rhode Island (Hasle &
Syvertsen, 1997), Yellow Sea (Lee et al., 2015), and Argentinean coastal waters (Sar et al., 2001).
Local occurrence:
Porosira pentaportula occurred occasionally in Kuwait’s waters, being recorded from 8% of all collected
MEDIOPHYCEAE
concentrations (2.0-3.9∙103 cells/l) recorded during the summer season from June to August along the
southern Kuwait’s shores. Porosira pentaportula is reported for the first time from Kuwait.
Genus Shionodiscus Alverson, Kang et Theriot

Shionodiscus oestrupii var. venrickiae (Fryxell et Hasle) Alverson, Kang et Theriot
Plate 91
Allen & Cupp, 1935: Fig. 4; Hustedt, 1930: Fig. 155; Hendey, 1964: p. 89 (all as Coscinosira oestrupii); Simonsen, 1974:
Pl. 1: 3-5 (as Thalassiosira oestrupii); Fryxell & Hasle, 1980: Figs. 11-19 (as Thalassiosira oestrupii var. venrickiae); Sar et
al., 2001: Figs. 45-47 (as Thalassiosira oestrupii var. venrickiae); Scott & Thomas, 2005: Fig. 2.56: a-d (as Thalassiosira
oestrupii); Tremarin et al., 2008: Figs. 22, 23 (as Thalassiosira oestrupii); García & Odebrecht, 2009b: Figs. 14, 47, 47 (as
Thalassiosira oestrupii var. venrickiae); Li et al., 2013: Figs. 101, 102 (as Thalassiosira oestrupii).
Synonymized names:
Thalassiosira oestrupii var. venrickiae Fryxell et Hasle 1980
Cells are solitary or united in short loose chains by thin mucilage thread running through the off-central
strutted process. Frustules are drum-shaped in girdle view, usually slightly shorter than broad. Valves are
circular in outline, nearly flat, 8.5-45 μm in diameter. Valve surface is coarsely areolated. Areolae are large,
hexagonal, closely packed, arranged in eccentric rows, 5-6 in 10 μm at the center, decrease in size towards
the margin. One fultoportula is present close to valve center, and one small rimoportula is located about
2-3 areolae away from the subcentral fultoportula, midway between the valve center and the margin. The
valve margin is furnished by a ring of fultoportulae, 3.5-6 μm apart. Externally, all portulae are without
tube extensions. Internally, subcentral fultoportula is surrounded by three satellite pores, and marginal
fultoportulae are with long internal tubes. Several plate-shaped chloroplasts are arranged around cell wall.
Nucleus is centrally located.
Distribution:
Shionodiscus oestrupii var. venrickae is a neritic warm water widely distributed species (Fryxell & Hasle,
1980). The species has been previously reported (as Thalassiosira oestrupii) from the Arabian Gulf
(Simonsen, 1974) and from Kuwait’s waters (Al-Kandari et al., 2009).
Local occurrence:
Shionodiscus oestrupii var. venrickae occasionally occurred in Kuwait’s waters, being recorded from 10% of
all collected samples. The species was present all year round with higher abundances (0.3-1.8∙106 cells/l)
recorded in the inshore waters of Kuwait Bay from June to November.
Plate 89. Porosira pentaportula.
a – live cells united to long 15-celled chain in girdle and oblique valve view; b – complete short 3-celled
chain in girdle view; c – two sibling cells; d – single cell in girdle view; e – complete short 5-celled chain in
girdle view; f – detailed valve surface showing areolation and chloroplasts; g – detailed valve margin with
ring of marginal fultoportulae (black arrowheads); h, i – acid cleaned valve (h) and its detailed part (i).
LM, BF illumination (a-e), DIC optic (f, g) and phase contrast (h, i); scale bars: a-c, e – 50 μm; d – 25
μm; f-i – 10 μm.

MEDIOPHYCEAE
Plate 90. Porosira pentaportula.

a – acid cleaned frustule in external valve view showing short tube of single rimoportula near valve
margin (arrow); b – collapsed valve, arrow points to marginal rimoportula; c, d, g – detail of valves
showing areolation, valve face fultoportulae (white arrowheads), ring of marginal fultoportulae
(black arrowheads) and single rimoportula (arrow); e – detail of margin showing rimoportula (arrow)
surrounded by two pairs of fultoportulae (black arrowheads); f – detailed central area.
SEM; scale bars: a, b – 10 μm; c, d – 5 μm; e-g – 2 μm.
Plate 91. Shionodiscus oestrupii var. venrickae.
a, b – live cells in valve view at different focal planes; c, d – acid cleaned frustules showing valve surface
areolation; e – detailed valve surface showing off-central fultoportula (arrowhead) and nearly located
rimoportula (arrow); f-h – frustules in external valve and oblique (g) view showing valve areolation, off-
central fultoportula (arrowhead), nearly located rimoportula (arrow), and marginal ring of fultoportulae
(black arrowheads); i – frustule in girdle view; j – internal valve view showing off-central fultoportula with
three satellite pores (arrowhead), nearly located rimoportula (arrow), and marginal ring of fultoportulae with
long internal tubes (black arrowheads); k – detail of valve in external view showing off-central fultoportula
(arrowhead), nearly located rimoportula (arrow), and marginal fultoportulae (black arrowheads).
LM, BF illumination (a, b) and phase contrast (c-e); SEM (f-k); scale bars: a-e – 10 μm; f-j – 5 μm; k – 2 μm.

Genus Thalassiosira Cleve
The diatom genus Thalassiosira has high species diversity worldwide, with more than 100 species described
(Round et al., 1990; Hasle & Syvertsen, 1997; Guiry & Guiry, 2017). Thalassiosira species are identified by
the presence of a marginal ring of fultoportulae, frequently with spine-like external development. The most
important characters for species recognition are the shape, position and number of strutted (fultoportulae)
and labiate (rimoportulae) processes on the valve face, the areolation pattern or other ornamentation of
valve, and the girdle band morphology (Fryxell & Hasle, 1977; Hasle & Syvertsen, 1997; Hasle, 2001). Gross
morphology including cells size, shape and chain formation are useful features for species identification
based on LM, whereas fine structural studies require the examination of acid cleaned frustules with light
and electron microscopy.
The diversity and abundance of Thalassiosira in Kuwait’s waters are greatly contributed by medium- and
small-sized species with weakly silicified frustules. In these delicate diatoms, the portulae pattern could
not be discerned from alive or Lugol-preserved material with LM. A wide range of Thalassiosira taxa were
recognized from Kuwait’s coastal waters based on their gross morphology, but not precisely identified during
routine phytoplankton analysis owing to challenges in the interpretation of fine frustule morphology and
required further LM examination of acid cleaned frustules mounted in a medium of a high refractive index
and electron microscopic studies.
Thalassiosira spp. occasionally occurred in Kuwait’s coastal waters. This species complex was present
in Kuwait’s inshore waters all year round in low to moderate concentrations, occasionally forming blooms
MEDIOPHYCEAE
during the warm season from April to September. Small-sized chain-forming Thalassiosira spp. can be
considered as potentially harmful nontoxic species due to their capacity to form intensive blooms in Kuwait’s
coastal waters. Dense blooms of these diatoms were associated with local fish mortality incidents along the
shores of Kuwait Bay in September 2014, June 2015, and April 2017 (for details see also Sect. 2, Vol. I).
Thalassiosira andamanica Gedde

Plate 92
Gedde, 1999: Figs. 2-16; Li et al., 2013: Figs. 6-12; Park et al., 2016a: Figs. 5, S1-S5.
Cells are solitary or united in short loose chains by a thin mucilage thread running through the central
fultoportula. Connecting thread between the sibling cells in a chain is relatively long, about twice the length
of cell diameter or slightly less. Frustules are drum-shaped in girdle view, shorter than broad. Valves are
circular in outline, slightly convex, 17-31 μm in diameter. Valve surface is finely areolated. Areolae are small,
hexagonal, closely packed, arranged in fasciculate pattern, 36-44 in 10 μm. The central fultoportula opens
externally by a simple pore with slightly raised rim, often next to large areola, and surrounded with 4 satellite
pores internally. Valve face fultoportulae are arranged distantly in 2 irregular rings. Marginal fultoportulae
are arranged in regular ring, 3.5-5 in 10 μm. Both the marginal and valve face fultoportulae open externally
by short tubes with distinct tulip-shaped ends. A single rimoportula with very long external tube is present
within the outermost ring of valve face fultoportulae, with radially oriented lip internally. The outmost valve
margin is furnished with ribs, 26-31 in 10 μm. Numerous plate-shaped chloroplasts are arranged alongside
the cell wall. Nucleus is centrally located.
Distribution:
Thalassiosira andamanica is a marine, neritic, principally warm water species. It has been reported from
the Indo-Pacific area in Thailand (Gedde, 1999), China (Li et al., 2013), and Korea (Park et al., 2016a).
Local occurrence:
Thalassiosira andamanica occasionally occurred in Kuwait’s inshore waters, and was observed in low to
moderate abundance, however, the species may be misidentified with other medium-sized Thalassiosira
species during routine phytoplankton analysis. The most reliable distinctive charater (pattern of strutted
processes) is clearly discernible by SEM only. Thalassiosira andamanica is reported for the first time from
Kuwait.
Plate 92. Thalassiosira andamanica.
a – two live cells united to chain by organic thread (arrowhead) in girdle view; b, c – cell in girdle view in
different focal planes, arrows point to marginal rimoportulae with long tube-like extensions; d – single cells
in valve view with marginal rimoportula (arrow); e – detailed valve margin of acid cleaned frustule showing
short outer tubes of marginal and valve face fultoportulae with tulip-shaped ends; f – acid cleaned frustule in
oblique valve view, arrow points to long outer extension of rimoportula; g, h – external valve view showing
pattern of fultoportulae and long extension of a single rimoportula (arrow); i – internal valve view showing
pattern of fultoportulae and radially oriented lip of rimoportula (arrow); j – detailed valve surface showing
central fultoportula (arrowhead); k, l – detailed valve margin in external (k) and internal (l) view showing
valve face (white arrowheads) and marginal (black arrowheads) fultoportulae and rimoportula (arrow).
LM, BF illumination (a) and DIC optic (b-d); SEM (e-l); scale bars: a – 10 μm; b-d, f-i – 5 μm; e, j-l – 1 μm.

Thalassiosira cathariensis García
Plate 93
García & Dutra, 2016: Figs. 1-14; Lavigne et al., 2018: Fig. 1.
Cells are very small, solitary or often occur in agglomerates united by mucilage and covered with attached
detritus particles. Frustules are weakly silicified, drum-shaped in girdle view, shorter than broad. Valves
are circular in outline, almost flat, 4.1-6.3 (occasionally 7.9-8.6) μm in diameter. Valve margin is coarsely
structured than central area. Areolae are small, round to elliptical, arranged in radial rows, decrease in
size from the center to the margin, 39-47 in 10 μm in the central area and 52-58 in 10 μm near the margin.
The central fultoportula opens externally by a short tube and surrounded with 3 satellite pores internally.
Marginal fultoportulae are arranged in regular ring, 3.5-4 (occasionally 4.5-5.5) in 10 μm. Fultoportulae
open externally by short tubes with distinct basal hyaline siliceous border, internally with 3 satellite pores.
A single rimoportula is located close to one fultoportula of the marginal ring, with external tube similar in
diameter and length to fultoportulae extensions. The advalvar edge of valvocopula is ornamented with a ring
of round poroids, 76 in 10 μm. The cytoplasm contains a few plate-shaped chloroplasts.
Distribution:
Thalassiosira cathariensis is a marine, neritic, recently described species. To date, it has been reported from
southern Brazil and Argentinean coastal waters (García & Dutra, 2016; Lavigne et al., 2018).
Local occurrence:
MEDIOPHYCEAE
Thalassiosira cathariensis is too small and delicate for accurate determination during routine phytoplankton
examination and may be easily overlooked. The structure of the valve surface is indistinct in the light
microscope. The species appears occasionally in electron microscopy preparations from Kuwait’s coastal
waters. The photographed specimens were isolated from the inshore waters of Kuwait Bay in April
2017. Thalassiosira cathariensis is recorded for the first time from Kuwait.
Thalassiosira cf. concava Semina

Plate 94
Semina, 1991: Figs. 1-8; Semina, 2003: Pl. 41: 1-5.
Cells are solitary or united in short loose chains by a thin mucilage thread running through the central
strutted process. Connecting thread between the sibling cells in a chain is relatively long, about twice the
length of cell diameter or slightly less. Frustules are lenticular in girdle view, shorter than broad. Valves are
circular in outline, convex, 16-39 μm in diameter. Valve surface is finely areolated. Areolae are small, closely
packed, arranged in fasciculate rows, 27-36 in 10 μm. A single central fultoportula is opening externally by a
simple pore with slightly raised rim, with 4 satellite pores internally (occasionally two closely spaced central
fultoportulae were observed). Valve face fultoportulae are arranged distantly in 2 irregular rings, with very
short external tubes, internally surrounded by 4 satellite pores. Marginal fultoportulae are arranged in regular
ring, 3-5.5 in 10 μm. A single rimoportula is present slightly offset internally from the valve margin within
the outermost ring of valve face fultoportulae, with long external tube and radially oriented lip internally.
The outmost valve margin is furnished with ribs, 18-20 in 10 μm. Numerous plate-shaped chloroplasts are
arranged alongside the cell wall. Nucleus is centrally located.
Remarks:
Studied Kuwait’s specimens well coincided in gross morphology with those described by Semina
as Thalassiosira concava (Semina, 1991) and with Thalassiosira andamanica (Gedde, 1999). Kuwait’s
specimens resembles these two species in size, areolation, and general pattern of portulae including
a single central fultoportula without external process, 2 irregular rings of valve face fultoportulae, one
regular ring of marginal fultoportulae, and a single rimoportula with long external tube. Despite the
above similarities, Thalassiosira cf. concava from Kuwait’s waters differs from Thalassiosira andamanica in
absence of tulip-shaped ends in fultoportulae, and has denser areolation (27-36 areolae in 10 μm) than
frustules of Thalassiosira concava (20-25 areolae in 10 μm), with more spaced marginal fultoportulae (12-
16 areolae apart versus 7-9 in Thalassiosira concava as it is seen from illustrations in Semina, 2003) and in
slightly different areolation pattern of cingular bands. Kuwait’s specimens were tentatively assigned to
Thalassiosira concava based on the structure of external processes of fultoportulae without tulip-shaped
Plate 93. Thalassiosira cathariensis.
a – live cells embedded in mucilage with attached detritus particles; b – live cells in valve view; c – frustule in
valve view embedded in mucilage and attached detritus; d – acid cleaned frustule in oblique valve view showing
areolation and portula pattern with marginal rimoportula (arrow) close to one marginal fultoportula; e-g –
external valve view showing areolation, marginal ring of fultoportulae (black arrowheads) and rimoportula
(arrow); h – valve in internal view showing marginal rimoportula (arrow) within the ring of marginal
fultoportulae (black arrowheads).
LM, DIC optic (a, b); SEM (c-h); scale bars: a, b – 5 μm; c-h – 2 μm.

MEDIOPHYCEAE
Plate 94. Thalassiosira cf. concava.

a – two live cells united to chain by organic thread (arrowhead) in girdle view; b – single cells in valve view;
c – acid cleaned frustule in girdle view with long tube-like extension of rimoportula (arrow) visible; d, e –
frustules in valve view, arrow points to long outer extension of rimoportula; f, g – detailed valve in external
(f) and internal (g) view showing central fultoportula (double arrowhead), valve face (white arrowheads)
and marginal (black arrowheads) fultoportulae and rimoportula (arrow); h – detailed cingulum; i – internal
valve view, arrow points to rimoportula.
LM, BF illumination (a) and DIC optic (b); SEM (c-i); scale bars: a – 20 μm; b – 10 μm; c-i – 5 μm.
ends, however, further studies are required to clarify whether it is a distinct species or a morphotype of
Thalassiosira andamanica or preparation artifact.
Distribution:
Thalassiosira concava is a marine tropical species that was originally described from the equatorial Pacific
and later reported from the Gulf of California (Semina, 2003).
Local occurrence:
Thalassiosira cf. concava may be misidentified with Thalassiosira andamanica and other medium-sized
Thalassiosira species during routine phytoplankton analysis. The most reliable distinctive character
(morphology of external processes of fultoportulae) is clearly discernible by SEM only. Thalassiosira
cf. concava appears occasionally in electron microscopy preparations from Kuwait’s coastal waters.
The photographed specimens were isolated from Kuwait’s southern inshore waters in April-May 2008.
Thalassiosira cf. concava is reported for the first time from Kuwait.
Thalassiosira concaviuscula Makarova

Plate 95
Harris et al., 1995: Figs. 5, 23; Hoppenrath et al., 2007: Figs. 8-10; Park & Lee, 2010: Pl.1: 1-8; Park et al., 2016a: Fig. 8.
Cells are small, united in short chains by a thin mucilage thread running through the central fultoportula.
Connecting thread between the sibling cells in a chain is about the length of cell diameter or slightly less.
Cells are often embedded in mucilage and covered with attached detritus particles. Frustules are drum-
shaped in girdle view, shorter than broad. Valves are circular in outline, almost flat, 10-22 μm in diameter.
Valve areolae are small, arranged in fasciculate pattern, decrease in size from the center to the margin,
29-35 in 10 μm in the central area and 31-38 in 10 μm near the margin. The external foramina of areolae
possess radial siliceous threads. Valve face is covered with minute siliceous spines arising from the edge of
each areola. The central fultoportula is located in a slight concavity of the valve, adjacent to a large areola,
opens externally by a short tube and surrounded with 3 satellite pores internally. Marginal fultoportulae are
arranged in regular ring, 2.5-3.4 μm apart, 5-7 in 10 μm. Fultoportulae open externally by short tubes with
funnel-shaped distal ends because siliceous collars encompass the tips. A single rimoportula is located close
to one fultoportula of the marginal ring, with external tube similar in diameter and length to fultoportulae
extensions but opening in different direction than the tubes of fultoportulae. The cytoplasm contains a few
plate-shaped chloroplasts.
Distribution:
Thalassiosira concaviuscula is a marine species widely distributed from temperate to tropical waters (Harris
et al., 1995; Hoppenrath et al., 2007; Park et al., 2016a).
Local occurrence:
Thalassiosira concaviuscula occurred in Kuwait’s inshore waters throughout the year in low to moderate
abundance, occasionally forming blooms (1.2-37.9∙106 cells/l) during the warm season from April to
September.
Harmful Effect:
Thalassiosira concaviuscula can be considered as a potentially harmful nontoxic bloom-forming species.
In bloom conditions, this species is mainly responsible for the high phytoplankton biomass causing
episodic hypoxia in the surrounding waters often resulting in sublethal to lethal effects on marine biota.
The photographed specimens were isolated from the inshore waters of Kuwait Bay in April 2017 during
a bloom incident associated with a local fish kill (for details see Sect. 2, Vol. I).

MEDIOPHYCEAE
Plate 95. Thalassiosira concaviuscula.

a, b – live cells in girdle view united to chains by organic threads lying in mucus with attached
detritus particles; c – live cells in valve view; d – frustule in valve view embedded in mucilage
and attached detritus; e, f – acid cleaned frustules of different size in external valve view
showing areolation, central fultoportula near large areola and marginal rimoportula (arrow)
close to marginal fultoportula; g – detail of valve showing central fultoportula near large areola
(arrowhead); h – detailed valve margin showing valve spinulation and marginal fultoportulae
(arrowheads); i – valve in internal view showing marginal rimoportula (arrow) within the ring
of marginal fultoportulae (black arrowheads).
LM, DIC optic (a-c); SEM (d-i); scale bars: a, b – 10 μm; c, f – 5 μm; d, e, h, i – 2 μm; g – 1 μm.
Thalassiosira decipiens (Grunow) Jørgensen
Plate 96: a-i
Hernández-Becerril & Tapia Peña, 1995: Figs. 10-15; Muylaert & Sabbe, 1996: Figs. 9, 10; Aké-Castillo et al., 1999:
Figs. 2-4; Sar et al., 2001: Figs. 14-16; Hoppenrath et al., 2007: Figs. 16-18; García & Odebrecht, 2009b: Figs. 3, 5, 6,
28-30; Park et al., 2009: Pl. 1: 1-6; Li et al., 2014: Figs. 20, 21; Lee et al., 2015: Fig. 8n; Park et al., 2016a: Fig. 10.
Synonymized names:
Coscinodiscus eccentricus var. decipiens Grunow 1878
Thalassiosira gelatinosa Hensen 1887
Cells are small, united in short loose chains by a thin mucilage thread running through the central fultoportula.
Connecting thread between the sibling cells in a chain is relatively long, about twice the length of cell
diameter or slightly less. Frustules are drum-shaped in girdle view, shorter than broad. Valves are circular in
outline, almost flat, 5.9-12.7 μm in diameter. Valve areolae are arranged in radial rows, decrease in size from
the center to the margin, 15.5-20 in 10 μm in the central area and 30-32 in 10 μm near the margin. Areolae on
the mantle are much smaller in size and arranged tightly, up to 36 in 10 μm. The external areolae foramina
have radial silicious threads. The entire valve surface is covered by tiny granules. The central fultoportula
is opening externally by a simple pore with slightly raised rim near the central areola and surrounded by
three satellite pores internally. Marginal fultoportulae with long external tubes are densely arranged in
regular ring, 1.3-1.4 μm apart, and surrounded by four satellite pores internally. A single rimoportula with
prominent external tube is present midway between two marginal fultoportulae. The cytoplasm contains a
few plate-shaped chloroplasts.
Distribution:
Thalassiosira decipiens is a neritic species widely distributed from temperate to tropical waters mainly in
shallow, coastal and estuarine environments (e.g., Park et al., 2016a, Guiry & Guiry, 2017).
Local occurrence:
Thalassiosira decipiens was occasionally found in Kuwait’s inshore waters, however, the species may be
misidentified with other small-sized Thalassiosira species during routine phytoplankton examination and
reliable identification requires SEM observations. Thalassiosira decipiens is reported for the first time from
Kuwait.
Plate 96. Thalassiosira spp.

a-i – Thalassiosira decipiens: a – live cells united to chain by organic threads in girdle view; b, c – single cells in
girdle view with long outer extension of marginal rimoportula visible (arrow); d, e – frustules in valve view,
external tube of rimoportula is arrowed; f – frustule in girdle view showing ring of marginal fultoportulae and
single rimoportula with long external tube (arrow) located midway between two fultoportulae; g – external
valve view showing areolation, central fultoportula, ring of marginal fultoportulae and single marginal
rimoportula (arrow); h – detailed central area showing valve surface ornamentation and central fultoportula;
i – internal valve view showing central fultoportula (white arrowhead), marginal fultoportulae (black
arrowheads), and marginal rimoportula (arrow); j – Thalassiosira nanolineata: valve in oblique view showing
areolation, central fultoportula (white arrowhead), ring of marginal fultoportulae (black arrowheads), and
single marginal rimoportula (arrow).
LM, DIC optic (a-c), SEM (d-i); scale bars: a-c – 10 μm; d-g, i, j – 2 μm; h – 1 μm.

MEDIOPHYCEAE
Thalassiosira delicatula Ostenfeld
Plate 97
Hallegraeff, 1984: Fig. 24a, b; Harris et al., 1995: Figs. 13, 30; Hoppenrath et al., 2007: Figs. 19-21; Park et al., 2016a:
Figs. 11, S6-S11.
Synonymized names:
Thalassiosira chilensis Krasske 1941
Thalassiosira coronata Gaarder 1951
Cells are small, united in short loose chains by a thin mucilage thread running through the central
fultoportula. Connecting thread between the sibling cells in a chain is relatively long, about twice the length
of cell diameter or more. Frustules are drum-shaped in girdle view, shorter than broad. Valves are circular
in outline, almost flat and slightly depressed at the center, 9.3-21.2 μm in diameter. Valve areolae are small,
irregularly shaped, arranged in radial rows with fasciculate pattern in large specimens, decrease in size from
the center to the margin, 28-31 in 10 μm in the central area and 32-34 in 10 μm near the margin. The central
fultoportula is opening externally by a short tube next to the central large areola, surrounded internally by
4 satellite pores. Vale face fultoportulae are arranged distantly in one irregular ring, opening externally by
short tubes, surrounded internally by 2-3 satellite pores. Occasionally 2 rings of valve face fultoportulae were
observed in large specimens. There are two dense marginal rings of alternating fultoportulae, 3.7-4.6 in 10
μm, with short external tubes and 3 satellite pores internally. A single rimoportula is located within the inner
ring of marginal fultoportulae, with prominent external tube that is longer and twice the diameter of the
adjacent fultoportulae. Additionally, there is a marginal ring of occluded processes. The cytoplasm contains
a few plate-shaped chloroplasts.
Distribution:
Thalassiosira delicatula is a marine, mainly warm water species (e.g., Park et al., 2016a, Guiry & Guiry,
2017).
Local occurrence:
Thalassiosira delicatula was occasionally found in Kuwait’s inshore waters, however, the species may be
misidentified with other small-sized Thalassiosira species during routine phytoplankton analysis, and
reliable identification requires SEM observations. In electron microscopy preparations, the species was
well represented in the material collected from the inshore waters of Kuwait Bay during the spring of
2017. Thalassiosira delicatula is reported for the first time from Kuwait’s waters.
Plate 97. Thalassiosira delicatula.

a – live cells united to chain by organic thread (arrowhead) in girdle view; b – live cells in girdle (left) and
valve (right) view; c – acid cleaned frustule in valve view; d-f – external valve view showing areolation and
portula pattern with central fultoportula (double arrowhead), valve face fultoportulae (white arrowheads),
marginal fultoportulae (black arrowheads), marginal rimoportula (white arrow), and occluded processes
(black arrows); g – detailed valve of large specimen; h – internal valve view showing valve face fultoportulae
(white arrowheads), marginal fultoportulae (black arrowheads) and rimoportula (arrow).
LM, DIC optic (a, b); SEM (c-h); scale bars: a, b, g – 5 μm; c-f, h – 2 μm.

MEDIOPHYCEAE
Thalassiosira diporocyclus Hasle
Plate 98
Simonsen, 1974: Pl.1: 2; Hallegraeff, 1984: Fig. 3; Hoppenrath et al., 2007: Figs. 22-24; Fernandes & Frassão-Santos,
2011: Figs. 1-23; Li et al., 2013: Figs. 36-42; Li et al., 2014: Figs. 22-28; Park et al., 2016a: Fig. 12.
Cells are solitary or often occur in agglomerates united by abundant mucilage. Frustules are drum-shaped in
girdle view, shorter than broad. Valves are circular in outline, convex, 15-24 μm in diameter. Valve areolae
are small, arranged radially in fasciculate sectors, 34-38 in 10 μm. A single fultoportula is located slightly
eccentrically, externally is opening in a small pore, internally as a short tube extension surrounded by three
satellite pores. On the valve face, there are 7-12 fultoportulae arranged in more or less regular ring almost
midway between the valve center and the margin. One marginal ring of regularly arranged fultoportulae is
present. Externally, fultoportulae are opening in simple pores with raised edges, with short internal tubes
surrounded by 2-3 (on valve face) or always 3 (on valve margin) satellite pores. A single rimoportula is
located within the ring of marginal fultoportulae, opening externally in an elongate aperture. The cytoplasm
contains numerous plate-shaped chloroplasts. The nucleus is centrally located.
Distribution:
Thalassiosira diporocyclus is a neritic widely distributed species from temperate to tropical waters (e.g.,
Park et al., 2016a, Guiry & Guiry, 2017). The species has been reported from the African and the Indian
sides of the Arabian Sea and from the Arabian Gulf (Simonsen, 1974).
Local occurrence:
Thalassiosira diporocyclus was occasionally found in Kuwait’s inshore waters, however, the species may be
misidentified with other medium-sized Thalassiosira species during routine phytoplankton analysis, and
reliable identification requires SEM observations. Thalassiosira diporocyclus is reported for the first time
from Kuwait.
Thalassiosira eccentrica (Ehrenberg) Cleve

Plate 99
Pavillard, 1925: Fig. 12 (as Coscinodiscus excentricus); Hustedt, 1930: Fig. 201 (as Coscinodiscus excentricus); Allen & Cupp,
1935: Fig. 5 (as Coscinodiscus excentricus); Crosby & Wood, 1958: Pl. 31: 10 (as Coscinodiscus excentricus); Hendey, 1964:
Pl. XXIV: 7 (as Coscinodiscus eccentricus); Simonsen, 1974: Pl. 2: 1-3; Hallegraeff, 1984: Fig. 15; Hernández-Becerril &
Tapia Peña, 1995: Figs. 16-20; Muylaert & Sabbe, 1996: Figs. 11, 12; Sar et al., 2001: Figs. 17-22; Aké-Castillo et al.,
1999: Figs. 5, 6; Hoppenrath et al., 2007: Figs. 25, 26; Throndsen et al., 2007: p. 136; Tremarin et al., 2008: Fig. 12;
García & Odebrecht, 2009b: Figs. 4, 7, 31-37; Hoppenrath et al., 2009: Figs. 5, 22 a-j; Park et al., 2009: Pl. 2: 7-12; Al-
Yamani & Saburova, 2011: Pl. 1: a-g; Li et al., 2013: Figs. 47-52; Li et al., 2014: Figs. 29-32; Lee et al., 2015: Fig. 8 o,
p; Park et al., 2016a: Fig. 14.
Synonymized names:
Coscinodiscus eccentricus Ehrenberg 1840
Coscinodiscus kryophilus Grunow 1884
Thalassiosira kryophila (Grunow) Jørgensen 1905
Thalassiosira excentrica Karsten 1905
Thalassiosira excentrica f. velata Cleve-Euler 1942
Thalassiosira excentrica var. fasciculata Chernov 1947
Thalassiosira excentrica f. major Jousé 1959
Cells are solitary or united in short loose chains by a thick mucilage thread running through the central strutted
process. Connecting thread between the sibling cells in a chain is relatively long, about twice the length of
cell diameter. Frustules are drum-shaped in girdle view, usually shorter than broad. Valves are circular in
outline, nearly flat or slightly concave, 39-52 μm in diameter. Valve surface is coarsely areolated. Areolae are
large, hexagonal, closely packed, arranged in curved tangential rows with tendency to fasciolation, decrease
in size towards the margin, 7-9 in 10 μm in the central area and 12-13 in 10 μm near the margin. The central
areola is surrounded by seven (occasionally 6 or 8) areolae. A single tiny central fultoportula is located
close to central areola, opening externally by small pore and surrounded by 5 satellite pores internally.

MEDIOPHYCEAE
Plate 98. Thalassiosira diporocyclus.

a, b – live cells in valve (a) and girdle (b) view; c – whole valve in external view showing areolation and marginal
rimoportula (arrow); d – collapsed frustule in valve view, arrow points to rimoportula; e, f – internal valve
view showing central fultoportula (double arrowhead), ring of valve face fultoportulae (white arrowheads),
ring of marginal fultoportulae (black arrowheads), and marginal rimoportula (arrow); g – detail of valve in
external view showing valve face (white arrowheads) and marginal (black arrowheads) fultoportulae; h – detail
of valve margin in external view showing rimoportula (arrow) between two marginal fultoportulae (black
arrowheads); i – detail of valve in internal view showing central fultoportula with four satellite pores (double
arrowhead) and valve face fultoportulae with two or three satellite pores (arrowheads); j – detail of valve
margin in internal view showing valve face (white arrowheads), marginal (black arrowheads) fultoportulae
and rimoportula (arrow).
LM, BF illumination (a, b); SEM (c-j); scale bars: a-f – 5 μm; g-j – 2 μm.
Fultoportulae are scattered over the valve face and in two marginal rings, with 4 satellite pores internally.
Valve margin is furnished with a ring of coarse marginal spines. A single prominent rimoportula with a long
tube-like extension is present at the valve margin. Numerous round to elongated plate-shaped chloroplasts
are arranged around cell wall. Nucleus is centrally located.
Distribution:
Thalassiosira eccentrica is a neritic cosmopolitan species. It has been previously reported from the Arabian
Gulf (Simonsen, 1974), and from Kuwait’s waters (Al-Kandari et al., 2009) and intertidal sediments
(Al-Yamani & Saburova, 2011).
Local occurrence:
Thalassiosira eccentrica commonly occurred in Kuwait’s waters, being recorded from 58.5% of all
higher concentrations (1.0-2.5∙105 cells/l) recorded in the inshore waters of Kuwait Bay and around
Bubiyan Island.
Thalassiosira exigua Fryxell et Hasle

Plate 100
Hernández-Becerril & Tapia Peña, 1995: Figs. 23-25; Aké-Castillo et al., 1999: Figs. 7-9; Li et al., 2013: Figs. 53, 54.
Cells are small, solitary. Frustules are drum-shaped in girdle view, much shorter than broad. Valves are
circular in outline, almost flat, 5.9-10 μm in diameter. Valve areolae are coarse, arranged in linear rows,
decrease in size from the center to the margin, 22-26 in 10 μm in the central area and 30-33 in 10 μm near the
margin. Areolae on the mantle are much smaller in size. The central areola is surrounded by silicified border
and bears a fultoportula inside. The central fultoportula is surrounded by three satellite pores internally.
Marginal fultoportulae are arranged in regular ring, 1.5-1.7 μm apart, 6-6.5 in 10 μm, with tooth-shaped
outer extensions and surrounded by four satellite pores internally. A single rimoportula with short external
tube is present midway between two marginal fultoportulae, opening internally by obliquely oriented lip.
Distribution:
Thalassiosira exigua is a marine warm water species (Hernández-Becerril & Tapia Peña, 1995; Guiry &
Guiry, 2017).
Local occurrence:
Thalassiosira exigua was occasionally found in Kuwait’s inshore waters, however, the species could be
reliable identification requires SEM observations. The photographed specimens were isolated from
Kuwait Bay in November 2017. Thalassiosira exigua is reported for the first time from Kuwait.
Plate 99. Thalassiosira eccentrica.

a – live cells united to short chain in girdle view; b, c – single cells in girdle (b) and valve (c) view with
long extension of marginal rimoportula visible (arrow); d – single cells in valve view showing areolation;
e – detail of acid cleaned valve showing areolation, central fultoportula (double arrowhead) and marginal
rimoportula (arrow); f – external valve view showing central fultoportula (double arrowhead), scattered
valve face fultoportulae (arrowheads), and marginal rimoportula (arrow); g – detailed part of valve showing
central fultoportula (double arrowhead), scattered valve face fultoportulae (white arrowheads), marginal
fultoportulae (black arrowheads) and rimoportula (arrow); h – internal valve view showing scattered valve
face fultoportulae (white arrowheads), marginal fultoportulae (black arrowheads) and rimoportula (arrow).
i – detailed central part of valve in internal view showing central fultoportula (double arrowhead) and
scattered valve face fultoportulae (white arrowheads); j – detailed marginal part of valve in internal view
showing scattered valve face fultoportulae (white arrowheads), marginal fultoportulae (black arrowheads),
and marginal rimoportula (arrow).
LM, BF illumination (a-c), DIC optic (d), and phase contrast (e); SEM (f-j) scale bars: a – 20 μm; b-h – 10 μm;
i, j – 5 μm; g – 2 μm.

MEDIOPHYCEAE
Plate 100. Thalassiosira exigua.
a, b, e – acid cleaned frustules in valve view showing size variability and areolation, external tube of
rimoportula is indicated by arrow; c – girdle view showing ring of marginal fultoportulae with tooth-shaped
extensions (arrowheads); d, f, g – internal valve view showing central fultoportula (white arrowhead),
marginal fultoportulae (black arrowheads), and marginal rimoportula (arrow); h – detailed central area
showing the central bordered areola with fultoportula inside.
SEM; scale bars: a-g – 2 μm; h – 1 μm.

Thalassiosira hendeyi Hasle et Fryxell
Plates 101, 102
Hasle & Fryxell, 1977: Figs. 35-45; Muylaert & Sabbe, 1996: Figs. 13, 14; Aké-Castillo et al., 1999: Fig. 10; Sar et al.,
2001: Figs. 26-29; Hoppenrath et al., 2007: Fig. 29; García & Odebrecht, 2009a: Figs. 3-5, 23-31; García & Odebrecht,
2009b: Figs. 9, 40; Hoppenrath et al., 2009: Fig. 22 m-o; Li et al., 2013: Figs. 59-66; Park et al., 2016a: Figs. 16, S19,
S20.
Synonymized names:
Thalassiosira hustedtii Poretzky et Anissimova 1933
Coscinodiscus hustedtii Müller-Melchers 1953
Cells are solitary or united in short loose chains by a thick mucilage thread running through the central
strutted process. Connecting thread between the sibling cells in a chain is relatively long, about twice the
length of cell diameter. Frustules are drum-shaped in girdle view, usually shorter than broad. Valves are
circular in outline, nearly flat, 18-65 μm in diameter. Valve surface is coarsely areolated. Areolae are large,
hexagonal, closely packed, regularly arranged in straight tangential rows, 5-7 in 10 μm. Valve margin is
furnished with ribs, 12-13 in 10 μm, with 2-3 vertical rows of pores in between. A single small central
fultoportula is adjacent to the central areola, opening externally by raised rimmed pore and surrounded
internally by 5 satellite pores. Marginal fultoportulae are arranged in irregular ring at the valve mantle,
closely spaced, 3-4 μm apart, with short outer tubes and surrounded by 4 satellite pores internally. Two
MEDIOPHYCEAE
prominent rimoportulae with long tube-like extensions are present on opposite sides of the valve margin,
opening internally by obliquely oriented lips. Numerous plate-shaped chloroplasts are arranged around cell
wall. Nucleus is centrally located.
Distribution:
Thalassiosira hendeyi is a neritic warm water species widely distributed in south temperate to tropical
localities (Park et al., 2016a).
Local occurrence:
Thalassiosira hendeyi occurred sporadically in Kuwait’s coastal waters. It may be easily confused with
morphologically similar Thalassiosira eccentrica during routine phytoplankton examination. The species
is reported for the first time from Kuwait’s waters.
Thalassiosira lineata Jousé

Plate 103: a-c
Simonsen, 1974: Pl. 1: 4-7; Hernández-Becerril & Tapia Peña, 1995: Figs. 30-32; Aké-Castillo et al., 1999: Fig. 16; Li
et al., 2014: Figs. 40-42; Park et al., 2016a: Fig. 19.
Cells are solitary. Frustules are rectangular in girdle view. Valves are circular in outline, flat, 23-28 μm in
diameter. Valve surface is areolated. Areolae are hexagonal, closely packed, regularly arranged in straight
rows throughout the whole valve face, 10-13 in 10 μm. Small fultoportulae are scattered over the whole valve
face and arranged in two marginal rings. A single rimoportula is present on the valve margin. Numerous
plate-shaped chloroplasts are arranged around cell wall. Nucleus is centrally located.
Distribution:
Thalassiosira lineata is regarded as a warm water species reported mainly from tropical or subtropical
localities (Hasle & Fryxell 1977; Hasle & Syvertsen, 1997; Park et al., 2016a). The species has been
previously reported from the Indian Ocean (Simonsen, 1974).
Local occurrence:
Thalassiosira lineata was sporadically observed in Kuwait’s waters in low numbers, however, its occurrence
can be underestimated due to possible misidentification with other small-sized Thalassiosira species
during routine phytoplankton analysis in Lugol’s preserved samples. The species is reported for the first
time from Kuwait’s waters.
Plate 101. Thalassiosira hendeyi.
a – two live cells in girdle view united by organic thread (arrowhead); b – two cells in girdle (top) and valve
(bottom) view, arrow points to long extension of marginal rimoportula; c, d – cells in girdle view with
marginal rimoportulae visible (arrows); e-h – cells in valve view showing valve areolation and chloroplasts
arrangement, two opposite rimoportulae are indicated by arrows; i – collapsed frustule; j – detailed central
part of valve showing central fultoportula (arrowhead); k, l – detailed marginal rimoportula with long tube-
like extension.
LM, DIC optic; scale bars: a, b – 20 μm; c-i – 10 μm; j-l – 2 μm.

MEDIOPHYCEAE
Plate 102. Thalassiosira hendeyi.

a – detail of valve margin in external view showing marginal fultoportulae (black arrowheads) and
rimoportula (arrow); b, c – detailed central part of valve in external (b) and internal (c) view showing central
fultoportula; d – internal valve view, two opposite rimoportulae are indicated by arrows; e, f – detail of valve
margin in internal view showing marginal fultoportulae (black arrowheads) and rimoportula (arrow).
LM, SEM; scale bars: d – 10 μm; a-c, e, f – 2 μm.
a-c – Thalassiosira lineata: a – live cell in valve view; b, c – acid cleaned frustules in valve view, double
arrowhead points to central fultoportula; d-k – Thalassiosira lundiana: d – two live cells in girdle view united
by organic thread; e-j – single cells in valve view showing marginal ring of occluded processes with long
tube-like extensions (black arrowheads) and single marginal rimoportula (i, arrow); k – acid cleaned valve.
LM, DIC optic (a-j) and phase contrast (k); scale bars: 10 μm.

Thalassiosira lundiana Fryxell
Plates 103: d-k, 104
Sar et al., 2001: Figs. 31-33; Hoppenrath et al., 2007: Figs. 31, 32; Lee & Park, 2008: Pl. II: 7-12; García & Odebrecht,
2009a: Figs. 6, 7, 32-38; García & Odebrecht, 2009b: Figs. 10, 39, 41; Hoppenrath et al., 2009: Fig. 23a; Li et al., 2013:
Figs. 70-74; Li et al., 2014: Figs. 43-47; Lee et al., 2015: Fig. 9c; Park et al., 2016a: Fig. 21.
Cells are united in short loose chains by a thin mucilage thread running through the central strutted process.
Connecting thread between the sibling cells in a chain is relatively long, about twice the length of cell
diameter. Frustules are lenticular in girdle view, much shorter than broad. Valves are circular in outline,
slightly convex, 17-36 μm in diameter. Valve surface is finely areolated. Areolae are arranged radially in
fasciculate sectors, 28-30 in 10 μm. The valve margin is ribbed, 20-22 ribs in 10 μm. A single small fultoportula
is located at the valve center, opening externally by a small pore surrounded internally by 4 satellite pores.
Valve face fultoportulae vary in number, are irregularly scattered, with very short outer tubes and 2 satellite
pores internally. Marginal fultoportulae are arranged in a ring, alternating in location and forming a zig-zag
pattern. A single rimoportula with a long tube-like extension is present between two marginal fultoportulae,
opening internally by large radially oriented lip. A ring of 7-11 widely spaced occluded processes with
long tube-like extensions is located alongside the valve margin, along the ring of marginal fultoportulae.
Numerous plate-shaped chloroplasts are arranged around cell wall. Nucleus is centrally located.
Distribution:
MEDIOPHYCEAE
Thalassiosira lundiana is a neritic species widely distributed in temperate to subtropical waters (Hasle &
Syvertsen, 1997; Park et al., 2016a).
Local occurrence:
Thalassiosira lundiana was sporadically recorded in Kuwait’s inshore waters in low numbers, however, its
occurrence can be underestimated due to possible misidentification with other small-sized Thalassiosira
species during routine phytoplankton examination in Lugol’s preserved samples. The species is reported
for the first time from Kuwait’s waters.
Thalassiosira mala Takano

Plate 105
Takano, 1976, Figs. 1-18; Hallegraeff, 1984: Fig. 2; Hernández-Becerril & Tapia Peña, 1995: Figs. 33. 34; Aké-Castillo
et al., 1999: Figs. 17-19; Sar et al., 2002: Figs. 7-10; Park et al., 2009: Pl. 3: 13-18; Fernandes & Frassão-Santos, 2011:
Figs. 48-69; Li et al., 2013: Figs. 75-77; Li et al., 2014: Figs. 48, 49; Lee et al., 2015: Fig. 9d; Park et al., 2016a: Fig. 22.
Cells are small, solitary or often occur in agglomerates united by abundant mucilage. Frustules are drum-
shaped in girdle view, shorter than broad. Valves are circular in outline, slightly concave, 3.4-7 μm in diameter.
Valve areolae are small, arranged in curved tangential lines, much smaller towards the margins, 30-42 in
10 μm in the central area, 58-65 in 10 μm close to margins. On valve face, there is a single fultoportula
located off center in front of rimoportula. One marginal ring of regularly arranged fultoportulae is present.
A single rimoportula is located within a ring of marginal fultoportulae. Externally, all processes are without
tube extensions. Internally, subcentral fultoportula is surrounded by three satellite pores, and marginal
fultoportulae with four satellite pores. The cytoplasm contains a few plate-shaped chloroplasts. The nucleus
is centrally located.
Distribution:
Thalassiosira mala is a neritic warm water species widely distributed in subtropical and tropical waters
(e.g., Park et al., 2016a, Guiry & Guiry, 2017).
Local occurrence:
Thalassiosira mala was occasionally found in Kuwait’s inshore waters, however, the species may be
reliable identification requires SEM observations. Thalassiosira mala is reported for the first time from
Kuwait.
Plate 104. Thalassiosira lundiana.
a-d – acid cleaned valves in external view showing areolation, portula pattern, and occluded
processes; e – detailed valve center; f – detailed valve margin; g, h – internal valve view; i-k
– detailed valve margin in internal view; central fultoportula (double arrowhead), valve face
fultoportulae (white arrowheads), marginal fultoportulae (black arrowheads), rimoportula
(white arrow), and occluded processes (black arrows) are indicated.
SEM; scale bars: a-d, g, h – 5 μm; e, f, i-k – 2 μm.

MEDIOPHYCEAE
Plate 105. Thalassiosira mala.

a – live cells attached to sediments particles; b-e – frustules in oblique (b, c, e) and valve (d) view showing
single off-central fultoportula (double arrowhead), ring of marginal fultoportulae (black arrowheads), and
single marginal rimoportula (arrow); f, g – collapsed frustule showing external and internal valve view (f)
and detailed internal valve view (g) showing single off-central fultoportula (double arrowhead), marginal
fultoportulae (black arrowheads), and single marginal rimoportula (arrow).
LM, BF illumination (a); SEM (b-g); scale bars: a – 10 μm; b-g – 2 μm.
Harmful Effect:
Thalassiosira mala can be considered as a harmful nontoxic bloom-forming species. A bloom of this
species discoloring the water of Tokyo Bay in September 1951 was considered responsible for shellfish
mortality. Mechanical closing of respiration by a gelatinous substance densely attached to the gills
exuded from the diatom, together with poor quality water, was regarded as responsible for the death of
the bivalves (Takano, 1956, 1976).
Thalassiosira minima Gaarder

Plate 106: a-g
Harris et al., 1995: Fig. 31; Hoppenrath et al., 2007: Figs. 35-37; Li et al., 2013: Figs. 80-84; Li et al., 2014: Figs. 50-54.
Synonymized names:
Coscinosira floridana Cooper 1958
Thalassiosira floridana (Cooper) Hasle 1972
Cells are small, solitary or united in short chains, often embedded in mucilage and covered with attached
detritus particles. Frustules are drum-shaped in girdle view. Valves are circular in outline, almost flat, often
slightly concave at the center, 3.7-10 μm in diameter. Valve areolae are arranged in radial rows, 35-42 in
10 μm. Walls between the foramina are covered by minute siliceous granules. Two central fultoportulae,
seldom one, are located near valve center and surrounded by several larger areolae, open externally by short
tubes and surrounded with 3 satellite pores internally. 7-12 marginal fultoportulae are arranged in regular
ring, 1.4-2.6 μm apart, with short outer tubes and small nodular siliceous process in front of each. A single
rimoportula is located close to one fultoportula of the marginal ring, with long external tube and radially
oriented internal lip. The cytoplasm contains a few plate-shaped chloroplasts.
Distribution:
Thalassiosira minima is a widely distributed marine species (Park et al., 2016a, Guiry & Guiry, 2017).
Local occurrence:
Thalassiosira minima may be misidentified with other small-sized Thalassiosira species during routine
phytoplankton analysis and reliable identification requires SEM observations. In electron microscope
preparations, the species was well represented in the material collected from the inshore waters of Kuwait
Bay during the spring of 2017. Thalassiosira minima is reported for the first time from Kuwait’s waters.
Thalassiosira minuscula Krasske

Plate 106: h-j
Hallegraeff, 1984: Fig. 4a, b; Hernández-Becerril & Tapia Peña, 1995: Figs. 39-41; Aké-Castillo et al., 1999: Fig. 20; Sar
et al., 2001: Figs. 34-36; Hoppenrath et al., 2007: Figs. 38-40; Lee & Park, 2008: Pl. III: 13-18; Fernandes & Frassão-
Santos, 2011: Figs. 24-47; Li et al., 2013: Figs. 85-88; Li et al., 2014: Figs. 55-59.
Synonymized names:
Thalassiosira monoporocyclus Hasle 1972
Cells are solitary or occur in agglomerates united by abundant mucilage. Frustules are drum-shaped in
girdle view, shorter than broad. Valves are circular in outline, convex, 11-14 μm in diameter. Valve areolae
are small, arranged radially in fasciculate sectors, 38-44 in 10 μm. A single fultoportula is located at the
valve center. One marginal ring of regularly arranged fultoportulae is present, 2.2-2.7 μm apart, 2 in 10
μm. Externally, fultoportulae are opening by simple pores with raised edges, surrounded by a hyaline band,
and then a ring of 8-10 elongated areolae. A single rimoportula with long external tube is slightly displaced
inward from the ring of marginal fultoportulae, closing to one or two fultoportulae.
Distribution:
Thalassiosira minuscula is a widely distributed marine species (Park et al., 2016a, Guiry & Guiry, 2017).

MEDIOPHYCEAE

a-g – Thalassiosira minima: a, b – live cells in girdle (a) and valve (b) view; c-e – acid cleaned valves in external
view with one (c) and two (d, e) central fultoportulae, arrow points to outer tube of rimoportula; f – detailed
valve margin showing marginal fultoportulae (black arrowheads) with adjacent nodular processes (white
arrowheads); g – internal valve view, rimoportulae is indicated by arrow; h-j – Thalassiosira minuscula: h –
detailed valve margin showing marginal fultoportulae (black arrowheads) and outer aperture of broken tube
of rimoportula (arrow); i – external valve view; j – internal valve view, rimoportula is indicated by arrow.
LM, BF illumination (a, b); SEM (c-j); scale bars: a, b – 5 μm; c-e, g, i, j – 2 μm; f, h – 1 μm.
Local occurrence:
Thalassiosira minuscula was not distinguished from the other Thalassiosira species during routine
phytoplankton analysis; reliable identification requires SEM observations. In electron microscopy
preparations, the species was found to occur sporadically in the material collected from the inshore
waters of Kuwait Bay. Thalassiosira minuscula is reported for the first time from Kuwait’s waters.
Thalassiosira nanolineata (Mann) Fryxell et Hasle

Plate 96: j
Hallegraeff, 1984: Fig. 19a-e; Sar et al., 2001: Figs. 37-40; Li et al., 2013: Figs. 89-94; Park & Lee, 2010: Pl. 2: 9-14;
Park et al., 2016a: Fig. 28.
Synonymized names:
Coscinodiscus nanolineatus Mann 1925
Valve is circular in outline, convex, 8 μm in diameter. Valve areolae are arranged in straight parallel rows,
decrease in size from the center to the margin, 10.5 in 10 μm in the central area and 25 in 10 μm near the
margin. The external areolae foramina have radial siliceous threads. The entire valve surface is covered by
tiny granules. The central fultoportula opens externally by a tiny simple pore adjacent to the central areola.
Marginal fultoportulae are densely arranged in regular ring, 1.4 μm apart, 6.8 in 10 μm, open externally by
short tubes with double layered distal ends, with outer layer as a flared skirt with crenulated edge. A single
rimoportula with long external tube is present midway between two marginal fultoportulae. The outmost
valve margin is furnished with coarse ribs, 19-20 in 10 μm.
Remarks:
The morphology of a single valve found in Kuwait’s waters matches well with previous reports of
Thalassiosira nanolineata. A single central fultoportula was observed in Kuwait’s material, however, one
or more (up to four) central fultoportulae were reported for this taxon (Sar et al., 2001; Park & Lee,
2010; Li et al., 2013). The valve isolated from Kuwait’s waters is included as belonging to Thalassiosira
nanolineata based on the characteristic morphology of double layered external processes of marginal
fultoportulae, however, the morphological variability of this species in Kuwait needs to be examined in
details.
Distribution:
Thalassiosira nanolineata is a warm water marine species distributed from temperate to tropical waters
(Park et al., 2016a; Guiry & Guiry, 2017).
Local occurrence:
The species was not seen alive and not as whole frustules. Only one valve of this taxon was found in
electron microscopy preparation isolated from Kuwait Bay in May of 2017. Thalassiosira nanolineata is
Thalassiosira simonsenii Hasle et Fryxell

Plate 107
Hallegraeff, 1984: Fig. 22 a-d; Hernández-Becerril & Tapia Peña, 1995: Figs. 52-55; Aké-Castillo et al., 1999: Fig. 29;
Sar et al., 2001: Figs. 54-56; Park & Lee, 2010: Pl. V: 29-34; Park et al., 2016a: Figs. 39, S55, S56.
Cells are solitary or united in short loose chains by a thick mucilage thread running through the central strutted
process. Connecting thread between the sibling cells in a chain is relatively long, about twice the length of
cell diameter. Frustules are drum-shaped in girdle view, usually shorter than broad. Valves are circular in
outline, nearly flat, 26-44 μm in diameter. Valve surface is coarsely areolated. Areolae are large, hexagonal,
closely packed, regularly arranged in straight tangential rows, 7-8 in 10 μm. Valve margin is furnished with
ribs, 16-18 in 10 μm, with 2 vertical rows of pores in between. A single small central fultoportula is adjacent
to the smaller central areola, opening externally by raised rimmed pore and surrounded internally by 4-5

MEDIOPHYCEAE
Plate 107. Thalassiosira simonsenii.

a, b – live cells in valve (a) and girdle (b) view, arrowhead points to organic thread; c – acid cleaned
frustule in girdle view; d – external valve view; e – internal valve view; f-h – detail of valve margin in
external view; i – detail of valve margin in internal view; j – detailed central part of valve in external
view showing central fultoportula; central fultoportula (white arrowhead), marginal fultoportulae
(black arrowheads), and rimoportulae (arrow) are indicated.
LM, BF illumination (a, b); SEM (c-j); scale bars: a-e – 10 μm; f – 5 μm; g-j – 2 μm.
satellite pores. Marginal fultoportulae are arranged in regular ring at the valve mantle, closely spaced, 3-3.2
μm apart, 3-3.5 in 10 μm, with short outer tubes with crenulated edges. Two prominent rimoportulae with
long tube-like extensions are present on opposite sides of the valve margin within the ring of marginal
fultoportulae, opening internally by radially oriented small lips. Two adjacent to each rimoportula marginal
fultoportulae differ in direction of their outer processes. No occluded processes were observed. Numerous
plate-shaped chloroplasts are arranged around cell wall. Nucleus is centrally located.
Remarks:
Most morphological characters of the present specimens match well the previous reports of Thalassiosira
simonsenii from the different geographic sites worldwide (Hallegraeff, 1984; Hernández-Becerril & Tapia
Peña, 1995; Aké-Castillo et al., 1999; Sar et al., 2001; Park & Lee, 2010; Park et al., 2016a), except the
presence of occluded processes at valve mantle that were not observed in Kuwait’s material and probably
could be physically eroded during SEM preparation.
Distribution:
Thalassiosira simonsenii is a marine species distributed from temperate to tropical waters (Park et al.,
2016a).
Local occurrence:
Thalassiosira simonsenii occurred sporadically in Kuwait’s coastal waters, but may be easily confused
with morphologically similar Thalassiosira hendeyi during routine phytoplankton analysis. Reliable
identification requires SEM observations. The photographed specimens were isolated from Kuwait Bay
in November of 2017. The species is reported for the first time from Kuwait’s waters.
Thalassiosira sinica Li et Guo

Plate 108
Li et al., 2018: Figs. 1-19.
Cells are solitary, weakly silicified. Frustules are lenticular in girdle view. Valves are circular in outline,
slightly convex, 29-48 μm in diameter. Valve surface is finely areolated. Areolae are small, closely packed,
arranged in fasciculate rows, 47-52 in 10 μm. The central fultoportula is opening externally by a simple
pore with slightly raised rim, with 4 satellite pores internally. Valve face fultoportulae are arranged distantly
in irregular pattern, opening externally by raised pores surrounded by hyaline rim, with 4 satellite pores
internally. Marginal fultoportulae are arranged in regular ring very close to the valve margin, 4-4.7 μm apart,
2.2-2.5 in 10 μm, with short external tubes and 4 satellite pores internally. A single rimoportula with short
funnel-shaped external tube is present close to the ring of marginal fultoportulae, with radially oriented
lip internally. The outmost valve margin is furnished with ribs, 26-29 in 10 μm. Numerous plate-shaped
chloroplasts are arranged alongside the cell wall. Nucleus is centrally located.
Distribution:
Thalassiosira sinica is a marine planktonic species that has been recently described from Zhanjiang
Harbour and Daya Bay of the South China Sea (Li et al., 2018).
Local occurrence:
Thalassiosira sinica may be misidentified with Thalassiosira andamanica and other medium-sized Thalas-
siosira species during routine phytoplankton analysis. The most reliable distinctive character (morphology
of external processes of fultoportulae) is clearly discernible by SEM only. This taxon appears occasionally
in electron microscopy preparations from Kuwait’s coastal waters. The photographed specimens were
isolated from Kuwait’s southern inshore waters in April-May of 2008. Thalassiosira sinica is reported for
the first time from Kuwait.

MEDIOPHYCEAE
Plate 108. Thalassiosira sinica.

a, b – live cells in valve (a) and girdle (b) view; c – acid cleaned frustule in girdle view; d – external
valve view; e, f – detail of valve margin in external view; g – detail of valve margin in internal view;
h – detailed cingulum; i, j – detailed central part of valve in external (i) and internal (j) view showing
central fultoportula; central fultoportula (double arrowhead), valve face fultoportulae (white
arrowhead), marginal fultoportulae (black arrowheads), and rimoportula (arrow) are indicated.
LM, BF illumination (a, b); SEM (c-j); scale bars: a-d – 10 μm; e, h – 5 μm; f, g – 2 μm; i, j – 1 μm.
Thalassiosira teleata Takano
Plate 109
Harris et al., 1995: Figs. 9, 27; Hernández-Becerril & Tapia Peña, 1995: Figs. 62-64, 66, 67; Aké-Castillo et al., 1999:
Fig. 37; Hoppenrath et al., 2007: Fig. 52; Li et al., 2013: Figs. 114, 115; Li et al., 2014: Figs. 73, 74; Park et al., 2016a:
Fig. 41.
Cells are small, solitary or united in short chains, often embedded in mucilage and covered with attached
detritus particles. Frustules are drum-shaped in girdle view, shorter than broad. Valves are circular in outline,
almost flat, slightly concave at the center, with high mantle, 5.2-7.1 μm in diameter. Valve areolae are arranged
in radial rows, 41-43 in 10 μm. Central fultoportula with short outer tube is located adjacent to large areola.
Marginal fultoportulae are arranged in regular ring, 1.6-2.2 μm apart, with long external ornamented by two
wings on the top. A single rimoportula is located close to one fultoportula of the marginal ring, with long
external tube and radially oriented internal small lip. The cytoplasm contains a few plate-shaped chloroplasts.
Distribution:
Thalassiosira teleata is a marine cosmopolitan species distributed from cold to tropical waters (Park et al.,
2016a).
Local occurrence:
Thalassiosira teleata occurred sporadically in Kuwait’s coastal waters, but may be easily confused with
morphologically similar small-sized Thalassiosira species during routine phytoplankton analysis. Reliable
identification requires SEM observations. The photographed specimens were isolated from Kuwait Bay
in November 2017. The species is reported for the first time from Kuwait’s waters.
Thalassiosira tenera Proschkina-Lavrenko

Plate 110
Aké-Castillo et al., 1999: Figs. 38-44; Sar et al., 2001: Figs. 58, 59; Hoppenrath et al., 2007: Figs. 53, 54; Lee & Park,
2008: Pl. VI: 35-42; Li et al., 2013: Figs. 116, 117; Li et al., 2014: Figs. 75-77; Lee et al., 2015: Fig. 9 j-l; Park et al.,
2016a: Fig. 42.
Cells are small, solitary. Frustules are drum-shaped in girdle view, shorter than broad. Valves are circular in
outline, almost flat, 5.2-8.1 μm in diameter. Valve areolae are coarse, arranged in linear rows, decrease in size
from the center to the margin, 22-27 in 10 μm in the central area and 45-52 in 10 μm near the margin. Areolae
on the mantle are much smaller in size, up to 68-75 in 10 μm. The central fultoportula is surrounded by
central areola and with three satellite pores internally. Marginal fultoportulae with tooth-shaped extentions
are arranged in regular ring and surrounded by four satellite pores internally. A single rimoportula with short
external tube is present within marginal ring directly next to fultoportula. Larger heavily silicified frustules
(18-22 μm in diameter) with much coarser aleolation (5-6 areolae in 10 μm) were ocassionaly observed (Pl.
110: i, k, l). The cytoplasm contains a few plate-shaped chloroplasts.
Distribution:
Thalassiosira tenera is a cosmopolitan widely distributed species (e.g., Hasle & Syvertsen, 1997; Park et
al., 2016a, Guiry & Guiry, 2017).
Local occurrence:
Thalassiosira tenera was occasionally found in Kuwait’s inshore waters, however, the species may be
misidentified with other small-sized Thalassiosira species during routine phytoplankton examination, and
reliable identification requires SEM observations. The illustrated specimens were isolated from Kuwait’s
southern inshore waters in March 2008. Thalassiosira tenera is reported for the first time from Kuwait.

MEDIOPHYCEAE
Plate 109. Thalassiosira teleata.

a, b – live cells in girdle view; c-e – acid cleaned valves in external view; f – internal valve view; g – detailed
central part of valve in external view showing central fultoportula; h – detail of valve margin in external
view; central fultoportula (white arrowhead), marginal fultoportulae (black arrowheads), and rimoportula
(arrow) are indicated.
LM, BF illumination (a, b); SEM (c-h); scale bars: a-f – 2 μm; g, h – 1 μm.
Plate 110. Thalassiosira tenera.
a – live cell in valve view; b – acid cleaned frustule in valve view; c – frustule in oblique valve
view; d, e – external valve view at different magnifications; f, g – internal valve view at different
magnifications; h, i – detailed valve center; j – small weakly silicified frustule; k, l – larger heavily
silicified valves; central fultoportula (white arrowhead), marginal fultoportulae (black arrowheads),
and rimoportula (arrow) are indicated.
LM, BF illumination (a) and phase contrast (b); SEM (c-l); scale bars: a, b – 10 μm; k, l – 5 μm; c, d,
f, i – 2 μm; e, g, h – 1 μm.

Genus Tryblioptychus Hendey
Tryblioptychus cocconeiformis (Grunow) Hendey
Plate 111
Hendey, 1957: Pl. 2: 10; Tynni, 1983: Fig. 1, Pls. 2: 18 a-m, 11 (as Cyclotella crassa); Prasad et al., 2002: Figs. 1-44; Al-
Handal, 2009: Fig. 12; Sar et al., 2010b: Figs. 10-13; Garibotti et al., 2011: Fig. 9.
Synonymized names:
Campylodiscus cocconeiformis Grunow 1883
Cyclotella crassa Tynni 1983
Frustules are drum-shaped in girdle view. Valves are subcircular to broadly elliptical, 16-18 μm long and
14-18 μm wide. Valve face is undulated, divided into two halves lying in different focal planes. Valve surface
is furnished with radially arranged fascicles of areolae, 9-12 per valve. Slender elevated interfascicles are
hyaline and alternate with wider depressed fascicles of areolae. Areolae are coarse, 11-13 in 10 μm. The
entire valve surface is covered by tiny granules. Valve edge is furnished by evenly spaced chambers with large
round external openings, 8-10 in 10 μm, giving the appearance of coarse striation in LM. A single valve
face fultoportula is located subcentrally in interfascicle, externally appears as a small pore. There is a ring
of marginal fultoportulae occurring at the mantle end of each interfascicle, 4.5-5 μm apart, with external
openings as raised pores. A single rimoportula is located marginally, opens externally by long tube with
flared distal end.
MEDIOPHYCEAE
Remarks:
In 1983, Cyclotella crassa was originally described from gastropod shells collected in Khawr Abdallah,
southern Iraq by Tynni (Tynni, 1983: Fig. 1, Pl. 2: 18 a-m, Pl. 11). This diatom regarded as sp. nov.
closely matches gross morphology of Tryblioptychus cocconeiformis (Grunow) Hendey 1958 in regards
to shape, size, and valve ornamentation. No morphological character separates these two species,
therefore, Cyclotella crassa Tynni 1983 may be synonymized with Tryblioptychus cocconeiformis, which has
nomenclatural priority (see also Prasad et al., 2002; Al-Handal, 2009).
Distribution:
Tryblioptychus cocconeiformis is a marine, primarily benthic species, secondarily displaced into the water
column in neritic habitats, with distribution confined to warm water areas (Prasad et al., 2002; Guiry &
Guiry, 2017). The species has been previously reported from the bottom sediments in the Shatt Al-Arab
Estuary, Iraq (Tynni, 1983, as Cyclotella crassa; Al-Handal, 2009).
Local occurrence:
Frustules of Tryblioptychus cocconeiformis were found in plankton net hauls from the inshore waters
around Bubiyan Island in low numbers. The species is reported for the first time from Kuwait’s waters.
Plate 111. Tryblioptychus cocconeiformis.
a, b – acid cleaned frustules in valve view; c – frustule in oblique valve view, arrows point to outer
tubes of opposite marginal rimoportulae; d, e – valve view, arrow points to marginal rimoportula; f, g
– detailed valve parts showing subcentral fultoportula (white arrowhead), marginal fultoportulae (black
arrowheads), and rimoportula (arrow).
LM, phase contrast (a, b); SEM (c-g); scale bars: a, b – 10 μm; c-e – 5 μm; f, g – 2 μm.

Class Bacillariophyceae Haeckel
Subclass Urneidophycidae Medlin
Order Rhaphoneidales Round
Family Rhaphoneidaceae Forti
Genus Asterionellopsis Round
Asterionellopsis glacialis (Castracane) Round ex Round, Crawford et Mann

Plate 112
Allen & Cupp, 1935: Fig. 98 (as Asterionella japonica); Cupp, 1943: Fig. 138 (as Asterionella japonica); Hasle & Syvertsen,
1997: Pl. 50; Hoppenrath et al., 2009: Fig. 44 m, n.
Synonymized names:
Asterionella glacialis Castracane 1886
Asterionella japonica Cleve 1882
Cells are united in star-shaped or spiral colonies by the corners of the enlarged base. In girdle view, frustules
are heteropolar, with the expanded triangular basal part, tapering abruptly to a slender and much longer head
region. Valves are very narrow, with a widened knob-like region at the base, 38-55 µm long. Valve surface is
finely striated. A single triangular plate-shaped chloroplast is located in the enlarged basal part of the cell.
Distribution:
Asterionellopsis glacialis is a cosmopolitan neritic species widely distributed in temperate and warm waters.
The species has been previously reported from the Arabian Gulf (Simonsen, 1974, as Asterionella glacialis)
and from Kuwait’s waters (Al-Kandari et al., 2009).
Local occurrence:
Asterionellopsis glacialis is a rarely observed species, recorded from 2% of all collected samples, principally
in the coastal waters of southern Kuwait, observed more abundantly (0.4-1.9∙105 cells/l) during the
warm season from April to September.
Genus Delphineis Andrews
Delphineis surirella (Ehrenberg) Andrews

Plate 113: e
Hendey, 1970: Pl. 3: 33 (as Rhaphoneis surirella); Andrews, 1981: Pls. 1, 2.
Synonymized names:
BACILLARIOPHYCEAE
Zygoceros surirella Ehrenberg 1840
Rhaphoneis surirella (Ehrenberg) Grunow 1881
Cells are small, solitary. Valves are elliptical, isopolar, with slightly produced bluntly rounded apices, 12-15
μm long, 7.5-10 μm broad. Valve surface is finely striated. Transapical striae are coarsely punctate, parallel
in the middle, becoming radiate towards the apices, 14-17 in 10 μm. Valves are araphid. Sternum is hyaline,
narrow, widening slightly near the apices. A single small rimoportula is present at each apex adjacent to the
opposite sides of sternum, but occasionally both rimoportulae are present at the same side.
Distribution:
Delphineis surirella is a primarily benthic and widely distributed species (Hendey, 1964) that occurs
occasionally in plankton samples. It has been previously reported from Kuwait’s Mina Al-Ahmadi shore
(Hendey, 1970; as Rhaphoneis surirella).
Local occurrence:
Delphineis surirella is sporadically found in plankton net hauls from Kuwait’s inshore waters.
Plate 112. Asterionellopsis glacialis.
a-c – live cells united to long spiral chains; d – chain in apical view; e – detailed part of chain showing
connection of sibling cells; f – detail of frustule in girdle view.
LM, BF illumination (a-e); SEM (f); scale bars: a-d – 50 μm; e – 20 μm; f – 2 μm.

Genus Diplomenora Blazé
Diplomenora cocconeiformis (Schmidt) Blazé

Plate 113: a-c
Foged, 1984: Pl. XXVII: 10 (as Rhaphoneis superba); Witkowski et al., 2000: Pls. 22: 1, 2, 23: 8-11; Hein et al., 2008: Fig.
7: 8; Al-Kandari et al., 2009: p. 85, Pl. 47: F-H; Al-Yamani & Saburova, 2011: Pl. 24: a-f.
Synonymized names:
Raphoneis superba (Janish) Grunow 1862
Coscinodiscus cocconeiformis Schmidt 1878
Valves are circular in outline, 18-25 μm in diameter. Sternum is narrow, linear. Valve surface is areolated,
areolae are arranged in concentric arcs radiating from sternum, 14-15 rows in 10 μm. Apical arcs form
circular structures. A single small rimoportula is present at each apex adjacent to the opposite sides of
sternum, but ocassionally both rimoportulae are present at the same side.
Distribution:
Diplomenora cocconeiformis is a common species of the Indian Ocean coasts, predominantly distributed
in warm waters. It has been recorded in Kuwait’s coastal waters (Al-Kandari et al., 2009), and was a
frequently observed species in Kuwait’s intertidal sandflats (Al-Yamani & Saburova, 2011).
Local occurrence:
Diplomenora cocconeiformis is a typically benthic species, which occasionally becomes stirred up from the
bottom sediments and carried into the water column. Frustules of this species were sporadically found
in plankton net hauls from Kuwait’s inshore waters.
Genus Perissonoë Andrews et Stoelzel
Perissonoë cruciata (Janisch et Rabenhorst) Andrews et Stoelzel

Plate 113: d
Hendey, 1970: Pl. 4: 41( as Rhaphoneis amphiceros var. tetragona); Round et al., 1990: p. 414-415; Hein et al., 2008: Pl.
11: 7; Lobban et al., 2012: Pl. 14: 1, 2.
Synonymized names:
Amphitetras cruciata Janisch et Rabenhorst 1863
Triceratium cruciatum (Janisch et Rabenhorst) Cleve 1878
BACILLARIOPHYCEAE
Rhaphoneis amphiceros var. tetragona Grunow in Van Heurck 1883
Rhaphoneis amphiceros var. cruciata (Janisch et Rabenhorst) Mereschkowsky 1902
Valves are quadrate in outline, with side of 22-25 μm, with round apices and slightly inflated sides. Four
sterna run diagonally from corners towards the center, forming a cruciform hyaline area. Valve surface is
areolated, areolae are arranged in concentric arcs radiating from each sternum, 7-8 rows in 10 μm.
Distribution:
Perissonoë cruciata is a marine benthic species distributed in subtropical and tropical regions (Lobban et
al., 2012; Guiry & Guiry, 2017). It has been previously reported from Kuwait’s Mina Al-Ahmadi shore
(Hendey, 1970; as Rhaphoneis amphiceros var. tetragona).
Local occurrence:
Perissonoë cruciata is a typically benthic species, which occasionally becomes stirred up from the bottom
sediments and carried into the water column. Frustules of this species were sporadically found in
plankton net hauls in Kuwait’s inshore waters.
Subclass Fragilariophycidae Round
Order Licmophorales Round
Family Licmophoraceae Kützing
Genus Licmophora Agardh
Licmophora abbreviata Agardh

Plate 113: f-l
Péragallo & Péragallo, 1897-1908: Pl. LXXXV: 8-13 (as Licmophora lyngbyei); Cupp, 1943: Fig. 127; Foged, 1984: Pl.
XXX: 9; Hustedt, 1985: Fig. 590; Witkowski et al., 2000: Pl. 20: 3-5; Al-Yamani & Saburova, 2011: Pl. 22: a-k.
Synonymized names:
Podosphenia abbreviata Ehrenberg 1838
Podosphenia lyngbyei Kützing 1844
Rhipidophora abbreviata Kützing 1844
Licmophora lyngbyei Grunow 1867
Licmophora lyngbyei var. abbreviata, var. elongata, var. minuta Grunow 1880
Cells in girdle view are wedge-shaped with rounded upper corners, deeply penetrating septa, and more or
less strongly waved intercalary bands. Valves are heteropolar, narrow, club-shaped, with rounded head-pole
and rostrate rounded foot-pole, 52-67 μm long, 7-8 μm broad (in the middle). Sternum is narrow, linear, and
distinct. Valve surface is coarsely striated. Transapical striae are robust, parallel, 13-15 in 10 μm. Numerous
small elongated chloroplasts are scattered throughout the cytoplasm.
Distribution:
Licmophora abbreviata is a cosmopolitan marine littoral species, but often found in plankton collections.
It has been previously reported from epiphytic assemblages on different substrata (macrophytes, rocks,
sedentary macrofauna) along Kuwait’s shores (Al-Yamani & Saburova, 2011).
Local occurrence:
Licmophora abbreviata occurred occasionally across Kuwait’s waters, being recorded from 14% of all
higher concentrations (0.6-1.3∙104 cells/l) recorded in the inshore waters during the months of November
and December.
Plate 113. Diplomenora cocconeiformis, Delphineis surirella, Perissonoë cruciata and Licmophora abbreviata.
a-c – Diplomenora cocconeiformis: a, b – acid cleaned valves; c – external valve view;
d – Perissonoë cruciata: acid cleaned valve;
e – Delphineis surirella: internal valve view;
f-l – Licmophora abbreviata: f – live cell in girdle view; g, h – cells in oblique view; i – acid cleaned valve; j-l
– acid cleaned frustules in girdle view.
LM, BF illumination (f-h) and phase contrast (a, b, d, i-k); SEM (c, e, l); scale bars: a, b, d, f-l – 10 μm; c – 5
μm; e – 2 μm.

BACILLARIOPHYCEAE
Order Rhabdonematales Round et Crawford
Family Grammatophoraceae Lobban et Ashworth
Genus Grammatophora Ehrenberg
Grammatophora oceanica Ehrenberg

Plate 114
Péragallo & Péragallo, 1897-1908: Pl. LXXXVII: 9-17; Hendey, 1970: p. 128; Hustedt, 1985: Figs. 573, 574; Podzorski
& Håkansson, 1987: Pl. 7: 4; Ricard, 1987: Fig. 662; Witkowski et al., 2000: Pls. 15: 13, 14; 16: 12; 17: 3, 4; Al-Handal,
2009: Fig. 49; Al-Kandari et al., 2009: p. 88, Pl. 49: F-I; Al-Yamani & Saburova, 2011: Pl. 26: a-g.
Synonymized names:
Grammatophora marina var. communis Grunow 1880
Cells are united by corners to form zigzag-shaped chains. Frustules are rectangular in girdle view, with
rounded corners and slightly wavy septa. Valves are linear, slightly widened in the middle, somewhat
constricted between middle and apices, 21-55 μm long, 5-6 μm wide. Apices are capitately rounded. A single
pseudosepta is present in the middle of the valve. Valve surface is finely striated. Transapical striae are finely
punctate, parallel, 25-28 in 10 μm. Several irregularly shaped plate-like chloroplasts are arranged along the
girdle.
Distribution:
Grammatophora oceanica is a cosmopolitan species of marine and brackish water coasts. It has been
previously reported from bottom sediments in the Shatt Al-Arab Estuary, Iraq (Al-Handal, 2009), from
Kuwait’s Mina Al-Ahmadi shore by Hendey (1970), and from Kuwait’s coastal waters (Al-Kandari et
al., 2009) and intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Grammatophora oceanica rarely occurred in Kuwait’s waters, being recorded from 4% of all collected
samples. The species was usually present in low to moderate concentrations, more abundantly (1.8-
8.1∙103 cells/l) in the southern coastal waters during the fall season from September to November.
Order Thalassionematales Round

Family Thalassionemataceae Round
Genus Thalassionema Grunow ex Mereschkowsky
Thalassionema frauenfeldii (Grunow) Tempère et Péragallo

Plate 115
Pavillard, 1925: Fig. 106A; Allen & Cupp, 1935: Fig. 97; Cupp, 1943: Fig. 135; Hendey, 1964: p. 165; Simonsen, 1974:
Pl. 24: 4; Hoppenrath et al., 2009: Fig. 45 j-l.
Synonymized names:
Asterionella frauenfeldii Grunow 1863
Thalassiothrix frauenfeldii (Grunow) Grunow 1880
Cells are very long, needle-shaped, united into star-shaped or zigzag colonies by small mucilage cushions
on one cell end. Frustules are narrowly-rectangular in girdle view. Valves are linear in outline, with parallel
sides, with more or less tapering one end with a spine, and broader rounded the other end, 110-140 µm long
and 3-4 µm wide. Sternum is wide. Marginal striae are regularly arranged, 7-9 in 10 µm. Numerous small
plate-like chloroplasts are arranged along valves.
Distribution:
Thalassionema frauenfeldii is an oceanic, south temperate to tropical species with widespread distribution.

BACILLARIOPHYCEAE
Plate 114. Grammatophora oceanica.

a – live cells in girdle view united corner to corner with pads of mucilage to zigzag chain; b, c – detailed
sibling cells; d-g – acid cleaned frustules in girdle view; h, i – acid cleaned frustules in valve view.
LM, BF illumination (a-c) and phase contrast (d-i); scale bars: a – 20 μm; b-i – 10 μm.
Plate 115. Thalassionema frauenfeldii.
a-d – live cells united by small mucilage cushions on one cell end to star-shaped, zigzag or combined chains;
e-g – ends of acid cleaned frustules in girdle view with spines (arrowheads); h – detailed live cell in girdle
view showing single spine at each valve end (arrowheads); i, j – detailed sibling cells, arrowheads point single
spines at each valve.
LM, BF illumination (a, b, d), DIC optic (c, h, i), and phase contrast (e-g); SEM (j); scale bars: a, b, d – 50
μm; c, i – 25 μm; f, h – 20 μm; e, g – 10 μm; j – 5 μm.

Local occurrence:
Thalassionema frauenfeldii occurred commonly in Kuwait’s waters, being recorded from 62% of all
period from May to November.
Thalassionema nitzschioides (Grunow) Mereschkowsky

Plate 116: e-i
Allen & Cupp, 1935: Fig. 96 (as Thalassiothrix nitzschioides); Cupp, 1943: Fig. 133; Hendey, 1964: p. 165; Hendey, 1970:
p. 127; Simonsen, 1974: p. 37; Ricard, 1987: Figs. 583-586; Throndsen et al., 2007: p. 186; Hoppenrath et al., 2009: Fig.
45 m-o.
Synonymized names:
Synedra nitzschioides Grunow 1862
Thalassiothrix nitzschioides (Grunow) Grunow 1881
Thalassiothrix nitzschioides var. javanica Grunow 1881
Thalassiothrix fraunfeldii var. nitzschioides (Grunow) Jörgensen 1900
Synedra nitzschioides var. minor Cleve 1883
Thalassiothrix curvata Castracane 1886
Cells are long, united into star-shaped or zigzag colonies by small mucilage cushions on one cell end. Frustules
are narrowly-rectangular in girdle view, sometimes slightly curved. Valves are linear to narrowly-lanceolate
in outline, with parallel sides and blunt-rounded ends, 40-90 μm long and 3-5 μm wide. Sternum is wide.
Areolae are regularly arranged in two marginal rows, 10-12 in 10 μm. The valvar structure is manifested in
SEM as a single row of small foramina internally, and external depressions partially occluded by Y-shaped
bars with a varying number of lateral branches. Rimoportula is located at each pole of valve. Externally,
rimoportulae open by apical protrusions, the more prominent is small arrow-shaped, and the other one at
the opposite pole is a short tube-shaped. More or less numerous small plate-like chloroplasts are arranged
along valves.
Distribution:
Thalassionema nitzschioides is a cosmopolitan, predominantly neritic species. It has been previously
reported from the Arabian Gulf (Simonsen, 1974), and from Kuwait’s shores (Hendey, 1970) and coastal
Local occurrence:
Thalassionema nitzschioides is the most common phytoplankton diatom species in Kuwait’s waters, being
BACILLARIOPHYCEAE
recorded from 79% of all collected samples. The species is present throughout the year in low to moderate
abundance with higher concentrations (0.6-1.8∙105 cells/l) recorded in the inshore waters during the
warm season from June to September.
Thalassionema pseudonitzschioides (Schuette et Schrader) Hasle

Plate 116: j, k
Hasle & Syvertsen, 1997: Pls. 56: 4, 57; Hasle, 2001: Figs. 28-42; Sar et al., 2007: Fig. 3 K-S; Sugie & Suzuki, 2015:
Figs. 28, 29.
Synonymized names:
Thalassiotrix pseudonitzschioides Schuette et Schrader 1982
Cells are long, united into star-shaped or zigzag colonies by small mucilage cushions on one cell end. Frustules
are narrowly-rectangular in girdle view, sometimes slightly curved. Valves are linear to narrowly-lanceolate
in outline, with parallel sides and blunt-rounded ends, 35-49 μm long and 3-4 μm wide. Sternum is wide.
Areolae are regularly arranged in two marginal rows, 10-12 in 10 μm. The valvar structure is manifested in
SEM as a single row of small foramina internally, crossed externally by simple Y-shaped bars. Rimoportula
is located at each pole of valve. Externally, rimoportulae open by apical protrusions, the more prominent is
small arrow-shaped, and the other one at the opposite pole is a short tube-shaped. Numerous small plate-like
chloroplasts are arranged along valves.
Remarks:
Thalassionema pseudonitzschioides is highly similar in morphology to Thalassionema nitzschioides (Grunow)
Mereschkowsky. Hallegraeff (1986) regarded the two species as conspecific, however, T. pseudonitzschioides
differs from T. nitzschioides by the structure of the external areolar occlusions (simple Y-shaped bars
versus Y-shaped bars with a varying number of lateral branches), the orientation of the labiate processes
and by having a more pronounced marginal ridge and an arrow-shaped protrusion (Hasle, 2001). The
specimens from Kuwait’s waters were identified as Thalassionema pseudonitzschioides based on external
structure of areolae as manifested in SEM.
Distribution:
Thalassionema pseudonitzschioides is a warm water species distributed from temperate to subtropical
localities (Hasle, 2001). In previous phytoplankton surveys in Kuwait’s waters (Al-Yamani et al., 2004;
Al-Kandari et al., 2009), the species might have been misidentified as Thalassionema nitzschioides, and
these records require verification with SEM.
Local occurrence:
Thalassionema pseudonitzschioides can be easily misidentified with Thalassionema nitzschioides during
phytoplankton analysis since critical diagnostic character (the morphology of the areola occlusions)
is not visible with light microscopy and requires SEM examination. Frustules of Thalassionema
pseudonitzschioides appear occasionally in electron microscope preparations from Kuwait’s coastal waters.
The photographed specimens were isolated from Kuwait’s southern waters in April 2008. This species is
reported for the first time from Kuwait.
Subclass Bacillariophycidae Mann

Order Cocconeidales Cox
Family Cocconeidaceae Kützing
Genus Anorthoneis Grunow
Anorthoneis excentrica (Donkin) Grunow

Plate 118: h
Witkowski et al., 2000: Pls. 42: 20, 54: 8, 10.
Synonymized names:
Cocconeis excentrica Donkin 1858
Valves are nearly circular in outline, 22-25 µm in diameter. Raph is straight, displaced from the apical axis,
with distant external central endings, and apical endings terminated at a certain distance from apices. Axial
area is very narrow, central area is small, nearly circular. Transapical striae are coarsely punctate, strongly
radiate, 11-12 in 10 µm.
Distribution:
Anorthoneis excentrica is a marine species inhabiting intertidal sandy sediments (Witkowski et al., 2000).
Local occurrence:
Anorthoneis excentrica is a typically benthic species, which sporadically occurs in the pelagic microalgal
assemblages. A few frustules of this species were found in plankton net hauls in Kuwait’s northern
inshore waters around Bubiyan Island. The species is reported for the first time from Kuwait.

BACILLARIOPHYCEAE
Plate 116. Thalassionema spp..

a-d – Thalassionema nitzschioides/pseudonitzschioides species complex: a-c – live cells united by small
mucilage cushions on one cell end to star-shaped, zigzag or combined chains; d – acid cleaned frustule
in valve view; e-i – Thalassionema nitzschioides: e – acid cleaned frustule in valve view; f, g – detail
of valve showing areolation pattern; h, i – detailed apices; j, k – Thalassionema pseudonitzschioides:
j – acid cleaned frustule in valve view; k – detail of valve showing areolation pattern.
LM, DF illumination (b), DIC optic (a, c), and phase contrast (d); SEM (e-k); scale bars: a, b – 20
μm; c, e – 10 μm; d, j, k – 5 μm; f, g – 2 μm; h, i – 1 μm.
Genus Cocconeis Ehrenberg
The species belonging to the genus Cocconeis are distinguished by heterovalvar frustules, valves with a very
narrow valve mantle, and ring near the edge of the raphid valve. Ornamentation of the raphe valve may differ
markedly from that of the rapheless valve. Species are identified by the shape and size of valve, the shape of
raphe, the type and density of striae, and the type, shape and density of areolae. The identification of Cocconeis
species requires a careful observation of both raphe and rapheless valves composing the frustule in outside
and inside view. Some morphological specificities of these relatively small taxa are hardly discernible in
frustules preserved on microscope slides in refractive mounting media with LM, and a number of diagnostic
features can be clearly observed using electron microscopy. The genus Cocconeis is known to be benthic, with
the valves firmly attached to various substrata, where they may constitute assemblages with high abundance
(Riaux-Gobin et al., 2010, 2011).
Cocconeis coronatoides Riaux-Gobin et Romero

Plate 117: j
Riaux-Gobin et al., 2010: Figs. 1-3; Riaux-Gobin et al., 2011: Pls. 34-36; Lobban et al., 2012: Pl. 39: 6.
Synonymized names:
Cocconeis coronata Riaux-Gobin et Romero 2010
Valves are elliptical with rounded apices, 28-32 μm long, 17-20 μm broad. On rapheless valve, sternum is
straight and narrow. Valve surface is coarsely areolated. Areolae are round, arranged in a regular hexagonal
pattern. Transapical striae are parallel in the middle, becoming radiate towards the apices, 14-16 in 10 μm.
A distinct submarginal hyaline rim is running throughout the valve parallel to the margin, marks the point
where the striae bifurcate and the valve slopes down to the margin, separating the outermost three rows of
small areolae. Numerous small spines are irregularly scattered around the areolae.
Distribution:
Cocconeis coronatoides was reported from the shores of tropical Indian Ocean and Pacific Islands (Riaux-
Gobin et al., 2010, 2011; Lobban et al., 2012).
Local occurrence:
A few frustules of Cocconeis coronatoides were found in plankton net hauls from Kuwait’s southern inshore
waters. The species is reported for the first time from Kuwait.
Cocconeis neothumensis var. marina De Stefano, Marino et Mazella

Plate 118: e
De Stefano et al., 2000: Figs. 53-65; Sar et al., 2003: Figs. 26-31.
Valves are elliptically lanceolate with rounded apices, 18-20 μm long, 9-10 μm broad. On rapheless valve,
sternum is narrow, straight. Transapical striae are parallel in the middle, becoming radiate towards apices,
21-22 in 10 μm. Areolae are transapically elongated, 4-5 in each hemivalve, arranged in wavy longitudinal
rows.
Distribution:
Cocconeis neothumensis var. marina has been originally described from the Mediterranean, Gulf of Naples
(De Stefano et al., 2000), and further was reported from the southwestern Atlantic Ocean (Sar et al.,
2003).
Local occurrence:
A few frustules of Cocconeis neothumensis var. marina were obtained from plankton net hauls in Kuwait’s
southern inshore waters. The species is reported for the first time from Kuwait.

BACILLARIOPHYCEAE
Plate 117. Cocconeis spp.

a-g, i – Cocconeis scutellum s.l.: a, b – live cells in valve view; c – empty frustule in rapheless valve view; d-f,
i – acid cleaned rapheless valves; g – acid cleaned raphe valve; h – Cocconeis scutellum var. posidoniae: acid
cleaned rapheless valve; j – Cocconeis coronatoides: acid cleaned rapheless valve.
LM, BF illumination (a-c) and phase contrast (d-g); SEM (h-j); scale bars: a-h – 10 μm; i, j – 5 μm.
Cocconeis placentula var. euglypta (Ehrenberg) Grunow
Plate 118: a-d
Hustedt, 1985: p. 308, fig. 802 c; Al-Yamani & Saburova, 2011: Pl. 53: a-c; Romero & Jahn, 2013: Figs. 9-18; Al-Handal
et al., 2014: Figs. 10-14.
Synonymized names:
Cocconeis euglypta Ehrenberg 1854
Cocconeis lineata var. euglypta (Ehrenberg) Grunow 1880
Cocconeis rouxii var. euglypta (Ehrenberg) Héribaud 1893
Cyclotella lineata var. euglypta Gutwinski 1895
Cocconeis placentula f. euglypta (Ehrenberg) Hustedt 1957
Valves are elliptically oblong with rounded apices, 20-25 μm long, 12-14 μm broad. On raphe valve, raphe
is straight, axial area is narrow, central area is small, circular. Transapical striae are composed of dot-like
areolae, slightly radiate, interrupted towards margin by raised siliceous rim. On rapheless valve, sternum is
narrow, linear, transapical striae are parallel in the middle, becoming radiate towards apices, 20-21 in 10 μm.
Areolae are transapically elongated, arranged in wavy longitudinal rows. Two plate-like irregularly shape
chloroplasts are arranged along valves on either side of the central nucleus.
Distribution:
Cocconeis placentula var. euglypta is a marine and brackish water cosmopolitan epiphytic species. It has
been previously reported from the epiphytic assemblages in the Mesopotamian wetland, South Iraq (Al-
Handal et al., 2014), from the bottom sediments in the Shatt Al-Arab Estuary, Iraq (Al-Handal, 2009),
and from Kuwait’s coastal waters (Al-Yamani et al., 2004; Al-Kandari et al., 2009) and intertidal habitats
Local occurrence:
Cocconeis placentula var. euglypta is a typically benthic species, regularly observed in epiphytic assemblages
on different substrata along Kuwait’s shores (Al-Yamani & Saburova, 2011). The species massively
covered filamentous green algae and other macrophytes, but sporadically occurred in the water column
and was observed in low numbers.
Cocconeis scutellum Ehrenberg sensu lato

Plates 117: a-g, i; 118: f, g
Péragallo & Péragallo, 1897-1908: Pl. IV: 1-3, 5-7; Hendey, 1964: Pl. XXVII: 8; Hustedt, 1985: Fig. 790; Ricard, 1987:
Figs. 1152, 1153; Witkowski et al., 2000: Pls. 36: 1-7, 38: 11; Riaux-Gobin & Romero, 2003: Pls. 26-29, Figs. 42-52;
Al-Handal, 2009: Fig. 74; Al-Yamani & Saburova, 2011: Pls. 54: a-g, 177: c.
Synonymized names:
Rhaphoneis scutellum Ehrenberg 1844
Cocconeis adriatica Kützing 1844
Cocconeis mediterrana Kützing 1844
Cocconeis peruviana Kützing 1844
Cocconeis morrisii Smith 1857
Rhaphoneis marginata Grunow 1862
Cocconeis pethöi Pantocsek 1893
Cocconeis haradaae Pantocsek 1893
Cocconeis baldjikiana Schmidt 1894
Cocconeis adjuncta Schmidt 1894
Remarks:
Cocconeis scutellum is presented here in the broad sense. Several varieties are grouped, including the
nominate variety C. scutellum var. scutellum (Pl. 117: a-g, i), C. scutellum var. posidoniae (Pl. 117: h), and C.
scutellum var. parva (Pl. 118: f, g). The reason for presenting such a wide concept of this species is the lack
of clearly established differentiating criteria between these taxa to aid in routine phytoplankton analysis.

BACILLARIOPHYCEAE
Plate 118. Cocconeis spp., Anorthoneis excentrica, Lyrella lyroides, and Mastogloia rostrata.
a-d – Cocconeis placentula var. euglypta: a – live cell in rapheless valve view; b, c – acid cleaned frustules in
rapheless valve view; d – acid cleaned frustule in raphe valve view; e – Cocconeis neothumensis var. marina:
– acid cleaned frustule in rapheless valve view; f, g – Cocconeis scutellum var. parva: acid cleaned frustules
in rapheless valve view; h – Anorthoneis excentrica: acid cleaned frustule in raphe valve view; i-k – Lyrella
lyroides: i – live cell in valve view; j – acid cleaned frustule in valve view; k – external valve view; l –
Mastogloia rostrata: acid cleaned frustule in valve view.
LM, BF illumination (a, i), DIC optic (j), and phase contrast (b-h); SEM (k); scale bars: a-h, k, l – 10 μm; i,
j – 20 μm.
Valves are broadly elliptical, 18-32 μm long, 12-17 μm broad. On raphe valve, raphe is straight; axial area
is linear, narrow; transapical striae are radial, 12-14 in 10 μm, marginal areolae are distinctly larger than
the others, separated by narrow hyaline line, which is parallel to margin. On rapheless valve, sternum is
narrow, linear; valve surface is coarsely areolated, areolae are rectangular, arranged in wavy longitudinal
rows; transapical striae are radiate, 8-9 in 10 μm. Two plate-like irregularly shaped chloroplasts are arranged
along valves on either side of the central nucleus.
Distribution:
Cocconeis scutellum is a cosmopolitan epiphytic species of marine and brackish water coasts. It has been
previously reported from the Indian Ocea (Simonsen, 1974), from the bottom sediments in the Shatt Al-
Arab Estuary, Iraq (Al-Handal, 2009), and from Kuwait’s shores (Al-Yamani & Saburova, 2011).
Local occurrence:
Cocconeis scutellum is a typically benthic species, regularly observed in epiphytic assemblages on different
substrata along Kuwait’s shores (Al-Yamani & Saburova, 2011). The species abundantly covered
filamentous green algae and other macrophytes, but sporadically occurred in the water column and was
observed in low numbers.
Order Lyrellales Mann

Family Lyrellaceae Mann
Genus Lyrella Karajeva [Karaeva]
Lyrella abrupta (Gregory) Mann

Plate 119: a-d
Péragallo & Péragallo, 1897-1908: Pl. XXI: 35-39 (as Navicula abrupta); Hendey, 1964: Pl. XXXIII: 12 (as Navicula
abrupta); Hendey, 1970: p. 135 (as Navicula abrupta); Simonsen, 1974: p. 41 (as Navicula abrupta); Witkowski et al., 2000:
Pls. 95: 4, 5, 99: 8; Al-Handal, 2009: Fig. 115; Al-Kandari et al., 2009: p. 89-90, Pl. 50: B; Al-Yamani & Saburova, 2011:
Pls. 30: a-f, 178: d.
Synonymized names:
Navicula lyra var. abrupta Gregory 1857
Navicula abrupta (Gregory) Donkin 1870
Cells are solitary. Valves are elliptic-lanceolate with rounded apices, 29-46 μm long, 14-21 μm broad. Raphe
is straight, axial area is narrow, wider at the middle distance, constricted near the central nodule, central area
is transversely extended, connected to lateral extensions to form lyre-shaped hyaline area. Lateral extensions
are narrow, crescent, constricted in the middle of the valve, widening at the middle distance, narrowing
toward apices, terminating before valve apices. Valve surface is furnished with striae, interrupted by lyre-
shaped area. Transapical striae are punctate, parallel in the middle, radiate toward apices, 12-14 in 10 μm.
The cytoplasm contains two large butterfly-shaped chloroplasts, one lying against each valve.
Distribution:
Lyrella abrupta is a common marine littoral species, occasionally occurs in plankton. The species has been
previously reported (as Navicula abrupta) from the Indian Ocean (Simonsen, 1974) and from Kuwait’s
shores (Hendey, 1970), coastal waters (Al-Kandari et al., 2009, as Lyrella cf. abrupta), and intertidal
sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Lyrella abrupta is a typically benthic species, regularly observed in the intertidal sediments along Kuwait’s
shores (Al-Yamani & Saburova, 2011), but admixtured in the phytoplankton samples, sporadically found
in Kuwait’s waters and observed in low concentrations.

Lyrella lyroides (Hendey) Mann
Plate 118: i-k
Hendey, 1957: Pl. V: 3; Hendey, 1964: Pl. XXXIII: 3, 4 (as Navicula lyroides); Al-Yamani & Saburova, 2011: Pl. 35: a-f.
Synonymized names:
Navicula elliptica Smith 1853 pro parte
Navicula lyra var. elliptica Schmidt 1874
Navicula lyra var. intermedia Péragallo 1908
Navicula lyroides Hendey 1958
Cells are solitary. Valves are elliptically lanceolate with bluntly rounded apices, 54-96 μm long, 29-49 μm
broad. Raphe is straight, axial area is narrowly linear; central area is rectangular, transversely extended,
connected to lateral extensions to form lyre-shaped hyaline area. Lateral extensions are narrow, slightly
constricted in the middle of the valve, gradually narrowing toward apices, reaching the valve apices. Valve
surface is furnished with striae that interrupted by lyre-shaped area. Transapical striae are coarsely punctate,
13-15 puncta in 10 μm, slightly radiate in the marginal part, 7-10 in 10 μm. Axial striae between raphe and
lateral extension are short, commencing at the central nodule as a single punctum, and ending in the same
way at the valve apex. The cytoplasm contains two large butterfly-shaped chloroplasts with strongly lobed
edges, one lying against each valve.
Distribution:
Lyrella lyroides is a marine littoral species. It has been previously reported from Kuwait’s intertidal
Local occurrence:
Lyrella lyroides is a typically benthic species, commonly observed in the intertidal sediments along Kuwait’s
shores (Al-Yamani & Saburova, 2011), but admixtured in the phytoplankton samples, sporadically found
in Kuwait’s coastal waters, and observed in low numbers.
Genus Petroneis Stickle et Mann
Petroneis granulata (Bailey) Mann

Plate 119: j, k
Péragallo & Péragallo, 1897-1908: Pl. XXVII: 2-5 (as Navicula granulata); Hendey, 1964: Pl. XXXI: 6 (as Navicula
granulata); Hendey, 1970: p. 133 (as Navicula granulata); Foged, 1984: Pl. XLVII: 1-3 (as Navicula brasilensis); Podzorski
& Håkansson, 1987: Pl. 33: 14 (as Navicula granulata); Ricard, 1987: Fig. 726 (as Navicula granulata); Witkowski et al.,
BACILLARIOPHYCEAE
2000: Pl. 97: 1, 2; Jones et al., 2005: Fig. 10; Al-Handal, 2009: Fig. 110; Al-Yamani & Saburova, 2011: Pl. 37: a-f.
Synonymized names:
Navicula granulata Bailey 1854
Navicula polysticta Greville 1859
Navicula brasiliensis Grunow 1863
Navicula baileyana Schmidt 1874
Cells are solitary. Valves are broadly lanceolate with obtusely rounded apices, 69-83 μm long, 32-35 μm
broad. Raphe is straight, axial area is narrow, bilanceolate, wider at the middle distance, central area is
rounded to transversely elliptical. Transapical striae are distinctly punctate in the marginal area, parallel
in the middle, radiate towards the apices, 11 in 10 μm. Puncta are rounded in shape, closer towards the
valve margin, but a little scattered towards the raphe. The cytoplasm contains two large butterfly-shaped
chloroplasts, one lying against each valve. Nucleus is rounded, centrally located.
Distribution:
Petroneis granulata is principally a marine littoral species, but is frequently found in the plankton. The
species has been previously reported from the bottom sediments in the Shatt Al-Arab Estuary, Iraq (Al-
Handal, 2009), and from Kuwait’s shores (Hendey, 1970, as Navicula granulata) and intertidal sediments
Local occurrence:
Petroneis granulata is a typically benthic species, commonly observed in the intertidal sediments along
Kuwait’s shores (Al-Yamani & Saburova, 2011), but occurred rarely in Kuwait’s waters as well, being
recorded from less than 1% of all collected samples in low abundance.
Petroneis marina (Ralfs) Mann

Plate 119: l-n
Péragallo & Péragallo, 1897-1908: Pl. XXVII: 10 (as Navicula marina); Hendey, 1964: Pl. XXXI: 1-3 (as Navicula marina);
Round et al., 1990: p. 674; Witkowski et al., 2000: Pl. 102: 1; Jones et al., 2005: Figs. 4-6, 16, 18, 20, 24, 29, 39, 40;
Al-Handal, 2009: Fig. 123; Al-Yamani & Saburova, 2011: Pl. 38: a-f.
Synonymized names:
Navicula punctulata Smith 1853
Navicula marina Ralfs 1861
Cells are solitary. Valves are elliptically lanceolate with obtusely rounded apices, 35-74 μm long, 19-32 μm
broad. Raphe is straight, with T-shaped central endings, axial area is area narrow, linear, central area is
rounded. Transapical striae are distinctly punctate, radiate throughout, 10-11 in 10 μm. The cytoplasm
contains two large butterfly-shaped chloroplasts, one lying against each valve. Nucleus is rounded, centrally
located.
Distribution:
Petroneis marina is principally a marine littoral species. It has been previously reported from the bottom
sediments in the Shatt Al-Arab Estuary, Iraq (Al-Handal, 2009), and from Kuwait’s intertidal sandflats
Local occurrence:
Petroneis marina is a typically benthic species, commonly observed in the intertidal sediments along
Kuwait’s shores (Al-Yamani & Saburova, 2011), but occurred sporadically in Kuwait’s coastal waters as
well.
Order Mastogloiales Mann

Family Mastogloiaceae Mereschkowsky
Genus Mastogloia Thwaites ex Smith
Mastogloia rostrata (Wallich) Hustedt

Plate 118: l
Simonsen, 1974: Pl. 26: 1; Gibson & Stephens, 1985: Pls. 1-3; Lobban et al., 2012: Pl. 36: 9, 10.
Synonymized names:
Stigmaphora rostrata Wallich 1860
Valves are linear-lanceolate with extremely long protracted apices, 63 μm long and 10 μm broad. Raphe is
straight, axial area is very narrow, central area is unresolved. Partectal ring is displaced interiorly toward

the middle line of the valve by a siliceous flange, with a pair of centrally located bead-shaped partecta on
each side of the valve. Apices bear small septa. Valve surface is finely striated. Transapical striae are parallel,
delicate, 27-29 in 10 μm.
Distribution:
Mastogloia rostrata is one of two the only known pelagic species of the principally benthic genus Mastogloia.
This species is widely distributed in tropical plankton. It has been previously reported from the Indian
Ocean (Simonsen, 1974).
Local occurrence:
A few frustules of Mastogloia rostrata were found in plankton net hauls from Kuwait’s southern inshore
waters. The species is reported for the first time from Kuwait.
Order Naviculales Bessey

Suborder Diploneidineae Mann
Family Diploneidaceae Mann
Genus Diploneis Ehrenberg ex Cleve
Diploneis weissflogii (Schmidt) Cleve

Plate 119: g-i
Péragallo & Péragallo, 1897-1908: Pl. XIX: 11; Allen & Cupp, 1935: Fig. 100 (as Navicula weissflogii); Hendey, 1970: p.
141; Foged, 1984: Pl. XL: 7, 8; Hustedt, 1985: Fig. 1085; Podzorski & Håkansson, 1987: Pl. 21: 6, 7; Witkowski et al.,
2000: Pls. 92: 4, 5, 94: 12, 13; Hein et al., 2008: Fig. 27: 9; Al-Kandari et al., 2009: p. 103, Pl. 56: E-G; Al-Yamani &
Saburova, 2011: Pl. 66: e, f
Synonymized names:
Navicula diversa Greville 1863
Navicula weissflogii Schmidt 1873
Cells are solitary, characteristically panduriform. Valves are linear-elliptical, with rounded apices, deeply
constricted in the middle, 35-41 μm long and 17-19 μm broad. Central nodule is small, apically elongated,
constricted in the middle due to penetration of middle chambers. Raphe sternum furrows are robust are,
close together and parallel. Longitudinal canals are linear, slightly convex towards the outside. Valve surface
is coarsely striated. Transapical striae are robust, radiate, 7.5-9.5 in 10 μm, crossed by numerous longitudinal
slightly wavy ribs. The cytoplasm contains two large plate-like irregularly shaped chloroplasts, one lying
against each valve.
BACILLARIOPHYCEAE
Distribution:
Diploneis weissflogii is principally a littoral species, but is frequently found in the plankton worldwide.
Species has been previously reported from Kuwait’s shores (Hendey, 1970), coastal waters (Al-Yamani
et al., 2004; Al-Kandari et al., 2009), and intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Diploneis weissflogii was commonly observed in Kuwait’s marine environment in both epipelic and neritic
habitats. The species was frequently recorded in the intertidal sediments as well as frequently occurred
across Kuwait’s waters, being recorded from 34% of all collected samples. The species was present
throughout the year in low abundance with higher concentrations (0.9-1.0∙103 cells/l) recorded in the
southern waters during the period from December to April.
Suborder Naviculineae Hendey
Family Naviculaceae Kützing
Genus Caloneis Cleve
Caloneis elongata (Grunow) Boyer

Plate 119: e, f
Hendey, 1970: p. 151; Simonsen, 1974: Pl. 26: 5; Witkowski et al., 2000: Pl. 152: 10; Al-Yamani & Saburova, 2011: Pl.
65: g-i.
Synonymized names:
Navicula elongata Grunow 1874
Caloneis liber var. elongata (Grunow) Cleve 1894
Cells are solitary. Valves are linear with rounded apices, 98-104 μm long, 10 μm broad. Raphe is straight, with
external central endings hooked in one side. Central area is small, apically elongate, with lunate markings on
either side of the central nodule. Transapical striae are parallel, 15 in 10 μm. Two large plate-like chloroplasts
are arranged along girdle on either side of the central nucleus.
Distribution:
Caloneis elongata is mainly a littoral cosmopolitan species, but is sporadically found in the plankton. It
has been previously reported from the Indian Ocean and the Arabian Gulf (Simonsen, 1974), and from
Kuwait’s shores (Hendey, 1970) and intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Caloneis elongata occurred occasionally across Kuwait’s waters, being recorded from 6.6% of all
collected samples. The species was present throughout the year usually in low abundance with higher
concentrations (0.4-1.0∙104 cells/l) recorded in the southern inshore waters in November.
Genus Gyrosigma Hassall
The species belonging to the genus Gyrosigma are characterized by linear or lanceolate and usually sigmoid
shape of valve. Valve surface is more or less finely striated. The striae are comprised of punctate areolae and
form rows that are perpendicular and parallel to the apical and transapical axes (Cupp, 1943; Hendey, 1964).
Gyrosigma balticum (Ehrenberg) Rabenhorst

Plate 120
Péragallo & Péragallo, 1897-1908: Pl. XXXIV: 8-12 (as Pleurosigma balticum); Hendey, 1964: Pl. XXXV: 9; Hendey,
1970: p. 153; Ricard, 1977: Pl. 5: 12; Foged, 1984: Pl. XL: 3; Hustedt, 1985: Fig. 331; Ricard, 1987: Figs. 797-802;
Sterrenburg et al., 2000: Figs. 1, 13-18; Reid & Williams, 2003: Figs. 45-50; Al-Yamani & Saburova, 2011: Pl. 97: a-c.
Synonymized names:
Navicula baltica Ehrenberg 1838
Pleurosigma balticum (Ehrenberg) Smith 1852
Cells are long, solitary. Valves are linear, with parallel, sometimes slightly wavy valve margins and oblique
rounded apices, gently sigmoid towards apices, 146-203 μm long and 22-24 μm broad. Raphe curvature is
similar to that of the valves. Raphe is central in the middle, slightly eccentric towards the apices, sometimes
curved or undulating as it approaches the central nodule, with external central raphe fissures deflected in
opposite directions. Central nodule is small, elliptical, encircled by distinct and obliquely rounded central
area. The striae are arranged in transapical and longitudinal lines, equidistant, 13-15 in 10 μm. Two large
deeply lobed plate-like chloroplasts are lying along valves.

BACILLARIOPHYCEAE
Plate 119. Lyrella abrupta, Caloneis elongata, Diploneis weissflogii, and Petroneis spp.
a-d – Lyrella abrupta: a – live cell in valve view; b, c – acid cleaned frustules in valve view; d – external
valve view; e, f – Caloneis elongata: e – live cell in valve view; f – part of acid cleaned frustule in valve
view; g-i – Diploneis weissflogii: g – live cell in valve view; h, i – acid cleaned frustules in valve view; j,
k – Petroneis granulata: j – live cell in valve view; k – acid cleaned frustules in valve view; l-n – Petroneis
marina: l, m – acid cleaned frustules in valve view; n – live cell in valve view.
LM, BF illumination (a, e, g, j, n) and phase contrast (b, c, f, h, k-m); SEM (d, i); scale bars: a-h, j-n –
10 μm; i – 5 μm.
Plate 120. Gyrosigma balticum.
a, b – live cells in valve view; c – acid cleaned frustule in valve view; d-f, k, l – detailed parts of acid cleaned
frustules in valve view; g, h – detailed central areas of valves; i, j – detailed valve ends.
LM, BF illumination (a-c) and phase contrast (d-j); scale bars: a-c, f, k, l – 20 μm; d, e, g-j – 10 μm.

Distribution:
Gyrosigma balticum is a marine to brackish water littoral species widely distributed mainly in temperate
areas, but is frequently found in the plankton. The species has been previously reported from Kuwait’s
Mina Al-Ahmadi shore by Hendey (1970) and was commonly observed in the intertidal mudflats (Al-
Local occurrence:
Gyrosigma balticum occurred occasionally across Kuwait’s waters, being recorded from 10% of all collected
(160-400 cells/l) recorded in the inshore waters during the warm season from June to October.
Gyrosigma tenuissimum (Smith) Griffith et Henfrey

Plate 123: a-d
Sterrenburg et al., 2014a: Figs. 1-23.
Synonymized names:
Pleurosigma tenuissimum Smith 1853
Gyrosigma tenuissimum var. genuina Cleve-Euler 1952
Cells are long, solitary. Valves are delicate, 62-115 μm long and 5.4-6.3 μm broad, parallel-sided and straight
for most of their length, becoming slightly sigmoid near the acute apex. Raphe is nearly straight over most
of its length, curving only near apices, with minute central nodule. The density of transverse striae is 21-23
in 10 μm. Two long plate-like chloroplasts are lying along valves.
Distribution:
Gyrosigma tenuissimum is a marine littoral widely distributed species (Sterrenburg et al., 2014a).
Local occurrence:
Gyrosigma tenuissimum is a typically benthic species, but occurred sporadically in Kuwait’s coastal waters
as well. The species is reported for the first time from Kuwait.
Genus Haslea Simonsen
The genus Haslea encompasses the diatoms with weakly silicified frustules, usually spindle-shaped with acute
ends and convex sides. Raphe is straight, with small, approximate central pores and little developed apical
pores. Axial and central areas are inconspicuous. The structure of the valve consists of transapical, parallel,
and straight apical rows of areolae (Simonsen, 1974). In a recent publication, definition of the genus Haslea
BACILLARIOPHYCEAE
was expanded significantly, and a range of new species were described or transferred to the genus (Massé et
al., 2001; Poulin et al., 2004; Sterrenburg et al., 2015a, b). Haslea species are characterized by a sandwich-
structured valve consisting of an internal grate-like basal layer and an external tegumental layer perforated
by continuous longitudinal fissures. These two layers are shored by longitudinal bulkhead-like structures
(termed as ‘saepes’) that are perforated so that they form fence-like rows of columns (Sterrenburg et al.,
2015a).
Haslea gigantea (Hustedt) Simonsen

Plate 121
Simonsen, 1974: Pl. 31: 1
Synonymized names:
Navicula gigantea Hustedt 1961
Navicula cf vitrea Schrader 1973
Cells are large, solitary. Valves are weakly silicified, lanceolate with acute apices, 360-395 μm long and 30-
40 μm broad. Raphe is straight, with extremely approximate central external endings depressed as two tiny
pits. Terminal raphe fissures are straight. Central and axial areas are inconspicuous. Valve surface is very
finely striated. Transapical and longitudinal striae are crossed at right angles. Transapical striae are parallel,
16-17 in 10 μm, longitudinal striae are more dense, 23-25 in 10 μm. The cytoplasm contains numerous small
elongate chloroplasts. The nucleus is centrally located. Light-yellow oil droplets are often accumulated at
the apices.
Distribution:
Haslea gigantea is a marine plankton warm water species with preference for more neritic habitats
(Simonsen, 1974). The species has been previously reported from the Indian Ocean and the Arabian
Gulf (Simonsen, 1974).
Local occurrence:
Haslea gigantea is of low occurrence in Kuwait’s waters, being recorded from 2% of all collected samples,
most abundantly (240-320 cells/l) recorded in the inshore southern waters during the winter in December.
This species is considered as a new record for Kuwait’s diatom flora.
Haslea howeana (Hagelstein) Giffen

Plate 122: a-e
Witkowski et al., 2000: Pl. 148: 12, 13; Al-Yamani & Saburova, 2011: Pl. 76: c-f; Lobban et al., 2012: Pl. 47: 6, 7.
Synonymized names:
Navicula howeana Hagelstein 1938
Cells are solitary. Valves are lanceolate, with acute apices, 39-60 μm long and 7-12 μm broad. Raphe is
straight, axial and central areas are inconspicuous. External central raphe endings are approximate, crook-
shaped. Terminate raphe fissures are curved.Valve surface is striated with transapical and longitudinal striae
crossed at right angles. Transapical striae are weakly radiationg at the center and almost parallel distally, 16-
19 in 10 μm, crossed by longitudinal striae, 18-20 in 10 μm. Two plate-like chloroplasts are lying along the
girdle on either side of the central nucleus.
Distribution:
Haslea howeana is a marine littoral warm water species. It has been previously reported from Kuwait’s
intertidal sandflats (Al-Yamani & Saburova, 2011).
Local occurrence:
Haslea howeana is a typically benthic species, regularly observed in epipelic assemblages Kuwait’s shores
(Al-Yamani & Saburova, 2011), but sporadically occurred in the water column, and was observed in low
numbers.
Plate 121. Haslea gigantea.

a-c – live cells in valve view; d – dividing cell in girdle view; e – detail of valve showing perpendicular
striation; f – detail of collapsed frustule in valve view; g – detail of collapsed frustule showing external
longitudinally striated tegumental layer and internal grate-like layer; h – detailed central area of external
valve surface showing proximal raphe endings; i – apex in external view showing terminal raphe fissure.
LM, DIC optic (a-e); SEM (f-i); scale bars: a-d – 50 μm; e – 25 μm; f – 5 μm; g-i – 2 μm.

BACILLARIOPHYCEAE
Haslea cf. meteorou Hinz et Sterrenburg
Plate 122: f-h
Sterrenburg et al., 2015a: Figs. 16, 17, 39-46.
Cells are solitary. Valves are lanceolate, with acute and slightly produced apices, 27-29 μm long and 5-6
μm broad. Raphe is straight, axial and central areas are inconspicuous. External central raphe endings are
approximate, crook-shaped. Internally, central raphe node is tilted sideways. Terminate raphe fissures are
curved. Valve surface is striated with transapical and longitudinal striae crossed at right angles. Transapical
striae are weakly radiate, 20.5-23 in 10 μm, crossed by longitudinal striae, 18-20 in 10 μm.
Distribution:
Haslea meteorou is a marine plankton species described recently from the sample of the “Meteor”
expedition (1964-1965) collected from the Arabian Sea (Sterrenburg et al., 2015a).
Remarks:
Specimens found in Kuwait’s waters are highly similar to Haslea meteorou including the crook-shaped
central external raphe endings but smaller, 27-29 μm long versus 38-45 μm (Sterrenburg et al., 2015a) and
more densily striated. Due to unknown range of morphological variability within the recently described
Haslea meteorou, the smaller specimens isolated from Kuwait’s waters are included as belonging to that
taxon with caution, until the morphological variability of this species is examined in details.
Local occurrence:
The illustrated specimens were isolated from Kuwait’s southern inshore waters in the spring. The species
is reported for the first time from Kuwait.
Haslea wawrikae (Hustedt) Simonsen emend. Sterrenburg et Hinz

Plate 122: i, j
Hasle & Syvertsen, 1997: Pl. 62; Sterrenburg et al., 2015b: Figs. 1-11.
Synonymized names:
Navicula wawrikae Hustedt 1961
Cells are long, solitary. Valves are filiform with almost parallel and gradually tapering ends with long hair-like
extensions, 350-465 μm long and 5-7 μm broad. Raphe is straight, central and axial areas are inconspicuous.
Transapical and longitudinal striae are crossed at right angles. Transapical striae are parallel, 27.5-29 in
10 μm, longitudinal striae are much more dense, 42-47 in 10 μm. The cytoplasm contains plate-shaped
chloroplasts.
Distribution:
Haslea wawrikae is a marine plankton warm water species (Hasle & Syvertsen, 1997).
Local occurrence:
Haslea wawrikae rarely occurred in Kuwait’s waters and was observed in low abundances. This species is
considered as a new record for Kuwait’s diatom flora.
Plate 122. Haslea spp.

a-e – Haslea howeana: a – live cell in valve view; b – acid cleaned frustule in valve view; c – frustule in oblique
view; d – detailed central area of external valve surface showing proximal crook-shaped raphe endings; e
– apex in external view showing distal raphe ending; f-h – Haslea cf. meteorou: f – frustule in valve view; g –
collapsed frustule showing internal (left) and external (right) valve view; h – detailed central area showing
sideways tilted central raphe node in internal view (left) and proximal crook-shaped raphe endings in external
view (right); i, j – Haslea wawrikae: i – Lugol-fixed specimen; j – detail of collapsed frustule showing external
longitudinally striated tegumental layer and internal grate-like layer.
LM, DIC optic (a), BF illumination (i), and phase contrast (b); SEM (c-h, j); scale bars: i – 50 μm; a, b – 10
μm; c, f, g – 5 μm; d, e, h, j – 2 μm.

BACILLARIOPHYCEAE
Genus Mastoneis Cleve
Mastoneis biformis (Grunow) Cleve

Plate 123: e-h
Péragallo & Péragallo, 1897-1908: Pl. VII: 5; Hendey, 1970: Pl. 6: 71; Witkowski et al., 2000: Pl. 101: 5; Hein et al.,
2008: Figs. 32: 3, 49: 1, 2; Al-Yamani & Saburova, 2011: Pl. 176: a-h.
Synonymized names:
Stauroneis biformis Grunow 1863
Navicula biformis (Cleve) Mann 1925
Cells are solitary. Valves are elliptic-oblong with rostrate, acutely rounded apices, 73-90 μm long and 25-
30 μm broad. Raphe is undulating, axial area is linear, very narrow, central area is in form of transversely
expanded short fascia. Valve surface is finely punctate, 17-18 puncta in 10 μm. Transapical striae are radiate
throughout, 19-20 in 10 μm. Marginal area of valve is furnished with numerous costae directed from margins
to about half of the distance to the raphe. Costae differ in length, 7.5-8.5 in 10 μm. Two large golden-brown
chloroplasts are arranged along apical axis on either side of the central nucleus.
Distribution:
Mastoneis biformis is principally a littoral species, but is sometimes found in the plankton. It has been
previously reported from Kuwait’s shores (Hendey, 1970) and intertidal sediments (Al-Yamani &
Saburova, 2011).
Local occurrence:
Mastoneis biformis rarely occurred across Kuwait’s waters, being recorded from less that 1% of all collected
samples in low abundances.
Genus Navicula Bory
Navicula directa (Smith) Ralfs

Plate 123: i-k
Péragallo & Péragallo, 1897-1908: Pl. XII: 6; Hendey, 1964: p. 202; Hendey, 1970: p. 132; Witkowski et al., 2000: Pls.
129: 1, 133: 10-12; Al-Handal, 2009: Fig. 104; Al-Kandari et al., 2009: p. 105, Pl. 57: C; Al-Yamani & Saburova, 2011:
Pl. 82: a-e.
Synonymized names:
Pinnularia directa Smith 1853
Cells are long, solitary. Valves are linear-lanceolate, with sub-acute apices, 67-89 μm long, 7-9 μm broad.
Raphe is straight, with central raphe endings expanded into pores. Axial area is very narrow, linear, central
area is small, slightly asymmetrical. Transapical striae are robust, parallel, 7-9 in 10 μm. Two long plate-like
chloroplasts are extending the entire length of the cell, appressed to the girdle bands. Nucleus is centrally
located.
Distribution:
Navicula directa is a marine cosmopolitan mainly littoral species, but is sporadically found in the plankton.
The species has been previously reported from the bottom sediments in the Shatt Al-Arab Estuary, Iraq
(Al-Handal, 2009), and from Kuwait’s shores (Hendey, 1970), coastal waters (Al-Yamani et al., 2004;
Al-Kandari et al., 2009), and intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Navicula directa rarely occurred in Kuwait’s waters, being recorded from less than 1% of all collected
samples in low abundance.

Navicula palpebralis Brébisson ex Smith
Plate 123: p-n
Hendey, 1964: Pl. XXXIV: 13-19; Witkowski et al., 2000: Pls. 139: 9, 140: 1-3; Al-Yamani & Saburova, 2011: Pl. 77:
e, f.
Synonymized names:
Schizonema palpebrale (Brébisson ex Smith) Kuntze 1898
Cells are solitary. Valves are elliptically lanceolate, with either broadly rounded or acute apices, 44-78 μm long
and 14-26 μm broad. Raphe is straight, with distant external central endings and apical endings hooked to
one side. Axial area is broad, widening to produce lanceolate to lanceolate-rhombic central area. Transapical
striae are arranged in sub-marginal band occupying about half or sometimes less than half the area between
the raphe and the valve margin, radiate, 11 in 10 μm. Two long plate-like chloroplasts are extending the
entire length of the cell, appressed to the girdle bands. Nucleus is centrally located.
Distribution:
Navicula palpebralis is a marine to brackish water widely distributed species inhabiting sandy sediments,
but is sporadically found in the plankton. It has been previously reported from Kuwait’s shores (Hendey,
1970) and intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Navicula palpebralis rarely occurred in Kuwait’s waters, being recorded from less than 1% of all collected
samples with low abundance.
Genus Seminavis Mann
Seminavis robusta Danielidis et Mann

Plate 161: h, i
Danielidis & Mann, 2002: Figs. 39-53; Hein et al., 2008: Fig. 57: 3, 5; Al-Yamani & Saburova, 2011: Pl. 89: a-g.
Synonymized names:
Amphora macilenta Gregory 1857
Cells are solitary. Frustules are rhombic-lanceolate with obtusely rounded apices. Valves are semilanceolate,
with straight or very weakly convex ventral margin and convex dorsal margin, 45-49 μm long and 8-11 μm
broad. Raphe is straight, approximate ventral margin, with branches deflected ventrally at the center. Axial
BACILLARIOPHYCEAE
area is more expressed on dorsal valve side, with central roundish expansion on ventral side. Transapical
striae on the dorsal side are radiate, 17-19.5 in 10 μm. Ventral striae are distinctly radiate in the middle,
becoming parallel towards apices, 16-18 in 10 μm. A single plate-like chloroplast is appressed to the dorsal
side.
Distribution:
Seminavis robusta is a marine littoral species. It has been previously reported from Kuwait’s intertidal
Local occurrence:
Seminavis robusta rarely occurred in Kuwait’s waters and found in low numbers.
Plate 123. Gyrosigma tenuissimum, Mastoneis biformis, and Navicula spp.
a-d – Gyrosigma tenuissimum: a – live cell in valve view; b-d – acid cleaned frustules in valve
view; e-h – Mastoneis biformis: e – live cell in valve view; f – live cell in girdle view; g, h – acid
cleaned frustules in valve view; i-k – Navicula directa: i – live cell in valve view; j, k – acid cleaned
frustules in valve view; l, m – Navicula spp.: live cells in valve view; n-p – Navicula palpebralis:
n – live cell in valve view; o, p – acid cleaned frustules in valve view.
LM, BF illumination (a, i, l-n), DIC optic (e, f, j, o), and phase contrast (b-d, g, h, k, p); scale
bars: a-c, e-k, n-p – 10 μm; d, l, m – 5 μm.

Genus Trachyneis Cleve
The species belonging to the genus Trachyneis are characterized by lanceolate, elliptic-lanceolate or elliptical
valves. Axial area is narrow, raphe is straight, central area is dilated to form an expanding stauros which may
or may not reach the valve margin. Valve surface is coarsely areolated, with an outer lamina bearing fine
perforations superimposed upon an inner loculate layer (Hendey, 1964).
Trachyneis antillarum (Cleve et Grunow) Cleve

Plate 124: a-c, k
Hendey, 1970: Pl. 4: 44; Simonsen, 1974: p. 43; Hustedt, 1985: Fig. T-18; Al-Handal, 2009: Fig. 190; Al-Kandari et al.,
2009: p. 106, Pl. 58: A-E; Al-Yamani & Saburova, 2011: Pl. 90: a-g.
Synonymized names:
Alloioneis antillarum Cleve et Grunow 1878
Cells are solitary. Frustules are rectangular in girdle view, with rounded corners and slightly convex margins.
Valves are broadly lanceolate to rhombic-lanceolate in outline, 67-139 μm long and 25-30 μm broad. Valve
face is strongly arched, median line is somewhat eccentric. Raphe is straight or slightly bent, with apical
endings hooked in the same side. Axial area is broad, unilateral. Central area is asymmetrical, rounded
and unilateral on the opposite side of the axial area. Valve surface is coarsely alveolate. Rectangular alveoli
form longitudinal irregularly oblique rows. Transverse rows are radiate, 9-11 in 10 μm. Two large plate-like
chloroplasts with slightly lobed edges are arranged along girdle on either side of the central transversely
elongated nucleus.
Distribution:
Trachyneis antillarum is mainly a littoral tropical species, but is frequently found in the plankton. It has
been previously reported from the Arabian Gulf (Simonsen, 1974), from the bottom sediments in the
Shatt Al-Arab Estuary, Iraq (Al-Handal, 2009), and from Kuwait’s shores (Hendey, 1970), coastal waters
(Al-Kandari et al., 2009), and intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Trachyneis antillarum occurred commonly in Kuwait’s waters, being recorded from 57% of all collected
concentrations (1.8-8.0∙103 cells/l) recorded in the inshore waters of Kuwait Bay during the period from
October to January.
Trachyneis aspera (Ehrenberg) Cleve
BACILLARIOPHYCEAE
Plate 124: d-g, l
Péragallo & Péragallo, 1897-1908: Pl. XXIX: 1-7; Hendey, 1964: Pl. XXIX: 11-13 (Trachyneis aspera var.); Hendey,
1970: p. 148; Simonsen, 1974: p. 43; Ricard, 1977: Pl. 4: 7; Foged, 1984: Pl. LI: 3, 4, 6, 7; Hustedt, 1985: Figs. T-1-T-12
(Trachyneis aspera var.); Podzorski & Håkansson, 1987: Pl. 40: 6; Ricard, 1987: Figs. 834-839; Witkowski et al., 2000: Pl.
159: 1-6, 9; Hein et al., 2008: Fig. 55: 3; Al-Handal, 2009: Figs. 194, 195; Al-Kandari et al., 2009: p. 107, Pl. 58: F-H;
Al-Yamani & Saburova, 2011: Pl. 91: a-h.
Synonymized names:
Navicula aspera Ehrenberg 1840
Stauroptera aspera Ehrenberg 1841
Stauroneis aspera (Ehrenberg) Kützing 1844
Stauroneis achnanthes (Ehrenberg) Kützing 1844
Stauroneis pulchella Smith 1853
Pinnularia aspera (Ehrenberg) Ehrenberg 1854
Navicula stauroneiformis Leuduger-Fortmorel 1892
Navicula aspera var. genuina (Cleve) Fricke 1902
Cells are solitary. Frustules are rectangular in girdle view, with rounded corners. Valves are linear to lanceolate,
with rounded apices, 54-101 μm long and 14-17 μm broad. Valve face is arched, slightly depressed in the
middle. Raphe is straight, with apical endings hooked in the same side. Axial area is narrow, central area is
in form of stauros, widened outwards and truncated before reaching the valve margins. Transapical striae
are composed of rectangular alveoli, arranged in transverse rows, radiate, 8-13 in 10 μm. Two large plate-
like chloroplasts with slightly lobed edges are arranged along girdle on either side of the central transversely
elongated nucleus.
Distribution:
Trachyneis aspera is a cosmopolitan marine littoral species. It has been previously reported from neritic
habitats of the northern Arabian Gulf (Simonsen, 1974), from the bottom sediments in the Shatt Al-
Arab Estuary, Iraq (Al-Handal, 2009), and from Kuwait’s shores (Hendey, 1970), coastal waters (Al-
Kandari et al., 2009), and intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Trachyneis aspera is a typically benthic species, regularly observed in epipelic assemblages Kuwait’s shores
(Al-Yamani & Saburova, 2011), but sporadically occurred in the water column, and was observed in low
numbers.
Trachyneis debyi (Leudiger-Fortmorel) Cleve

Plate 124: h-j, m
Hendey, 1970: p. 149; Simonsen, 1974: Pls. 27, 28: 1; Hustedt, 1985: Fig. T-15; Al-Handal, 2009: Figs. 191, 193; Al-
Yamani & Saburova, 2011: Pl. 93: a-j.
Synonymized names:
Alloioneis debyi Leudiger-Fortmorel 1892
Trachyneis debyi var. osculifera Cleve 1894
Cells are slender, solitary. Frustules are rectangular in girdle view, with rounded corners and slightly convex
margins. Valves are linear-lanceolate with rounded apices, 87-149 μm long and 21-28 μm broad. Valve face is
strongly arched. Axial area is narrow, linear. Central area is roundish, with two semilunate furrows on either
side of the central nodule. Valve surface is strongly alveolate. Alveoli are rectangular, arranged in transverse
and oblique rows. Transverse rows are parallel, 9-10 in 10 μm. Two large plate-like chloroplasts with lobed
edges are arranged along girdle on either side of the central transversely elongated nucleus.
Distribution:
Trachyneis debyi is mainly a littoral tropical and subtropical species, but is sporadically found in the
plankton. It has been previously reported from the coastal waters of the northern Arabian Gulf
(Simonsen, 1974), from the bottom sediments in the Shatt Al-Arab Estuary, Iraq (Al-Handal, 2009), and
from Kuwait’s shores (Hendey, 1970) and intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Trachyneis debyi occurred occasionally in Kuwait’s waters, being recorded from 9.4% of all collected
(100-200 cells/l) recorded during the warm season in April and May.
Plate 124. Trachyneis spp.

a-c, k – Trachyneis antillarum: a – live cell in valve view; b – live cell in girdle view; c – acid cleaned frustules
in valve view; k – detailed central area of valve; d-g, l – Trachyneis aspera: d – live cell in valve view; e – live
cell in girdle view; f – detailed part of cell in valve view; g – acid cleaned valve; l – detailed central area of
valve; h-j, m – Trachyneis debyi: h – live cell in valve view; i – live cell in girdle view; j – collapsed cell in girdle
view with partially detached valve; m – detailed central area of valve.
LM, BF illumination (a, b, d-f, h-j) and phase contrast (c, g, k-m); scale bars: a-c, f, g, k-m – 10 μm; d, e, h-j
– 20 μm.

BACILLARIOPHYCEAE
Family Plagiotropidaceae Mann
Genus Ephemera Paddock
Ephemera planamembranacea (Hendey) Paddock

Plate 125: a-d
Hendey, 1964: Fig. 8 (as Navicula planamembranacea); Paddock, 1988: Pl. 31: 1-8; Hasle & Syvertsen, 1997: Pl. 64.
Synonymized names:
Navicula planamembranacea Hendey 1964
Cells are solitary, weakly silicified, with pervalvar axis of 15-30 µm, apical axis of 66-90 µm, and transapical
axis of 8-10 µm. Valves are highly vaulted and flattened in transapical plane. Raphe is separating the valve
face into two unequal parts. Central nodule is slightly depressed, with a few stronger and more widely spaced
interstriae, reaching valve margin and producing the impression of a pseudostauros. No striation is visible
by LM. The cytoplasm contains numerous small elongated chloroplasts radiating from central nucleus and
lying along the raphe.
Distribution:
Ephemera planamembranacea is a marine plankton scarcely reported species known from the North
Atlantic (Hendey, 1964; Paddock, 1988; Hasle & Syvertsen, 1997).
Local occurrence:
Ephemera planamembranacea sporadically occurred in Kuwait’s waters, and was observed in low numbers.
Genus Plagiolemma Paddock
Plagiolemma confusa (Hendey) Paddock

Plate 125: e-i
Hendey, 1964: Pl. XXXVII: 15 (as Tropidoneis confusa); Paddock, 1988: Pl. 27: 1-4; Al-Yamani & Saburova, 2011: Pl.
112: e, f.
Synonymized names:
Tropidoneis confusa Hendey 1964
Cells are solitary, transapically flattened, weakly siliceous. Frustules are nearly circular in outline in girdle
view, with strongly convex margins, 50-54 μm long and 40-42 μm broad. Valves are semicircular, with a
strongly arcuate dorsal margin which is occupied by raphe. Central area is transversely dilated to form a
single narrow costa (fascia) which is nearly attained the valve margin. Transapical striae are very gentle, 28-
30 in 10 μm. A single (?) plate-like deeply four-lobed chloroplast extends along the girdle.
Distribution:
Plagiolemma confusa is a marine neritic warm water species. It has been previously reported from Kuwait’s
intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Plagiolemma confusa sporadically occurred in Kuwait’s waters, and was observed in low numbers.

BACILLARIOPHYCEAE
Plate 125. Ephemera planamembranacea and Plagiolemma confusa.

a-d – Ephemera planamembranacea: a, b – live cell in oblique view at different focal planes; c, d –
frustules in girdle view; e-i – Plagiolemma confusa: e, f – acid cleaned valves; g-i – live cell in oblique
(g), valve (h) and girdle (i) views.
LM, BF illumination (a, b, g-i) and phase contrast (c-f); scale bars: 10 μm.
Genus Meuniera Silva
Meuniera membranacea (Cleve) Silva

Plate 126
Pavillard, 1925: Fig. 109 (as Navicula membranacea); Cupp, 1943: Fig. 142 (as Navicula membranacea); Hendey, 1964: Pl.
XXI: 3 (as Stauroneis membranacea); Simonsen, 1974: p. 40 (as Stauroneis membranacea); Throndsen et al., 2007: p. 188;
Hoppenrath et al., 2009: Fig. 46 g-i.
Synonymized names:
Navicula membranacea Cleve 1897
Stauropsis membranacea (Cleve) Meunier 1910
Stauroneis membranacea (Cleve) Hustedt 1959
Cells are united in straight ribbon-shaped chains by valve face. Frustules are weakly silicified, rectangular in
girdle view, 60-75 µm long and 38-46 µm broad. Valves are narrowly elliptical in outline with pointed ends,
nearly flat or slightly concave in the center. Central nodule is dilated into a distinct narrow stauros extending
to valve margin. Four ribbon-like and folded chloroplasts are radiating from the centrally located nucleus.
Distribution:
Meuniera membranacea is a neritic, truly planktonic species of wide distribution. It has been previously
reported from the Arabian Gulf (Simonsen, 1974, as Stauroneis membranacea) and from Kuwait’s waters
Local occurrence:
Meuniera membranacea occurred frequently in Kuwait’s waters, being recorded from 34% of all collected
winter season in February.
Genus Plagiotropis Pfitzer
Plagiotropis lepidoptera (Gregory) Kuntze

Plate 127: a-f
Péragallo & Péragallo, 1897-1908: Pl. XXXIX: 1-18 (as Tropidoneis lepidoptera); Cupp, 1943: Fig. 149 (as Tropidoneis
lepidoptera); Hendey, 1964: Pl. XXXVI: 2, 4 (as Tropidoneis lepidoptera); Hendey, 1970: p. 149 (as Tropidoneis lepidoptera);
Podzorski & Håkansson, 1987: Pl. 38: 2; Ricard, 1987: Figs. 843-851; Paddock, 1988: Pl. 13: 1-11; Witkowski et al.,
2000: Pl. 174: 1, 2; Hein et al., 2008: Fig. 54: 1, 2; Al-Kandari et al., 2009: p. 110, Pl. 62: E-I; Al-Yamani & Saburova,
2011: Pls. 113: a-g, 114: a-g.
Synonymized names:
Amphiprora lepidoptera Gregory 1857
Tropidoneis lepidoptera (Gregory) Cleve 1894
Orthotropis lepidoptera (Gregory) Van Heurck 1896
Cells are solitary. Frustules are rectangular, oblong, constricted in the middle, with distinct wings projecting
equally above and below the central nodule, 59-113 μm long and 21-47 μm broad. Longitudinal fold is
strongly marked. Girdle is simple. Valves are linearly lanceolate with apiculate apices. Raphe is central,
raised upon a keel. Central area is transversely dilated, small. Transapical striae are finely punctate, parallel,
20-22 in 10 μm. Two, rarely four plate-like multi-lobed chloroplasts are arranged along the girdle.
Distribution:
Plagiotropis lepidoptera is a littoral species, but reported occasionally in the plankton. It has been previously
reported (as Tropidoneis lepidoptera) from Kuwait’s shores (Hendey, 1970), coastal waters (Al-Kandari et
al., 2009), and intertidal sediments (Al-Yamani & Saburova, 2011).

BACILLARIOPHYCEAE
Plate 126. Meuniera membranacea.

a – live cells in broad girdle view united to ribbon-like chain; b, g – detailed part of chain in broad girdle
view; c – chain in valve view; d – chain in narrow girdle view; e – single cell in broad girdle view; f – two
sibling cells in broad girdle view; h – acid cleaned sibling frustules in broad girdle view.
LM, BF illumination (a-g) and phase contrast (h); scale bars: a-g – 20 μm; h – 10 μm.
Local occurrence:
Plagiotropis lepidoptera is a typically benthic species, commonly observed in the intertidal sediments along
Kuwait’s shores (Al-Yamani & Saburova, 2011), but frequently occurred in Kuwait’s waters as well,
being recorded from 35.5% of all collected samples. The species was present throughout the year in low
to moderate abundance with higher concentrations (1.0-2.4∙103 cells/l) recorded in the inshore waters in
Kuwait Bay and the adjacent waters during the period from June to November.
Plagiotropis tayrecta Paddock

Plate 127: g-i
Paddock, 1988: Pl. 5: 1-8; Witkowski et al., 2000: Pls. 173: 1, 174: 8-11; Al-Yamani & Saburova, 2011: Pl. 115: a-h.
Synonymized names:
Amphiprora recta Gregory 1857
Plagiotropis recta (Gregory) Grunow ex Cleve et Möller 1882
Tropidoneis recta (Gregory) Cleve 1894
Cells are solitary. Frustules are rectangular in girdle view, with rounded corners, slightly constricted in the
middle, 54-61 μm long and 16-17 μm broad. Girdle is simple. Valves are asymmetrical, longitudinal fold is
indistinct. Transapical striae are very finely punctate, parallel, 24-25 in 10 μm. Two plate-like chloroplasts
are arranged along the girdle.
Distribution:
Plagiotropis tayrecta is a marine species with wide distribution, mainly in the temperate areas. It has been
previously observed in Kuwait’s marine intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Plagiotropis tayrecta is a typically benthic species, but sporadically occurred in Kuwait’s waters and was
observed in low numbers.
Family Pleurosigmataceae Mereschowsky

Genus Pleurosigma Smith
Pleurosigma is a marine or brackish water genus typical of epipelic habitats, with just a few species, and
appear more or less regularly in the plankton. This genus is closely related to Gyrosigma, but differs in plastid
structure, decussate striae and the prominent sternum on both sides of the raphe. The species belonging to
the genus Pleurosigma are characterized by lanceolate, more or less sigmoid valves, and central and usually
sigmoid raphe. Striae are arranged in decussate pattern composed of evenly spaced transapical rows and
two opposing oblique rows (Cupp, 1943; Hendey, 1964; Simonsen, 1974). The species within the genus
Pleurosigma cannot be reliably distinguished based on living specimens, and from light microscopy alone.
Pleurosigma decorum Smith

Plates 128: a-f; 130: a-c
Hendey, 1964, p. 242; Sar et al., 2013: Figs. 2-4; Sterrenburg et al., 2014b: Figs. 2, 3 b, c.
Cells are solitary. Valves are narrowly lanceolate and gently sigmoid, vaulted, with sub-acute apices, 205-
230 μm long and 25-37 μm broad. Raphe-sternum is sigmoid, eccentric, crossing the valve diagonally and
approaching opposite sides toward the ends. Central area is small, nearly circular, with bilateral dilatation
around the central nodule. External central raphe fissures are markedly projecting into the central area,
undulated toward the same side and overlapping in parallel for some distance. Striation pattern is decussate,
oblique striae are intersecting at nearly 90°, 15-17 in 10 μm, crossed by transverse striae, 16-18 in 10 μm. Two
multilobed chloroplasts extend on either side of the central nucleus.

BACILLARIOPHYCEAE
Plate 127. Plagiotropis spp.

a-f – Plagiotropis lepidoptera: a – live cell in girdle view; b – live cell in oblique valve view; c – acid cleaned
frustule in valve view; d, e – detailed part of acid cleaned frustule in girdle view; f – detail of central
nodule; g-i – Plagiotropis tayrecta: g – live cell in girdle view; h – acid cleaned frustule in girdle view; i –
detail of frustule in girdle view.
LM, DIC optic (a, b), BF illumination (g) and phase contrast (c, d, h); SEM (e, f, i); scale bars: a-e, g, h –
10 μm; i – 5 μm; f – 2 μm.
Distribution:
Pleurosigma decorum is a marine littoral species with a global distribution, but may be easily confused with
morphologically similar Pleurosigma formosum (Hendey, 1964; Sterrenburg et al., 2014b).
Local occurrence:
Pleurosigma decorum occurred sporadically in Kuwait’s waters, and was observed in low numbers. The
species is reported for the first time from Kuwait.
Pleurosigma diversestriatum Meister

Plates 128: g-k; 129
Hendey, 1970: Pl. 6: 62; Sterrenburg, 2001: Figs. 6, 15-18; Al-Handal, 2009: Fig. 153; Al-Kandari et al., 2009: p. 108,
Pls. 59: I, J, 60: A-F; Al-Yamani & Saburova, 2011: Pl. 102: a-d.
Synonymized names:
Pleurosigma aequatoriale Cleve 1878
Cells are solitary. Valves are broadly lanceolate, gently tapering to slightly rostrate apices, slightly sigmoid
at apices, 88-104 μm long and 18-22 μm broad. Raphe is median, strongly sigmoid, and eccentric near
the apices. Raphe curvature is much greater than that of valve. External terminal raphe ends are hook-
shaped, oppositely bent, internally are terminating in helictoglossae turned toward the concave margin of
the valve. Central area is distinct and circular, internally saddle-shaped, with central bars surrounding the
elongate central nodule. External central raphe endings protrude far into the central area; their shape and
type of deflection are variable. Internal central raphe endings are coaxial and slightly dilated. Valve surface
is coarsely areolated, areolae are round rather than slit-like. Striation pattern is decussate, strongly curved in
the central portion of the valve around the central nodule. Oblique striae are intersecting at nearly 60-70°,
18-20 in 10 μm, crossed by transverse striae, 24-25 in 10 μm. Two multilobed chloroplasts extend on either
side of the central nucleus.
Distribution:
Pleurosigma diversestriatum is mainly a littoral species, but is frequently found in the plankton. The species
has been previously reported from Kuwait’s shores (Hendey, 1970), coastal waters (Al-Yamani et al.,
2004; Al-Kandari et al., 2009), and intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Pleurosigma diversestriatum commonly occurred across Kuwait’s waters, being recorded from 60% of all
collected samples. The species was present throughout the year in low to moderate abundance with higher
concentrations (0.6-3.7∙104 cells/l) recorded in the inshore waters during the period from September to
January.
Plate 128. Pleurosigma spp.

a-f – Pleurosigma decorum: a – live cell in valve view; b, c – acid cleaned frustules in valve view; d – detailed
part of acid cleaned frustule in valve view; e – detailed end of cleaned valve; f – detailed central part of
cleaned valve; g-k – Pleurosigma diversestriatum: g – live cell in valve view; h – acid cleaned frustule in valve
view; i – detailed part of acid cleaned frustule in valve view; j – detailed end of cleaned valve; k – detailed
central part of cleaned valve.
LM, BF illumination (a, g) and phase contrast (b-f, h-k); scale bars: a-c – 25 μm; d-h – 10 μm; i-k – 5 μm.

BACILLARIOPHYCEAE
Plate 129. Pleurosigma diversestriatum.
a – external valve view; b – internal valve view; c – detailed central part in external valve view;
d – detailed central part in internal valve view; e – apex in external view showing terminal
fissure; f – apex in internal view; g – detail of valve in external view showing central raphe
endings; h – detail of valve in internal view showing central nodule.
SEM; scale bars: a, b – 10 μm; c-f – 5 μm; g, h – 2 μm.

BACILLARIOPHYCEAE

a-c – Pleurosigma decorum: a – frustule in external valve view; b – apex in external valve view; c – detail of
valve in external view showing central raphe endings; d-h – Pleurosigma inflatum: d, e – internal valve view;
f, g – detail of valve in internal view showing central nodule; h – apex in internal view showing terminal
fissure.
SEM; scale bars: a – 20 μm; d, e – 10 μm; b, c, f-h – 5 μm.
Pleurosigma elongatum Smith
Plate 131: a-f
Reid, 2002: Figs. 22-26; Hein et al., 2008: Fig. 55: 2; Al-Kandari et al., 2009: p. 109, Pl. 61: A-E; Al-Yamani &
Saburova, 2011: Pl. 103: a-d; Sar et al., 2014: Figs. 48-55; Al-Handal et al., 2014: Fig. 55.
Synonymized names:
Gyrosigma elongatum (Smith) Griffith et Henfrey 1855
Pleurosigma angulatum var. elongatum (Smith) Van Heurck 1885
Cells are solitary. Valves are slender, linear-lanceolate, gently tapering to acutely rounded apices, 180-212
μm long and 22-28 μm broad. Raphe is median, almost straight for most of its length, becoming slightly
sigmoid and eccentric near the apices. Central area is small, rounded. External central raphe endings are
twice deflected to the same side, becoming broadly W-shaped. Striation pattern is decussate. Oblique striae
are intersecting at nearly 60°, 18-19.5 in 10 μm, crossed by transverse striae, 22 in 10 μm. Two large folded
ribbon-shaped chloroplasts are lying in the valvar plane.
Distribution:
Pleurosigma elongatum is a marine littoral species distributed in temperate and warm water regions. The
species has been previously reported from the epiphytic assemblages in the Mesopotamian wetland,
South Iraq (Al-Handal et al., 2014), and from Kuwait’s coastal waters (Al-Kandari et al., 2009) and
Local occurrence:
Pleurosigma elongatum sporadically occurs in Kuwait’s waters in low numbers.
Pleurosigma falx Mann

Plate 131: g-k
Mann, 1925: Pl. 30: 1; Stidolph, 2002: Pl. 3: 1-4; Al-Yamani & Saburova, 2011: Pl. 104: a-d.
Cells are solitary. Valves are narrowly lanceolate, sigmoid, gently tapering to acutely rounded apices, 128-
143 μm long and 16-18 μm broad. Raphe is median, more sigmoid than the valve, eccentric near the apices.
Central nodule is small, nearly circular and transversely dilated. Striation pattern is decussate. Oblique striae
are intersecting at nearly 70°, 20-21 in 10 μm, crossed by transverse striae, 18 in 10 μm. Two multilobed
chloroplasts extend on either side of the central nucleus.
Remarks:
According to Stidolph (2002), Pleurosigma falx closely resembles specimens of Pleurosigma normanii Ralfs
ex Pritchard and may be synonymized with P. normanii, which has nomenclatural priority. Valves of P.
falx from Kuwait’s material are more slender than lanceolate P. normanii. There is no difference between
these species discerned with LM except in valve shape and proportions, therefore, further studies with
SEM need to be performed to clarify the species identity.
Distribution:
Pleurosigma falx is a marine littoral warm water species, occasionally reported from neritic habitats. It has
been previously reported from Kuwait’s intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Pleurosigma falx sporadically occurred in Kuwait’s waters, and was observed in low numbers.

BACILLARIOPHYCEAE

a-f – Pleurosigma elongatum: a – live cell in valve view; b – detailed part of cell in valve view; c –
detailed part of acid cleaned frustule in valve view; d, e – detailed central part of cleaned valve;
f – detailed end of cleaned valve; g-k – Pleurosigma falx: g – live cell in valve view; h – acid cleaned
frustule in valve view; i – detailed central part of cleaned valve; j, k – detailed ends of cleaned valve.
LM, BF illumination (a, b, g) and phase contrast (c, d, f, h-k); SEM (e); scale bars: a, g, h – 20 μm; b,
c – 10 μm; d, i-k – 5 μm; e – 2 μm.
Pleurosigma formosum Smith
Plate 132
Péragallo & Péragallo, 1897-1908: Pl. XXX: 1-5; Hendey, 1964: p. 242: Hendey, 1970: p. 151; Podzorski & Håkansson,
1987: Pl. 35: 1, 1a; Ricard, 1987: Figs. 787, 788; Reid, 2002: Figs. 54-58; Hein et al., 2008: Fig. 56: 1; Al-Kandari et al.,
2009: p. 109, Pl. 61: F-H; Al-Yamani & Saburova, 2011: Pl. 105: a-i; Sterrenburg et al., 2014b: Figs. 1, 3a.
Synonymized names:
Gyrosigma formosum (Smith) Griffith et Henfrey 1856
Pleurosigma australicum Witt 1873
Pleurosigma tahitense Witt 1873
Scalprum formosum (Smith) Kuntze 1891
Cells are large, robust, solitary. Valves are linear with almost parallel margins and obliquely rounded and
slightly curved sigmoid apices, 160-340 μm long and 20-35 μm broad. Raphe is strongly sigmoid, eccentric
towards the apices, extending across the valve at about middle distance and becoming almost coincident
with the convex margin as it approaches the apices. Central area is small, transversely expanded. Striation
pattern is decussate. Oblique striae are intersecting at nearly 85-90°, 14 in 10 μm, crossed by transverse striae,
16-18 in 10 μm. Two large folded ribbon-shaped chloroplasts are lying in the valvar plane.
Distribution:
Pleurosigma formosum is a marine mainly littoral species widely distributed from temperate to tropical
regions, sporadically found in the plankton as well. It has been previously reported from Kuwait’s shores
(Hendey, 1970), coastal waters (Al-Yamani et al., 2004; Al-Kandari et al., 2009), and intertidal sediments
Local occurrence:
Pleurosigma formosum commonly occurred across Kuwait’s waters, being recorded from 56% of all
higher concentrations (6.30-9.98∙103 cells/l) recorded in the inshore waters of Kuwait Bay and around
Bubiyan Island during the period from October to February.
Pleurosigma inflatum Shadbolt

Plates 130: d-h; 133: a-d
Allen & Cupp, 1935: Fig. 103; Hendey, 1964: p. 243; Simonsen, 1974: p. 46; Foged, 1984: Pl. XXXVIII: 3; Podzorski &
Håkansson, 1987: Pl. 35: 3 (as Pleurosigma naviculaceum); Sterrenburg, 2001: Figs. 1-5, 11-14; Al-Yamani & Saburova,
2011: Pl. 108: a-h (as Pleurosigma naviculaceum); Sar et al., 2013: Figs. 7-9.
Synonymized names:
Pleurosigma naviculaceum Brébisson 1854
Pleurosigma transversale Smith 1856
Cells are robust, solitary. Valves are broadly lanceolate, moderately sigmoid, gently tapering to acutely
rounded apices, 37-85 μm long and 10-17 μm broad. Valve surface is broadly arched. Raphe is median,
sigmoid, and eccentric near the apices. External terminal raphe ends are hook-shaped, oppositely bent,
internally are terminating in helictoglossae turned toward the concave margin of the valve, with a large apical
pore. Central area is distinct and circular, internally transverse elongated, with central bars surrounding the
central nodule. Internal central raphe endings are slightly dilated. Striation pattern is decussate, striae are
more distant near the center than at the ends, but the terminal striae are at a more acute angle. Oblique striae
are intersecting at nearly 60°, 20-21 in 10 μm, crossed by transverse striae, 24 in 10 μm. Two multilobed
chloroplasts extend on either side of the central nucleus.
Remarks:
Pleurosigma inflatum closely resembles specimens of Pleurosigma diversestriatum (see above), but differs by
more arched valves, denser striation, larger central area, and less eccentric raphe position.

BACILLARIOPHYCEAE
Plate 132. Pleurosigma formosum.

a, b – live cells in valve view; c – detailed part of cell in valve view; d, e – detailed parts of
acid cleaned frustules in valve view; f-h – detailed central parts of valves; i, j – detailed ends
of valves.
LM, BF illumination (a-c, f, g, i) and phase contrast (d, e, h, j); scale bars: a, b – 25 μm; c – 20
μm; d-i – 10 μm; j – 5 μm.
Distribution:
Pleurosigma inflatum is mainly a littoral species, but is frequently found in the plankton. It has been
previously reported as regularly occurring in coastal waters of the northern Arabian Gulf (Simonsen,
1974, as Pleurosigma naviculaceum), and from Kuwait’s intertidal sediments (Al-Yamani & Saburova,
2011, as Pleurosigma naviculaceum).
Local occurrence:
Pleurosigma inflatum frequently occurred across Kuwait’s waters, being recorded from 48.5% of all
higher concentrations (3.6-4.4∙103 cells/l) recorded in the inshore waters of Kuwait Bay and the adjacent
waters.
Pleurosigma inscriptura Harper

Plate 133: e-h
Sar et al., 2013: Figs. 10, 11.
Cells are solitary. Valves are lanceolate with broadly rounded apices, 166-180 μm long and 23-26 μm broad.
Raphe is sigmoid, almost straight in the central area and gradually deviated in the opposite direction towards
the apices. Central area is small, circular. Striation pattern is decussate. Oblique striae are intersecting at
nearly 60-65°, 18-19 in 10 μm, crossed by transverse striae, 20 in 10 μm. Two large deeply lobed chloroplasts
are lying in the valvar plane.
Distribution:
Pleurosigma inscriptura is a marine neritic species known from New Zealand and South America,
Argentina (Sar et al., 2013).
Local occurrence:
Pleurosigma inscriptura sporadically occurred in Kuwait’s waters, and was observed in low numbers. The
species is reported for the first time from Kuwait.
Pleurosigma intermedium Smith

Plate 134: a-e
Péragallo & Péragallo, 1897-1908: Pl. XXXII: 19-21; Hendey, 1964: p. 244; Al-Yamani & Saburova, 2011: Pl. 106: a-d;
Lobban et al., 2012: Pl. 51: 2-4; Sar et al., 2012: Figs. 1-12.
Synonymized names:
Pleurosigma nubecula var. intermedium (Smith) Cleve 1894
Pleurosigma intermedium var. nubecula (Smith) Grunow ex Van Heurck 1896
Cells are solitary. Valves are slender lanceolate with acute apices, 123-147 μm long and 13-17 μm broad.
Raphe is median, straight for most of its length, becoming a little eccentric with minimal curvature near the
apices. Central area is transversely elliptic. Striation pattern is decussate. Oblique striae are intersecting at
nearly 60-65°, 20-21 in 10 μm, crossed by transverse striae, 21-23 in 10 μm. Two large folded ribbon-shaped
chloroplasts are lying in the valvar plane.
Distribution:
Pleurosigma intermedium is a marine neritic and littoral species with a worldwide distribution in warm
waters. The species has been observed epiphytically on different substrata along Kuwait’s shores as well
as in the intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Pleurosigma intermedium is of low occurrence in Kuwait’s southern offshore waters, being recorded from
less that 1% of all collected samples, always in low numbers.

BACILLARIOPHYCEAE

a-d – Pleurosigma inflatum: a – live cell in valve view; b, c – acid cleaned frustules in valve
view; d – detailed central part of cleaned valve; e-h – Pleurosigma inscriptura: e – live cell in
valve view; f – detailed part of cell in valve view; g – acid cleaned frustules in valve view;
h – detailed central part of cleaned valve.
LM, BF illumination (a, e, f) and phase contrast (b-d, g, h); scale bars: e – 20 μm; a-c, f-h –
10 μm; d – 5 μm.
Pleurosigma javanicum Grunow
Plate 135
Cleve, 1894: p. 42.
Synonymized names:
Pleurosigma angulatum var. javanicum Grunow ex Cleve et Möller 1878
Cells are solitary. Valves are lanceolate, sigmoid, gently tapering to acutely rounded apices, 105-119 μm long
and 19-25 μm broad. Valve surface is broadly arched. Raphe is median, sigmoid, and eccentric towards the
apices. External central raphe fissures are markedly projecting into the central area, overlap and curve sharply
away from each other. Central area is circular, often with bilateral dilatation around the central nodule.
Striation pattern is decussate. Oblique striae are intersecting at nearly 60-65°, 21-23 in 10 μm, crossed by
transverse striae, 24-25 in 10 μm. Two large folded ribbon-shaped chloroplasts are lying in the valvar plane.
Remarks:
The Pleurosigma species illustrated here (Pl. 135) corresponds closely to Pleurosigma angulatum (Quekett)
Smith, which is morphologically highly variable species (Sterrenburg, 1991), and whose occurrence in
Kuwait’s marine environment has been reported previously (Al-Yamani & Saburova, 2011: Pl. 100: a-f).
The reason for presenting Pleurosigma javanicum separately is the observed morphological and ecological
distinctions between these two species in Kuwait’s material. P. javanicum has more lanceolate valves
compared to rhombic-lanceolate shape of P. angulatum, and differs in striae density (21-25 in 10 μm in
P. javanicum versus 17-18 in P. angulatum). Moreover, in contrast to P. angulatum inhabiting Kuwait’s
intertidal sediments (Al-Yamani & Saburova, 2011), P. javanicum has a pelagic lifestyle.
Distribution:
Pleurosigma javanicum is a marine pelagic warm water species (Cleve, 1894).
Local occurrence:
Pleurosigma javanicum was occasionally observed in Kuwait’s waters, at times was quite abundant, but
easily misidentified with P. inflata and P. diversestriata in routine phytoplankton monitoring. Further
morphological study needs to be performed to clarify the species identification.
Pleurosigma speciosum var. javanicum Péragallo

Plate 134: f-j
Reid, 2002: Figs. 42-46 (type material); Al-Yamani & Saburova, 2011: Pl. 110: a-c (as Pleurosigma speciosum).
Synonymized names:
Pleurosigma speciosum var. sumatricum Péragallo 1891
Cells are solitary. Valves are broadly linear with almost parallel valve margins and obliquely rounded and
slightly curved sigmoid apices, 112-134 μm long, 22-25 μm broad. Raphe is median, with straight central
part, strongly eccentric towards the apices. Central nodule is small, rounded. Striation pattern is decussate.
Oblique and transapical striae are almost equidistant, 19-21 in 10 μm. Oblique striae are intersecting at
nearly 60°. Two large folded ribbon-shaped chloroplasts are lying in the valvar plane.
Remarks:
Pleurosigma speciosum var. javanicum differs from the nominate variety (Pleurosigma speciosum Smith) in
size, valve shape, central hyaline area outline, and habitat. The variety is smaller, more lanceolate than
linear in valve outline, with nearly circular than transversely expanded central hyaline area. Pleurosigma
speciosum var. javanicum is warm water forms compared to records of nominate variety limited to the
northern temperate localities (Reid, 2002). The specimens illustrated here (Pl. 134: f-j) corresponds to
Pleurosigma speciosum var. javanicum rather than to the nominate variety in respect to its size range, valve
outline, shape of central hyaline area, and subtropical habitat.
Distribution:
Pleurosigma speciosum var. javanicum is a marine warm water scarcely reported species (Reid, 2002). It
has been previously reported from Kuwait’s intertidal sediments (Al-Yamani & Saburova, 2011, as
Pleurosigma speciosum).
Local occurrence:
Pleurosigma speciosum var. javanicum sporadically occurred in Kuwait’s waters, always in low numbers.

BACILLARIOPHYCEAE

a-e – Pleurosigma intermedium: a – live cell in valve view; b – detailed part of cell in valve view;
c – acid cleaned frustule in valve view; d – detailed central part of cleaned valve; e – detailed end
of valve; f-j – Pleurosigma speciosum: f – live cell in valve view; g, h – acid cleaned frustules in valve
view; i – detailed end of cleaned valve; j – detailed central part of cleaned valve.
LM, BF illumination (a, b, f) and phase contrast (c-e, g-j); scale bars: f – 20 μm; a-c, g, h – 10 μm;
d, e, i, j – 5 μm.
Plate 135. Pleurosigma javanicum.
a, b – live cells in valve view; c – detailed part of cell in oblique valve view; d – detailed part of acid cleaned
frustule in valve view; e-h – detailed central parts of valves; i – detailed end of valve.
LM, BF illumination (a-c, e-g) and phase contrast (d, h, i); scale bars: a-d – 10 μm; e-i – 5 μm.

Pleurosigma strigosum Smith
Plate 136: a-f
Péragallo & Péragallo, 1897-1908: Pl. XXXII: 22, 23 (as Pleurosigma angulatum var. strigosa); Hendey, 1964: Pl. XXXVI:
7; Sterrenburg, 2003: Figs. 1-4, 22-25 (type material); Al-Kandari et al., 2009: p. 109, Pl. 62: A-C; Al-Yamani &
Saburova, 2011: Pl. 111: a-d; Al-Handal et al., 2014: Fig. 56.
Synonymized names:
Gyrosigma strigosum (Smith) Griffith et Henfrey 1856
Pleurosigma angulatum v. strigosa Van Heurck 1885
Pleurosigma strigosum var. genuinum Cleve-Euler 1952
Cells are solitary. Valves are lanceolate, sigmoid, with obtusely rounded and curved sigmoid apices, 122-
140 μm long and 24-26 μm broad. Raphe is median, almost straight for most of its length, gently curving
and slightly eccentric towards the apices. Central nodule is small, roundish. Striation pattern is decussate.
Oblique and transapical striae are almost equidistant, 18-21 in 10 μm. Oblique striae crossed each over at an
angle of about 60-65º. Two large folded ribbon-shaped chloroplasts are lying in the valvar plane.
Distribution:
Pleurosigma strigosum is a marine mainly littoral species, but is sporadically found in the plankton. It has
been previously recorded from the epiphytic assemblages in the Mesopotamian wetland, South Iraq (Al-
Handal et al., 2014), and from Kuwait’s coastal waters (Al-Kandari et al., 2009) and intertidal sediments
Local occurrence:
Pleurosigma strigosum is of low occurrence in Kuwait’s inshore waters, and found in low numbers.
Genus Toxonidea Donkin
Toxonidea insignis Donkin

Plate 136: g-j
Péragallo & Péragallo, 1897-1908: Pl. XXXVI: 23, 24; Hendey, 1964: p. 251; Reid, 2006: Figs. 58-70.
Cells are solitary. Valves are strongly asymmetrical on the apical axis, arcuate, with nearly straight ventral
margin and strongly convex dorsal margin, tapering into sub-rostrate apices, 86-97 μm long and 20-22 μm
broad. Raphe is arcuate, lying alongside ventral margin. Valve surface is very finely striated. Transapical and
oblique striae are almost equidistant, about 22-24 in 10 μm. Large folded ribbon-shaped chloroplasts are
BACILLARIOPHYCEAE
lying through the valve. Nucleus is median, appressed to the dorsal side of the valve.
Distribution:
Toxonidea insignis is a marine species distributed in temperate coastal waters of European seas (the shores
of the British Isles and North Sea coastline of Europe, up into the Kattegat, Denmark) (Reid, 2006).
Local occurrence:
A few cells of Toxonidea insignis were observed from the water samples collected in Kuwait’s southern
waters in February. The species is recorded for the first time from Kuwait’s subtropical waters.
Plate 136. Pleurosigma strigosum and Toxonidea insignis.
a-f – Pleurosigma strigosum: a – live cell in valve view; b, c – acid cleaned frustules
in valve view; d, e – detailed central part of valve; f – detailed end of valve; g-j –
Toxonidea insignis: g-i – live cells in valve view; j – collapsed cell showing eccentric
raphe (arrowheads).
LM, BF illumination (a, d), DIC optic (b, g-j), and phase contrast (c, e, f); scale bars:
a-c, g-j – 10 μm; d-f – 5 μm.

Suborder Neidiineae Mann
Family Amphipleuraceae Grunow
Genus Amphiprora Ehrenberg
Amphiprora angustata Hendey

Plate 137: j-l
Hendey, 1964: Pl. XLIII: 1-6.
Cells are large, solitary. Frustules are deeply constricted in the middle in girdle view, 62-77 µm long and 32-
36 µm broad at the widest part. Valves are linear-lanceolate, slightly constricted in the middle, with acute
apices. Junction line between valve surface and the keel is weakly arcuate. Keel is furnished with a strongly
developed marginal rib. Valve surface is finely striated. Transapical striae are parallel, continuous through to
the margin of the keels, 24 in 10 µm. Striae on the keels are slightly radiate. Girdle is composed of numerous
narrow longitudinal bands. A single large plate-like chloroplast extends along the girdle.
Distribution:
Amphiprora angustata is a marine neritic species recorded scarcely from temperate localities (Europe,
British coastal waters, Australia, and New Zealand) (Hendey, 1964; Guiry & Guiry, 2017).
Local occurrence:
Amphiprora angustata sporadically occurred in Kuwait’s coastal waters, and was observed in low numbers.
Amphiprora gigantea var. sulcata (O’Meara) Cleve

Plate 137: a-i
Péragallo & Péragallo, 1897-1908: Pl. XXXVIII: 1-5 (as Amphiprora sulcata); Allen & Cupp, 1935: Fig. 113; Cupp, 1943:
Fig. 151; Simonsen, 1974: p. 45 (as Amphiprora sulcata); Foged, 1984: Pl. LII: 2 (as Amphiprora sulcata); Al-Kandari et
al., 2009: p. 120, Pls. 77: D-F, 78: A, B (as Entomoneis sulcata); Al-Yamani & Saburova, 2011: Pl. 168: a-h (as Amphiprora
sulcata).
Synonymized names:
Amphiprora sulcata O’Meara 1871
Cells are large, solitary. Frustules are narrowly rectangular in girdle view, strongly constricted in the middle,
with biarcuated sides, 108-131 μm long and 31-38 μm broad at the widest part, with four distinct marginal
wings. Keel separation line is weakly biarcuated. Valve surface is finely striated. Transapical striae in the
middle are slightly curved, divergent from central nodule, becoming parallel towards apices, 11-12 in 10 μm.
Wings are furnished with puncta arranged in obliquely decussating rows, 12 in 10 μm. Girdle is composed
BACILLARIOPHYCEAE
of numerous coarsely punctate intercalary bands, 10-11 puncta in 10 μm. A single large elongate chloroplast
with wavy edges lies along the girdle.
Distribution:
Amphiprora gigantea var. sulcata is principally a marine littoral warm water species, occasionally occurring
in the plankton. It has been previously reported from the Indian Ocean and the Arabian Gulf (Simonsen,
1974, as Amphiprora sulcata), and from Kuwait’s waters (Al-Yamani et al., 2004; Al-Kandari et al., 2009,
as Entomoneis sulcata) and intertidal sediments (Al-Yamani & Saburova, 2011, as Amphiprora sulcata).
Local occurrence:
Amphiprora gigantea var. sulcata frequently occurred in Kuwait’s waters, being recorded from 44.5% of
all collected samples. The species was present throughout the year in low to moderate abundance with
period from February to October.
Plate 137. Amphiprora spp.
a-i – Amphiprora gigantea var. sulcata: a-e – live cells in girdle view; f-h – acid cleaned
frustules in girdle view; i – detailed end of valve; j-l – Amphiprora angustata: j – live cell in
girdle view; k – part of acid cleaned frustule in girdle view; l – acid cleaned valve.
LM, BF illumination (a-e, j) and phase contrast (f-i, k, l); scale bars: a, b, i-l – 0 μm; c-f – 25
μm; g, h – 20 μm.

Family Berkeleyaceae Mann
Genus Climaconeis Grunow
Climaconeis lorenzii Grunow

Plate 138: a
Podzorski & Håkansson, 1987: Pl. 31: 1. 2 (as Navicula lorenzii); Prasad et al., 2000: Figs. 35-42; Hein et al., 2008: Pl.
26: 1; Lobban et al., 2010: Figs. 11, 33, 34.
Synonymized names:
Climaconeis frauenfeldii Grunow 1862
Climacosphenia linearis Janisch et Rabenhorst 1863
Stictodesmis australis Greville 1863
Navicula famelica Castracane 1889
Denticula paucicostata Meister 1932
Navicula lorenzii (Grunow) Hustedt 1961
Frustules are narrowly linear in girdle view, slightly gibbous in the middle, with rounded apices, 75-112 µm
long and 6-7 µm broad. Numerous transverse septa (craticular bars) are widely spaced or absent at the center,
and more closely distributed toward the cell apices. Valve surface is striated. Transapical striae are punctate,
18 in 10 µm.
Distribution:
Climaconeis lorenzii is a marine widespread warm water species (Lobban et al., 2010).
Local occurrence:
Climaconeis lorenzii is a very rare species in Kuwait’s material. Several frustules were found in plankton
hauls from Kuwait’s northern waters in the vicinity of Bubiyan Island. The species is recorded for the
first time in Kuwait’s waters.
Climaconeis undulata (Meister) Lobban, Ashworth et Theriot

Plate 138: b-d, g-j
Lobban et al., 2010: Figs. 5-7, 19-25.
Synonymized names:
Gomphocaloneis undulata Meister 1932
Navicula scopulorum Brébisson var. triundulata Meister
Navicula scopulorum Brébisson f. triundulata Meister
Climaconeis mabikii Park, Khim et Lee 2016 ?
BACILLARIOPHYCEAE
Cells are long, slender and solitary. Valves are elongate, linear, 60-92 μm long and 6-8 μm broad. Valve sides
are slightly inflated at the center, halfway between the centre and apex, and again at the apices, giving a
slightly undulating outline. Raphe is straight, axial area is very narrow, central area is small, oblong. Valve
surface is distinctly striated. Transapical striae are punctate, slightly radiate at the center, becoming parallel
in the middle, and convergent near the apices, 18 in 10 μm. Shorter striae are intercalated between the main
striae alongside the central area.
Distribution:
Climaconeis undulata is a marine warm water tropical species, reported from the western Pacific
(Southern China, Sumatra, Guam) (Lobban et al., 2010). The species has been previously reported from
Kuwait’s intertidal sediments (Al-Yamani & Saburova, 2011, as Berkeleya scopulorum). Recently, a very
morphologically similar species, Climaconeis mabikii Park, Khim et Lee 2016, has been described from
northeast Asia (Park et al., 2016b), which differs from C. undulata practically in size only.
Local occurrence:
Frustules of Climaconeis undulata were occasionally found in plankton hauls from Kuwait’s northern
waters in the vicinity of Bubiyan Island. The species is recorded for the first time in Kuwait’s waters.
Climaconeis sp.
Plate 138: e, f
Cells are long, slender, solitary, and slightly arcuate in valve view, with weakly capitate apices, 84-115 μm
long and 4-6 μm broad. Two large H-shaped chloroplasts with conspicuous pyrenoids are located on each
side of the central nucleus. Valve striation was not observable with LM from live cells.
Local occurrence:
Cells of Climaconeis sp. were occasionally found in Kuwait’s southern waters around Kubbar Island.
Plate 138. Climaconeis spp.

a – Climaconeis lorenzi: acid cleaned frustule; b-d, g-j – Climaconeis undulata:
b, c, g, h – acid cleaned frustules in valve view; d, i, j – detailed parts of
cleaned valves; e, f – Climaconeis sp.: live cells showing chloroplasts with
pyrenoids (arrowheads).
LM, BF illumination (e, f) and phase contrast (a-d, g-j); scale bars: a-i – 10
μm; j – 5 μm.

Order Bacillariales Hendey
Family Bacillariaceae Ehrenberg
Genus Bacillaria Gmelin
Bacillaria paxillifera (Müller) Marsson

Plate 139
Pavillard, 1925: Fig. 112 (as Bacillaria paradoxa); Allen & Cupp, 1935: Fig. 117 (as Bacillaria paradoxa); Cupp, 1943: Fig.
159 (as Nitzschia paradoxa); Hendey, 1964: Pl. XXI: 5; Simonsen, 1974: p. 49 (as Bacillaria paradoxa); Ricard, 1987: Figs.
1125-1128 (as Bacillaria paradoxa); Throndsen et al., 2007: p. 192; Hoppenrath et al., 2009: Fig. 46 j-o; Al-Handal et al.,
2014: Fig. 93.
Synonymized names:
Vibrio paxillifer Müller 1786
Bacillaria paradoxa Gmelin 1791
Oscillaria paxillifera (Müller) Schrank 1823
Oscillatoria paxillifer (Müller) Schrank 1823
Diatoma paxillifera (Müller) Brébisson 1838
Nitzschia paxillifera (Müller) Heiberg 1863
Nitzschia paradoxa (Gmelin) Grunow 1880
Oscillatoria paxillifera (Müller) Schrank ex Gomont 1892
Homoeocladia paxillifer (Müller) Elmore 1921
Cells are united valve to valve to form block-shaped movable colony. Within colony, each cell slides along the
surface of the sibling cell, for full length of the raphe. General shape of colony is very variable, rectangular
to filamentous due to cells motility. Frustules are narrowly rectangular in girdle view, 120-135 μm long
and 6-7 μm broad. Valves are linear-lanceolate with produced apices. Keeled raphe is central. Fibulae are
nearly evenly spaced, 7-9 in 10 μm. Transapical striae are finely punctate, 20-21 in 10 μm. Numerous small
irregularly shaped plate-like chloroplasts are distributed along the girdle. Nucleus is centrally located.
Distribution:
Bacillaria paxillifera is a very common and widely distributed marine plankton species. It has been
previously reported (as Bacillaria paradoxa) from the Arabian Gulf (Simonsen, 1974), from the epiphytic
assemblages in the Mesopotamian wetland, South Iraq (Al-Handal et al., 2014), and from Kuwait’s
Local occurrence:
Bacillaria paxillifera frequently occurred across Kuwait’s waters, being recorded from 46% of all collected
samples. The species was present all year round in low to moderate concentrations, more abundantly
BACILLARIOPHYCEAE
(1.4-4.3∙104 cells/l) in the inshore waters around Bubiyan Island from February to September.
Genus Cylindrotheca Rabenhorst
Cylindrotheca closterium (Ehrenberg) Reimann et Lewin

Plate 140: a-e
Péragallo & Péragallo, 1897-1908: Pl. LXXIV: 15 (as Nitzschia closterium); Allen & Cupp, 1935: Fig. 122 (as Nitzschia
closterium); Cupp, 1943: Fig. 153 (as Nitzschia closterium); Hendey, 1964: Pl. XXI: 8 (as Nitzschia closterium); Simonsen,
1974: p. 51; Ricard, 1987: Figs. 1094, 1095 (as Nitzschia closterium); Witkowski et al., 2000: Pl. 212: 4-6 (as Nitzschia
closterium); Throndsen et al., 2007: p. 192; Al-Kandari et al., 2009: p. 115, Pl. 69: E; Hoppenrath et al., 2009: Fig. 46 p,
q (as Ceratoneis closterium); Al-Yamani & Saburova, 2011: Pl. 160: e-h.
Synonymized names:
Ceratoneis closterium Ehrenberg 1839
Nitzschia closterium (Ehrenberg) Smith 1853
Nitzschiella tenuirostris Mereschkowsky 1901
Plate 139. Bacillaria paxillifera.
a – large colony in movement; b, c – live cells in girdle view united valve to valve to block-shaped colonies;
d, e - cells in colony in valve view; f – acid cleaned frustule in valve view; g, h – detailed central part of
cleaned valve showing keeled raphe with fibulae.
LM, BF illumination (a-e) and phase contrast (f, g); SEM (h); scale bars: a – 50 μm; c – 20 μm; b, d-f – 10
μm; g – 5 μm; h – 2 μm.

Cells are solitary, weakly silicified, slightly bent. Valves are lanceolate, abruptly narrowed into flexible
long rostrate slightly bent or straight apices, 28-33(up to 67) μm long and 3-3.5 μm broad. Keeled raphe is
eccentric, fibulae are about 7 in 10 µm, transapical striae are indistinct. Two elongate or irregularly shaped
chloroplasts extend on either side of the central nucleus, not in cell apices.
Distribution:
Cylindrotheca closterium is a very common marine and brackish water species, widely distributed in
temperate to tropical areas in the littoral zone (mud-dwelling) and neritic habitats. It has been previously
reported (as Nitzschia closterium) from the Indian Ocean (Simonsen, 1974), and from Kuwait’s coastal
waters (Al-Yamani et al., 2004; Al-Kandari et al., 2009) and intertidal sediments (Al-Yamani & Saburova,
2011).
Local occurrence:
Cylindrotheca closterium frequently occurred across Kuwait’s waters, being recorded from 44% of all
collected samples. The species was present throughout the year in moderate to high abundance with
higher concentrations (0.9-2.4∙106 cells/l) recorded in the southern inshore waters during the spring
season in March.
Harmful Effect:
Cylindrotheca closterium was occasionally found in bloom proportions in Kuwait’s marine environment.
Due to its capacity to form high biomass blooms, the species has been included in the list of potentially
harmful species for Kuwait’s marine environment (Al-Yamani et al., 2012).
Genus Cymatonitzschia Simonsen
Cymatonitzschia marina (Lewis) Simonsen

Plate 140: f, g
Hendey, 1957: Pl. I: 9 (as Nitzschia antillarum); Simonsen, 1974: p. 56, Pl. 41: 5-9; Ricard, 1987: p. 281, Figs. 1129, 1130;
Al-Yamani & Saburova, 2011: Pl. 146: g-i.
Synonymized names:
Cymatopleura marina Lewis 1861
Denticula antillarum Cleve et Grunow 1878
Nitzschia antillarum Meister 1937
Cells are solitary. Valves are lanceolate with acute apices, 37-41 μm long and 8-9 μm broad. Keeled raphe is
eccentric, located near one margin. Parapical axis of the valves is conspicuously undulated, undulations of
BACILLARIOPHYCEAE
the two valves are opposing each over. Valve surface is furnished with irregularly areolated and depressed
areas, alternating with elevated hyaline zones.
Distribution:
Cymatonitzschia marina is obviously a tropical species, found in the plankton samples as only allochthonous.
It has been previously reported from the Arabian Gulf (Simonsen, 1974) and from Kuwait’s intertidal
Local occurrence:
Frustules of Cymatonitzschia marina were found in plankton net hauls from the inshore waters around
Bubiyan Island, and in low numbers.
Genus Giffenia Round et Basson
Giffenia cocconeiformis (Grunow) Round et Basson

Plate 140: h-k
Round et al., 1990: p. 678 (as Tryblionella cocconeiformis); Round & Basson, 1997: Figs. 1-12; Witkowski et al., 2000: Pl.
188: 8, 9 (as Nitzschia cocconeiformis); Al-Yamani & Saburova, 2011: Pl. 164: e-g (as Tryblionella cocconeiformis).
Synonymized names:
Nitzschia cocconeiformis Grunow 1880
Tryblionella cocconeiformis (Grunow) Mann 1990
Valves are broadly lanceolate to elliptic with rounded apices, 29-32 μm long and 9-10 μm broad. Valve surface
is transapically undulated, with slightly raised edge. Keeled raphe is eccentric, marginal. Transapical striae
are multiseriate, alternating with robust transverse costae, parallel in the middle, becoming radiate towards
apices, 8.5-9 in 10 μm, interrupted by hyaline longitudinal wavy fold (sternum), lying along the apical axis.
Off-center sternum is moderately broad in the middle, tapering towards apices, sometimes curved. Fibulae
are as many as striae.
Distribution:
Giffenia cocconeiformis is a marine to brackish water cosmopolitan littoral species. It was sporadically
observed in Kuwait’s marine sediments (Al-Yamani & Saburova, 2011, as Tryblionella cocconeiformis).
Local occurrence:
Frustules of Giffenia cocconeiformis were found in plankton net hauls from the inshore Kuwait’s waters in
low numbers.
Genus Gomphotheca Hendey et Sims
Gomphotheca sinensis (Skvortzov) Hendey et Sims

Plate 147: j-m
Hendey & Sims, 1982: Figs. 1-12; Tynni, 1983: Pl. 10: 102; Al-Handal, 2009: Figs. 50-52.
Synonymized names:
Nitzschia chinensis Skvortzow 1932
Frustules are slender, extremely long, solitary. Valves are straight, linear, with acutely rounded apices and
several flattened areas, which appear as undulation of the valve margins, 460-486 μm long and 12-13 μm
broad. Raphe is central, bearing two grooves, one on either side of the raphe and parallel to it. Fibulae are
unevenly spaced, connect with one or two thickened transapical rib-like costae extending across the valve,
5.5-6 in 10 μm.
Distribution:
Gomphotheca sinensis is a marine to brackish water species scarcely recorded in subtropical and tropical
habitats (Round et al., 1990). The species has been previously reported from the bottom sediments in the
Shatt Al-Arab Estuary, Iraq (Tynni, 1983; Al-Handal, 2009).
Local occurrence:
Frustules of Gomphotheca sinensis were occasionally recorded from plankton hauls collected from
Kuwait’s northern waters around Bubiyan Island, and observed in low numbers. The species is recorded
for the first time from Kuwait.

BACILLARIOPHYCEAE
Plate 140. Cylindrotheca closterium, Cymatonitzschia marina, and Giffenia cocconeiformis.

a-e – Cylindrotheca closterium: live cells; f, g – Cymatonitzschia marina: acid cleaned frustules in valve view;
h-k – Giffenia cocconeiformis: acid cleaned frustules in valve view.
LM, BF illumination (a-d), DIC optic (h, i), and phase contrast (e-g, j, k); scale bars: a, b – 10 μm; c-k – 5 μm.
Genus Nitzschia Hassall
The species belonging to the genus Nitzschia are distinguished by the presence of an eccentric unbroken
keeled raphe on the opposite side of each valve, uniseriate striae and distinct keel puncta (fibulae). The
species are identified by the size, shape of valve and valve apices. Number of fibulae and striae, and the
quotient between them are important characters at species level (Round et al., 1990; Hasle & Syvertsen,
1997).
Nitzschia amabilis Suzuki

Plate 141: i-o
Witkowski et al., 2000: Pl. 190: 1-6 (as Nitzschia laevis); Suzuki et al., 2010: Fig. 1; Al-Yamani & Saburova, 2011: Pl.
153: f-k (as Nitzschia laevis).
Synonymized names:
Nitzschia laevis Husted 1939
Cells are small, solitary. Valves are linear-elliptic with acutely rounded apices, 12-16 μm long and 3-6
μm broad. Keeled raphe is eccentric, marginal. Fibulae are unevenly spaced, 13-15 in 10 μm, the median
two being further apart to form a central nodule. Valve surface ornamentation is indiscernible with LM.
Transapical striae are finely punctate, 34-37 in 10 μm as discerned with SEM. Two plate-like chloroplasts
extend on either side of the central nucleus toward the apices, lying obliquely and appressed to opposite
valves.
Distribution:
Nitzschia amabilis is a marine cosmopolitan widespread species (Witkowski et al., 2000). It has been
previously reported from Kuwait’s intertidal sediments (Al-Yamani & Saburova, 2011, as Nitzschia laevis).
Local occurrence:
Nitzschia amabilis occasionally occurred in Kuwait’s coastal waters, being recorded from 8% of all
collected samples, sometimes in bloom proportions (Al-Yamani et al., 2012, as Nitzschia laevis).
Harmful Effect:
Nitzschia amabilis was found to form nontoxic high biomass harmful blooms in Kuwait’s marine
environment. The conspicuous bloom of this species resulted in water discoloration was detected in
the coastal waters of Kuwait Bay in March 2006 with cells concentration of more than 108 cells/l (for
details see Sect. 2, Vol. I). Due to its capacity to form blooms, the species has been included in the list of
potentially harmful species for Kuwait’s marine environment (Al-Yamani et al., 2012, as Nitzschia laevis).
Nitzschia bicapitata Cleve

Plate 141: a-e
Simonsen, 1974: Pl. 35: 3-15; Hasle & Syvertsen, 1997: Pl. 74.
Cells are solitary. Valves are lanceolate in outline, with more or less long capitate apices, 11-38 μm long and
4-6 μm broad. Keeled raphe is eccentric, marginal. Fibulae are evenly spaced, 16-18 in 10 μm. Central pair
of fibulae is more widely separated than the others. Valve surface ornamentation is indiscernible with LM.
Transapical striae are finely punctate, 40-42 in 10 μm as discerned with SEM. Two plate-like chloroplasts
extend on either side of the central nucleus toward the apices.
Distribution:
Nitzschia bicapitata is a marine cosmopolitan plankton species (Hasle & Syvertsen, 1997). It has been
previously reported from the Arabian Gulf (Simonsen, 1974).

BACILLARIOPHYCEAE
Plate 141. Nitzschia spp.

a-e – Nitzschia bicapitata: a – small cells (arrowheads) attached to floating detritus; inset (2x
enlargement) – single live cell; b – external valve view showing areolation; c-e – acid cleaned frustules;
f – Nitzschia braarudii: acid cleaned frustule; g, h – Nitzschia fluminensis: g – live cell; h – acid cleaned
frustule; i-o – Nitzschia amabilis: i-k – live cells; l-o – acid cleaned frustules.
LM, BF illumination (a, g, i-k) and phase contrast (c-f, h, l-n); SEM (b, o); scale bars: a, g, h – 10 μm;
b-f, i-o – 5 μm.
Local occurrence:
The present observations constitute the first record of Nitzschia bicapitata from Kuwait’s waters. The
species occasionally occurred, mostly in the inshore waters of Kuwait Bay but can be easily misidentified
with other small-sized Nitzschia species during routine phytoplankton examination.
Nitzschia braarudii Hasle

Plate 141: f
Simonsen, 1974: Pl. 35: 2; Hasle & Syvertsen, 1997: Pl. 77.
Synonymized names:
Nitzschia capitata Heiden 1928
Cells are solitary. Valves are linearly lanceolate in outline, gradually tapering into produced and slightly
capitate apices, 32-35 μm long and 3.5-4 μm broad. Keeled raphe is eccentric, marginal. Fibulae are evenly
spaced, 10-12 in 10 μm. Valve surface is finely striated. Transapical striae are distinctly punctate, 25-26 in 10
μm. Central pair of fibulae is more widely separated than the others.
Distribution:
Nitzschia braarudii is a marine warm water plankton species (Simonsen, 1974; Hasle & Syvertsen, 1997).
It has been previously reported from the Arabian Gulf (Simonsen, 1974).
Local occurrence:
The present observations constitute the first record of Nitzschia braarudii from Kuwait’s waters. The
species sporadically occurred but can be easily misidentified with other small-sized Nitzschia species
during routine phytoplankton examination.
Nitzschia dissipata (Kützing) Rabenhorst

Plate 147: f
Manoylov, 2010; Lobban et al., 2012: Pl. 60: 1-3.
Synonymized names:
Synedra dissipata Kützing 1844
Nitzschia palea f. dissipata (Kützing) Rabenhorst 1864
Homoeocladia dissipata (Kützing) Kuntze 1898
Synedra fasciculata Ehrenberg 1832
Nitzschia palea var. dissipata (Kützing) Schönfeldt 1907
Nitzschia dissipata var. genuina Mayer 1913
Cells are solitary. Valves are linearly lanceolate in outline, with rostrate apices, 30-33 μm long and 4-4.5 μm
broad. Canal raphe is prominent, central to slightly eccentric. Fibulae are short, rib-like, distant and evenly
spaced, 6.5-7 in 10 μm, linked at their bases by faint longitudinal lines on either side of the raphe, which
mark the boundaries of extra flap of silica (conopeum). Valve surface ornamentation is indiscernible with
LM.
Distribution:
Nitzschia dissipata is an ecologically eurytopic and mainly of fresh to brackish water form. In the northern
Arabian Gulf, this species has been reported from the Shatt Al-Arab River system, Iraq (Maulood et al.,
2013).
Local occurrence:
Frustules of Nitzschia dissipata were sporadically found in plankton hauls from Kuwait’s northern waters
in the vicinity of Bubiyan Island, and may be of secondary origin indicating the intrusion of fresh waters
from the Shatt Al-Arab River system. The species is recorded for the first time in Kuwait’s waters.

Nitzschia distans Gregory
Plate 142: a-c
Péragallo & Péragallo, 1897-1908: Pl. LXXIII: 1-3; Hendey, 1964: p. 281; Foged, 1984: Pl. LVIII: 13; Ricard, 1987:
Figs. 1069-1072; Witkowski et al., 2000: Pl. 203: 7-9; Al-Yamani & Saburova, 2011: Pl. 152: a-d.
Cells are solitary. Frustules are linearly rectangular in girdle view, with slightly inflated corners furnished
with small wind-like expansions. Valves are narrowly lanceolate, with acutely rounded apices, 66-71 μm
long and 7-8 μm broad. Raphe is central. Fibulae are short, rib-like, distant and unevenly spaced, 3.5-4 in 10
μm, linked at their bases by faint longitudinal lines on either side of the raphe. Valve surface ornamentation
is indiscernible with LM. Two large elongated and deeply lobed chloroplasts extend on either side of the
central nucleus toward the apices.
Distribution:
Nitzschia distans is a marine mainly littoral widely distributed species. It has been previously reported from
the Indian Ocean (Simonsen, 1974) and from Kuwait’s intertidal sediments (Al-Yamani & Saburova,
2011).
Local occurrence:
Nitzschia distans is a typically benthic species that observed in epipelic assemblages along Kuwait’s shores
(Al-Yamani & Saburova, 2011), and rarely occurred in Kuwait’s waters, being recorded from less than
1% of all collected samples in low numbers.
Nitzschia distans var. tumescens Grunow

Plate 142: d, e
Witkowski et al., 2000: Pl. 203: 10.
Cells are solitary. Frustules are linearly rectangular in girdle view, with slightly inflated corners furnished
with small wind-like expansions, 245-260 μm long and 50-55 μm broad. Valves are narrowly lanceolate, with
acutely rounded apices. Raphe is central. Fibulae are short, rib-like, distant and unevenly spaced, 2.5-3 in 10
μm, linked at their bases by faint longitudinal lines on either side of the raphe. Valve surface ornamentation
is indiscernible with LM. Girdle is longitudinally striated. Two large elongated and perforated chloroplasts
Distribution:
Nitzschia distans var. tumescens is a marine littoral species known from the Atlantic coasts of Europe and
North America, from South America, Colombia and Australia (Witkowski et al., 2000; Guiry & Guiry,
2017).
BACILLARIOPHYCEAE
Local occurrence:
Nitzschia distans var. tumescens was sporadically recorded from plankton hauls collected from Kuwait’s
waters, and observed in low numbers. The variety is recorded for the first time from Kuwait.
Nitzschia fluminensis Grunow

Plate 141: g, h
Péragallo & Péragallo, 1897-1908: Pl. LXXI: 9, 10; Foged, 1984: Pl. LVIII: 4; Witkowski et al., 2000: Pl. 202: 6-9; Hein
et al., 2008: Fig. 58: 12; Al-Kandari et al., 2009: p. 117, Pls. 71: B-F, 72: A, B; Al-Yamani & Saburova, 2011: Pl. 153: a-e.
Synonymized names:
Nitzschia neogena Grunow 1882
Cells are solitary. Valves are lanceolate, tapering gradually into produced apices, 86-91 μm long and 12-13
μm broad. Keeled raphe is central. Fibulae are moderately long, prolonged into transapical ribs, somewhat
unevenly spaced, 6-7 in 10 μm. Valve surface is delicately striated. Transapical striae are finely punctate, 18-
19 in 10 μm. Two large elongated and deeply lobed chloroplasts extend on either side of the central nucleus
toward the apices.
Distribution:
Nitzschia fluminensis is a marine species inhabiting mainly warm water coasts. It has been previously
reported from Kuwait’s waters (Al-Kandari et al., 2009) and intertidal sediments (Al-Yamani & Saburova,
2011).
Local occurrence:
Nitzschia fluminensis is a typically benthic species commonly observed in epipelic assemblages along
Kuwait’s shores (Al-Yamani & Saburova, 2011), and rarely occurred in Kuwait’s waters, being recorded
from 4.2% of all collected samples. The species was present throughout the year in low abundance with
higher concentrations (80-640 cells/l) recorded from the southern offshore waters around Kubbar Island.
Nitzschia hybrida Grunow

Plate 142: f-k
Péragallo & Péragallo, 1897-1908: Pl. LXX: 29; Witkowski et al., 2000: Pl. 191: 12-14; Al-Handal, 2009: Fig. 150; Al-
Yamani & Saburova, 2011: Pl. 154: a-d; Al-Handal et al., 2014: Fig. 92.
Cells are solitary. Frustules are linearly oblong in girdle view, slightly constricted in the middle, with rounded
corners. Valves are linear, with slightly arched margin opposite very weakly biarcuate side, with produced
and bent apices, 110-116 μm long and 6-7 μm broad. Keeled raphe is eccentric, marginal. Fibulae are small,
elongated, somewhat unevenly spaced, 12-13 in 10 μm, the median two being further apart to form a central
nodule. Central nodule is distinct, small. Valve surface is delicately striated. Transapical striae are very finely
punctate, parallel throughout the valve, 27-31 in 10 μm. Two large elongated irregularly shaped chloroplasts
Distribution:
Nitzschia hybrida is a marine to brackish water littoral species, primarily occurring in temperate regions.
It has been previously reported from the epiphytic assemblages in the Mesopotamian wetland, South
Iraq (Al-Handal et al., 2014), from the bottom sediments in the Shatt Al-Arab Estuary, Iraq (Al-Handal,
2009), and from Kuwait’s intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Nitzschia hybrida is a mud-dwelling species, commonly observed in great concentrations along Kuwait’s
northern shores around Bubiyan Island (Al-Yamani & Saburova, 2011), but also occurred rarely in
Kuwait’s neritic habitats, where was observed in low numbers.
Nitzschia lanceolata Smith

Plate 143: a-c
Péragallo & Péragallo, 1897-1908: Pl. LXXIII: 16-22; Foged, 1984: Pl. LVII: 5-7; Witkowski et al., 2000: Pl. 194: 1-5;
Al-Yamani & Saburova, 2011: Pl. 155: f-h.
Synonymized names:
Homoeocladia lanceolata (Smith) Kuntze 1898
Cells are solitary. Frustules are narrow, rectangular. Valves are narrowly lanceolate, with produced apices,
47-51 μm long and 6-7 μm broad. Keeled raphe is eccentric, marginal. Fibulae are unevenly spaced, 12 in 10
μm. Valve surface ornamentation is indiscernible with LM. Two irregularly shaped chloroplasts extend on
either side of the central nucleus toward the apices.
Distribution:
Nitzschia lanceolata is a marine to brackish water species widely distributed mainly in temperate areas. It
has been previously reported from Kuwait’s intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Nitzschia lanceolata sporadically occurred in Kuwait’s waters, and was observed in low numbers.

BACILLARIOPHYCEAE

a-c – Nitzschia distans: a – live cell in girdle view; b – acid cleaned frustule in girdle view; c – acid
cleaned valve; d, e – Nitzschia distans var. tumescens: part of live cell in oblique (d) and girdle (e)
view; f-k – Nitzschia hybrida: f – live cell in girdle view; g – acid cleaned valve; i – acid cleaned
frustule in girdle view; h, j, k – detailed parts of cleaned valves.
LM, BF illumination (a, d-f), DIC optic (b, i, j), and phase contrast (c, g, h, k); scale bars: a-c,
f-k – 10 μm; d, e – 20 μm.
Nitzschia longissima (Brébisson) Ralfs
Plate 143: i, j
Péragallo & Péragallo, 1897-1908: Pl. LXXIV: 20; Pavillard, 1925: Fig. 114; Allen & Cupp, 1935: Fig. 121; Cupp, 1943:
Fig. 154; Hendey, 1964: p. 283; Hendey, 1970: p. 157; Simonsen, 1974: p. 53; Foged, 1984: Pl. LVIII: 7, 8; Podzorski
& Håkansson, 1987: Pls. 44: 8, 45: 2; Witkowski et al., 2000: Pl. 207: 6, 7; Throndsen et al., 2007: p. 200; Hein et al.,
2008: Fig. 59: 1; Al-Kandari et al., 2009: p. 117, Pls. 72: C-F, 73: A, B; Hoppenrath et al., 2009: Fig. 47 j; Al-Yamani
& Saburova, 2011: Pl. 156: a-c.
Synonymized names:
Ceratoneis longissima Brébisson 1849
Nitzschia birostrata Smith 1853
Cells are long, weakly silicified, solitary. Valves are linearly lanceolate with very long rostrate apices, 216-225
μm long, 6-7 μm broad. Keeled raphe is eccentric, marginal. Fibulae are unevenly spaced, 5-6 in 10 μm, the
median two ones are distant. Central nodule is small, distinct. Valve surface ornamentation is indiscernible
with LM. Two elongate chloroplasts extend on either side of the central nucleus, not in cell apices.
Distribution:
Nitzschia longissima is a marine widespread littoral species, frequently found in the plankton. The species
has been previously reported from the Arabian Gulf (Simonsen, 1974), and from Kuwait’s shores
(Hendey, 1970), coastal waters (Al-Yamani et al., 2004; Al-Kandari et al., 2009), and intertidal sediments
Local occurrence:
Nitzschia longissima occasionally occurred in Kuwait’s waters, being recorded from 12.6% of all collected
period from June to November.
Nitzschia lorenziana Grunow

Plate 144: a-i
Péragallo & Péragallo, 1897-1908: Pl. LXXIV: 24, 25; Allen & Cupp, 1935: Fig. 123; Hendey, 1970: p. 158; Simonsen,
1974: Pl. 41: 4; Podzorski & Håkansson, 1987: Pl. 46: 1; Witkowski et al., 2000: Pls. 210: 24, 25, 211: 3, 212: 1-3; Al-
Kandari et al., 2009: p. 118, Pl. 73: C-I; Al-Yamani & Saburova, 2011: Pl. 157: a-f; Al-Handal et al., 2014: Fig. 97.
Synonymized names:
Nitzschia incurvata var. lorenziana Ross 1986
Cells are solitary. Frustules are linearly rectangular in girdle view, with truncate ends. Valves are linearly
lanceolate, sigmoid, gradually tapering into long rostrate apices, 87-160 μm long and 5-6 μm broad. Keeled
raphe is eccentric, marginal. Fibulae are evenly spaced, 7 in 10 μm, the median two ones are slightly distant.
Valve surface is delicately striated. Transapical striae are parallel, 20-21 in 10 μm. Two large bar-shaped
chloroplasts extend on either side of the central nucleus toward the apices.
Distribution:
Nitzschia lorenziana is a cosmopolitan and primarily benthic species, but is occasionally found in plankton.
It has been previously reported from the Indian Ocean (Simonsen, 1974), from the epiphytic assemblages
in the Mesopotamian wetland, South Iraq (Al-Handal et al., 2014), and from Kuwait’s shores (Hendey,
1970), coastal waters (Al-Yamani et al., 2004; Al-Kandari et al., 2009), and intertidal sediments (Al-
Local occurrence:
Nitzschia lorenziana commonly occurred in Kuwait’s waters, being recorded from 54% of all collected
concentrations (1.3-6.1∙104 cells/l) recorded in the inshore waters of Kuwait Bay and around Bubiyan
Island during the fall season in October and November.

BACILLARIOPHYCEAE

a-c – Nitzschia lanceolata: a, b – live cell in girdle view at different focal planes; c – acid cleaned
frustule in girdle view; d-h – Nitzschia reversa: d, e – live cells; f, g – acid cleaned frustules; h –
detail of valve showing fine areolation and central nodule (arrow); i, j – Nitzschia longissima:
live cells.
LM, BF illumination (a, b, d, e), DIC optic (i, j), and phase contrast (c, f); SEM (g, h); scale
bars: i, j – 20 μm; a-g – 10 μm; h – 5 μm.
Nitzschia pellucida Grunow
Plate 144: j-l
Witkowski et al., 2000: Pl. 191: 1-7, 11; Al-Yamani & Saburova, 2011: Pl. 159: e-h.
Cells are solitary. Frustules are linearly oblong in girdle view, slightly constricted in the middle, with rounded
corners, 52-56 μm long and 10-11 μm broad. Valves are linearly lanceolate, with slightly arched margin
opposite very weakly biarcuate side, with capitately produced apices. Keeled raphe is eccentric, marginal.
Fibulae are small, elongated, unevenly spaced, 13-15 in 10 μm, the median two being further apart to form a
central nodule. Central nodule is distinct, small. Valve surface ornamentation is indiscernible with LM. Two
large plate-like chloroplasts extend on either side of the central nucleus toward the apices.
Distribution:
Nitzschia pellucida is a marine littoral cosmopolitan species. It has been previously reported from Kuwait’s
Local occurrence:
Nitzschia pellucida is a typically benthic species commonly observed in epipelic assemblages along
Kuwait’s shores (Al-Yamani & Saburova, 2011), but sporadically occurred in Kuwait’s waters as well,
and in low numbers.
Nitzschia reversa Smith

Plate 143: d-h
Péragallo & Péragallo, 1897-1908: Pl. LXXIV: 19 (as Nitzschia longissima var. reversa); Witkowski et al., 2000: Pl. 210:
26, 27; Al-Yamani & Saburova, 2011: Pl. 160: a-d.
Synonymized names:
Nitzschia longissima var. reversa Grunow 1880
Cells are solitary. Valves are lanceolate, abruptly narrowed into long rostrate sigmoid apices, bent to opposite
sides, 137-151 μm long and 7-8 μm broad. Keeled raphe is eccentric, marginal. Fibulae are small, evenly
spaced, 14 in 10 μm, the median two ones are distant. Central nodule is small, distinct. Valve surface
ornamentation is indiscernible with LM. Transapical striae are very finely punctate, 52-53 in 10 μm as
discerned with SEM. Two elongated chloroplasts extend on either side of the central nucleus, not in cell
apices.
Distribution:
Nitzschia reversa is a marine cosmopolitan species, often occurs in the plankton. It has been previously
reported from Kuwait’s intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Nitzschia reversa occasionally occurred in Kuwait’s waters, being recorded from 7% of all collected samples.
The species was present throughout the year in low to moderate abundance with higher concentrations
(0.2-1.5∙104 cells/l) recorded in the inshore waters of Kuwait Bay and around Bubiyan Island during the
fall season from September to November.

BACILLARIOPHYCEAE

a-i – Nitzschia lorenziana: a-c – live cells in valve view; d-g – acid cleaned frustules; h – detailed part of
acid cleaned frustule; i – detailed central part of frustule with two central fibulae distant (arrowhead);
j-l – Nitzschia pellucida: j – live cell in girdle view; k, l – acid cleaned frustules in girdle view.
LM, BF illumination (a-c, j) and phase contrast (d-i, k, l); scale bars: a – 20 μm; b-h, j-l – 10 μm;
i – 5 μm.
Nitzschia sigma (Kützing) Smith
Plate 145: a-i
Péragallo & Péragallo, 1897-1908: Pl. LXXIV: 4-7; Allen & Cupp, 1935: Fig. 120; Hendey, 1964: Pl. XLII: 1; Hendey,
1970: p. 158; Simonsen, 1974: p. 55; Foged, 1984: Pls. XVI: 1, LIX: 5; Podzorski & Håkansson, 1987: Pl. 42: 2;
Witkowski et al., 2000: Pl. 206: 1-10; Hein et al., 2008: Fig. 60: 3; Al-Handal, 2009: Fig. 143; Al-Kandari et al., 2009:
p. 118, Pl. 74: A, B; Al-Yamani & Saburova, 2011: Pl. 161: a-g; Lobban et al., 2012: Pls. 2: 9, 10, 61: 4 (Nitzschia sigma
var. intercedens); Al-Handal et al., 2014: Fig. 91.
Synonymized names:
Synedra sigma Kützing 1844
Homoeocladia sigma (Kützing) Kuntze 1898
Sigmatella sigma (Kützing) Frenguelli 1923
Nitzschia sigma var. genuina Grunow 1878
Nitzschia sigma var. undulata Hohn et Hellerman 1966
Cells are long, solitary. Frustules are sigmoid in girdle view. Valves are long, linear-lanceolate and weakly
sigmoid, with long cuneate to opposite sides apices, 200-223 μm long and 8-11 μm broad. Keeled raphe
is eccentric, marginal. Fibulae are short, robust, evenly spaced, 7-8.5 in 10 μm. Valve surface is delicately
striated. Transapical striae are finely punctate, parallel, 28-30 in 10 μm. Two large plate-like chloroplasts
Remarks:
The extremely wide range of dimensions and structures’ density in Nitzschia sigma points to a heterogeneous
complex of different species under the name of this taxon, divided into many varieties by some authors
(Witkowski et al., 2000). In Kuwiat’s material, the largest strongly sigmoid specimens were assigned to
Nitzschia sigma var. intercedens Grunow (Pl. 145: c, d, h, i). The frustules were larger (270-320 μm long),
more strongly sigmoid and more finely striated (ca 30-32 in 10 μm), with closer fibulae (8-10 in 10 μm)
than the nominate variety.
Distribution:
Nitzschia sigma is a cosmopolitan benthic marine and extremely euryhaline species, often found in the
plankton. It has been previously reported from the Indian Ocean (Simonsen, 1974), from the epiphytic
assemblages in the Mesopotamian wetland, South Iraq (Al-Handal et al., 2014), from the bottom
sediments in the Shatt Al-Arab Estuary, Iraq (Al-Handal, 2009), and from Kuwait’s shores (Hendey,
1970), coastal waters (Al-Kandari et al., 2009), and intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Nitzschia sigma frequently occurred in Kuwait’s waters, being recorded from 38% of all collected samples.
The species was present throughout the year, more abundantly (2.0-4.0∙103 cells/l) in the inshore waters
of Kuwait Bay and around Bubiyan Island during the period from May to November.
Nitzschia sigmaformis Hustedt

Plate 145: j-m
Hustedt, 1955: Pl. 16: 2, 3; Simonsen, 1974: Pl. 40: 5; Witkowski et al., 2000: Pl. 206: 19, 20.
Cells are long, slender, solitary. Frustules are linearly lanceolate and sigmoid in girdle view, gradually tapering
into long protracted apices curved in opposite directions, 180-220 μm long and 6-8 μm broad. Keeled raphe
is eccentric, marginal. Fibulae are unevenly spaced, 6-8 in 10 μm, the median two ones are distant. Valve
surface is delicately striated. Transapical striae are parallel, 18-20 in 10 μm. Several small irregularly-shaped
chloroplasts are distributed through the cell length except the slender apices.
Distribution:
Nitzschia sigmaformis is a marine benthic species, but also found admixtured in the phytoplankton samples.
The species has been previously reported from the northern Arabian Gulf (Simonsen, 1974).

BACILLARIOPHYCEAE

a, b, e-g – Nitzschia sigma: a, b – live cells; e – detailed central part of cell; f – detailed
end of valve; g – part of acid cleaned frustule; c, d, h, i – Nitzschia sigma var. intercedens:
c – dividing live cell; d – detailed cell end; h – detailed part of acid cleaned frustule; i –
detailed central part of acid cleaned frustule; j-m – Nitzschia sigmaformis: j, k – live cells;
l – detailed part of acid cleaned frustule; m – detailed central part of acid cleaned frustule.
LM, BF illumination (a-e, j, k) and phase contrast (f-i, l, m); scale bars: a-d, j, k – 20 μm;
e-h, l – 10 μm; i, m – 5 μm.
Local occurrence:
Nitzschia sigmaformis frequently occurred in Kuwait’s waters, being recorded from 36% of all collected
concentrations (4.4-9.0∙103 cells/l) recorded in the inshore waters of Kuwait Bay during the fall season
in October. The species is recorded for the first time from Kuwait.
Nitzschia spathulata Brébisson

Plate 146: a-d
Péragallo & Péragallo, 1897-1908: Pl. LXXIII: 4, 5; Hendey, 1964: p. 281; Witkowski et al., 2000: p. 405; Hein et al.,
2008: Fig. 59: 5; Al-Yamani & Saburova, 2011: Pl. 163: a-e.
Cells are solitary. Frustules are linearly rectangular, slightly convex in the middle, with corners furnished
with small wind-like expansions and truncate ends. Valves are narrowly lanceolate with acutely rounded
apices, 44-53 μm long and 5-7 μm broad. Keeled raphe is central. Fibulae are unevenly spaced, 5.5-6.5 in 10
μm. Valve surface ornamentation is indiscernible with LM. Two large plate-like chloroplasts extend on either
side of the central nucleus toward the apices.
Distribution:
Nitzschia spathulata is a marine littoral species widely distributed mainly in temperate areas. It has been
previously reported from Kuwait’s intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Nitzschia spathulata is a typically benthic species commonly observed in epipelic assemblages along
Kuwait’s shores (Al-Yamani & Saburova, 2011), and occurred sporadically in Kuwait’s waters as well,
and in low numbers.
Nitzschia ventricosa Kitton

Plate 147: a-e
Ricard, 1977: Pl. 4: 15; Podzorski & Håkansson, 1987: Pl. 45: 1a-c (as Nitzschia longissima f. costata); Ricard, 1987: Figs.
1096-1098; Witkowski et al., 2000: Pl. 204: 8; Al-Yamani & Saburova, 2011: Pl. 156: d-g; Lobban et al., 2012: Pl. 62:
1, 2.
Synonymized names:
Nitzschia longissima (Brébisson) Ralfs f. costata Hustedt ex Schmidt 1921
Cells are long, solitary. Valves are lanceolate, abruptly narrowed into very long rostrate apices, 171-188 μm
long and 13 μm broad. Keeled raphe is eccentric, marginal. Fibulae are unevenly spaced, extend to form
filiform transverse costae, reaching the other valve margin, 6-8 in 10 μm. Central nodule is small, distinct.
Transapical striae are fine, 24-26 in 10 μm. Numerous small irregularly shaped chloroplasts are scattered
through the cell including slender apices.
Distribution:
Nitzschia ventricosa is a marine mainly littoral species. It has been commonly observed in Kuwait’s marine
sediments, and often in the composition of epiphytic assemblages on different substrata (Al-Yamani &
Saburova, 2011).
Local occurrence:
Nitzschia ventricosa is a typically benthic species commonly observed along Kuwait’s shores (Al-Yamani &
Saburova, 2011), and occurred sporadically in Kuwait’s waters as well, and in low numbers.

BACILLARIOPHYCEAE

a-d – Nitzschia spatulata: a – live cell in girdle view; b – acid cleaned frustule in valve view; c, d – acid cleaned
frustules in girdle view; e-j – Nitzschia sp. 1: e-g – live cell aggregated in dense irregularly shaped clusters;
h – detached cells; i, j – acid cleaned frustule in valve view.
LM, BF illumination (a, e-h) and phase contrast (b-d, i, j); scale bars: f – 50 μm; e, g, h – 20 μm; a-d – 10 μm;
i, j – 5 μm.
Nitzschia sp. 1
Plate 146: e-j
Cells are solitary or aggregated in dense irregularly shaped clusters. Valves are lanceolate in outline, gradually
tapering into produced and slightly capitate apices, 35-39 μm long and 3.5-4 μm broad. Keeled raphe is
eccentric, marginal. Fibulae are evenly spaced, 15-18 in 10 μm. Valve surface is finely striated. Transapical
striae are distinctly punctate, 29-30 in 10 μm. Two plate-like chloroplasts extend on either side of the central
nucleus toward the apices.
Local occurrence:
Nitzschia sp. 1 occurred sporadically in Kuwait’s waters, often forming dense irregularly shaped clusters,
or epiphytically on floating detritus.
Nitzschia sp. 2
Plate 147: g-i
Valves are lanceolate in outline, gradually tapering into produced and capitate apices, 23-62 μm long and
3-7 μm broad. Keeled raphe is eccentric, marginal. Fibulae are unevenly spaced, extend to form filiform
transverse costae, reaching the other valve margin, 12-13 in 10 μm. Transapical striae are indiscernible with
LM.
Local occurrence:
Frustules of Nitzschia sp. 2 were occasionally recorded from plankton hauls collected from Kuwait’s
waters, and observed in low numbers.
Genus Psammodictyon Mann
Psammodictyon panduriforme (Gregory) Mann

Plate 148: a-i
Péragallo & Péragallo, 1897-1908: Pl. LXX: 1-7, 11-13; Hendey, 1964: p. 279; Hendey, 1970: Pl. 5: 56; Simonsen, 1974:
p. 54; Foged, 1984: Pls. LVI: 4, 9, LVII: 1; Podzorski & Håkansson, 1987: Pl. 43: 10, 11; Ricard, 1987: Figs. 1029-1035;
Witkowski et al., 2000: Pls. 183: 6, 184: 13, 14, 186: 1-3; Al-Kandari et al., 2009: p. 118-119, Pl. 74: F, G (all as Nitzschia
panduriformis); Al-Handal, 2009: Fig. 142; Al-Yamani & Saburova, 2011: Pl. 165: a-f; Lobban et al., 2012: Pl. 62: 3, 4.
Synonymized names:
Nitzschia panduriformis Gregory 1857
Cells are solitary. Valves are broadly elliptical, constricted in the middle, with broadly cuneate apices, 67-84
μm long and 24-29 μm broad. Keeled raphe is eccentric, marginal. Fibulae are short, rib-like, evenly spaced,
8-9 in 10 μm, the median two ones are distant. Transapical striae are arranged in decussate lines of puncta,
15-16 in 10 μm, crossed by irregularly shaped longitudinal hyaline fold displaced slightly toward the keel
margin. Two large irregularly shaped deeply lobed chloroplasts extend on either side of the central nucleus,
appressed to the opposite valves.
Remarks:
According to Witkowski et al. (2000), the whole complex around Nitzschia panduriformis needs a general
taxonomic revision due to a wide range of variation in shape and striae density reported worldwide.
The smaller specimens of Psammodictyon panduriforme were observed in Kuwait’s material among the
nominate variety (Pl. 148: d-i). Specimens illustrated differ from the nominate variety by considerably
smaller size (17-25 μm long and 8-11 μm broad), striae density (24-30 in 10 μm), and closely spaced
fibulae (12-14 in 10 μm).

BACILLARIOPHYCEAE
Plate 147. Nitzschia spp. and Gomphotheca sinensis.

a-e – Nitzschia ventricosa: a – live cell; b – detailed central part of live cell; c-e – detailed central parts of
acid cleaned frustules; f – Nitzschia dissipata: acid cleaned frustule in valve view; g-i – Nitzschia sp. 2: acid
cleaned frustules; j-m – Gomphotheca sinensis: j-l – parts of acid cleaned frustules; m – acid cleaned frustule.
LM, BF illumination (a, b) and phase contrast (c-m); scale bars: m – 25 μm; j, k – 20 μm; a-e, l – 10 μm;
f-i – 5 μm.
Distribution:
Psammodictyon panduriforme is a marine widespread littoral species. It has been previously reported from
neritic plankton of the northern Arabian Gulf (Simonsen, 1974), from the bottom sediments in the Shatt
Al-Arab Estuary, Iraq (Al-Handal, 2009), and from Kuwait’s shores (Hendey, 1970), coastal waters (Al-
Yamani et al., 2004; Al-Kandari et al., 2009), and intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Psammodictyon panduriforme is of low occurrence in Kuwait’s coastal waters, being recorded from less
than 1% of all collected samples, and observed in low abundance.
Psammodictyon roridum (Giffen) Mann

Plate 148: j-l
Round et al., 1990: p. 676; Witkowski et al., 2000: Pl. 184: 9-12 (as Nitzschia rorida); Al-Yamani & Saburova, 2011: Pl.
167: a-c.
Synonymized names:
Nitzschia rorida Giffen 1975
Cells are solitary. Valves are linear-elliptic, weakly constricted in the middle, with shortly produced rostrate
apices, 28-38 μm long and 9-10 μm broad. Keeled raphe is eccentric, marginal. Fibulae are short, almost
evenly spaced, 10-11 in 10 μm, the median two ones are distant. Transapical striae are finely punctate,
arranged in decussate lines, 24 in 10 μm, interrupted by irregularly shaped longitudinal hyaline fold in apical
axis. Two large plate-like chloroplasts extend obliquely on either side of the central nucleus, appressed to the
opposite valves.
Distribution:
Psammodictyon roridum is a marine widely distributed littoral species. It has been previously reported
from Kuwait’s intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Psammodictyon roridum sporadically occurred in Kuwait’s coastal waters, being recorded from less than
1% of all collected samples, and observed in low abundance.
Genus Tryblionella Smith
Tryblionella coarctata (Grunow) Mann

Plate 149: a-c
Péragallo & Péragallo, 1897-1908: Pl. LXIX: 26-30 (as Nitzschia punctata var. coarctata); Hendey, 1964: p. 278 (as
Nitzschia punctata var. coarctata); Round et al., 1990: p. 678; Witkowski et al., 2000: Pls. 183: 13, 186: 4-13 (as Nitzschia
coarctata); Al-Handal, 2009: Fig. 147 (as Nitzschia cf coarctata); Al-Kandari et al., 2009: p. 117, Pls. 70: E, F, 71: A (as
Nitzschia coarctata); Al-Yamani & Saburova, 2011: Pl. 163: f-h.
Synonymized names:
Nitzschia coarctata Grunow 1880
Nitzschia punctata var. coarctata (Grunow) Hustedt 1921
Cells are solitary. Valves are elliptical, constricted in the middle, with broadly cuneate apices, 39-50 μm long
and 13-14 μm broad. Keeled raphe is eccentric, marginal. Number of fibulae is the same as of transapical
striae. Transapical striae are coarsely punctate, 14-15 in 10 μm, crossed by longitudinal fold in the apical axis.
Two large irregularly shaped deeply lobed chloroplasts extend on either side of the central nucleus toward
the apices, each lying beneath one valve face.

BACILLARIOPHYCEAE
Plate 148. Psammodictyon spp.

a-i – Psammodictyon panduriforme: a, b – live cells in valve view; c – acid cleaned frustule in valve
view; d – live small-sized cell in valve view; e-i – acid cleaned small-sized frustules in valve view; j-l –
Psammodictyon roridum: j – live cell in valve view; k, l – acid cleaned frustules in valve view.
LM, BF illumination (a, b, d, j) and phase contrast (c, e-g, k, l); SEM (h, i, l); scale bars: a-c – 10 μm;
d-l – 5 μm.
Remarks:
Tryblionella (Nitzschia) coarctata complex exhibits a wide range of variation in size, shape and striae
density and contains very likely some independent species or varieties (Witkowski et al., 2000). The
smaller specimens of Tryblionella coarctata were observed in Kuwait’s material among the nominate
variety, differed by smaller size (10-12 μm long and 5-6 μm broad) and striae density (28-30 in 10 μm).
Distribution:
Tryblionella coarctata is a marine to brackish water cosmopolitan species complex. It has been previously
reported from the bottom sediments in the Shatt Al-Arab Estuary, Iraq (Al-Handal, 2009), and from
Kuwait’s coastal waters (Al-Kandari et al., 2009) and intertidal sediments (Al-Yamani & Saburova,
2011).
Local occurrence:
Tryblionella coarctata sporadically occurred in Kuwait’s coastal waters, being recorded from less than 1%
of all collected samples, and observed in low abundance.
Tryblionella compressa (Bailey) Poulin

Plate 149: d-f
Witkowski et al., 2000: Pls. 182: 3, 4, 185: 18-21 (as Nitzschia compressa var. compressa).
Synonymized names:
Tryblionella punctata Smith 1853
Nitzschia punctata (Smith) Grunow 1880
Nitzschia compressa (Bailey) Boyer 1916
Valves are elliptic to broadly lanceolate with cuneate slightly produced apices, 18-27 μm long and 11-13 μm
broad. Keeled raphe is strongly eccentric, marginal. Number of fibulae is the same as of transapical striae,
14-18 in 10 μm. Striae are distinctly punctate.
Distribution:
Tryblionella compressa is a cosmopolitan marine to brackish water littoral species (Witkowski et al., 2000).
Local occurrence:
Frustules of Tryblionella compressa were sporadically found in plankton hauls from Kuwait’s coastal
waters, and observed in low numbers.

BACILLARIOPHYCEAE
Plate 149. Tryblionella spp.

a-c – Tryblionella coarctata: a – live cell in valve view; b, c – acid cleaned frustules in valve view; d-f –
Tryblionella compressa: d, e – acid cleaned frustules in valve view; f – external valve view.
LM, BF illumination (a) and phase contrast (b-e); SEM (f); scale bars: 5 μm.
Genus Pseudo-nitzschia Péragallo
The species belonging to the genus Pseudo-nitzschia are marine, planktonic diatoms with a worldwide
distribution. These species were previously included in Nitzschia given the formal rank of section, but
following morphological, physiological and genetic investigations of the type species of Nitzschia reinstated
Pseudo-nitzschia at the generic level (Mann, 1986; Hasle, 1994). Presently, the genus comprises nearly 50
species.
Pseudo-nitzschia species are characterized by narrowly lanceolate to fusiform and linear cells united into
stepped chains by overlap of valve ends. The shape and symmetry of the cells, cell dimensions, and the detailed
morphology of the frustules and girdle bands are important features for identification of Pseudo-nitzschia
species. Only some of these details can be seen in the LM, and electron microscopy is usually necessary
for species examination in order to be accurately identified (Hasle & Syvertsen, 1997; Larsen & Nguyen,
2004; Throndsen et al., 2007). Based on LM and limited SEM examination, five distinct morphotypes were
recognized within the Pseudo-nitzschia complex in Kuwait’s waters, however, further morphological studies
using electron microscopy need to be performed to clarify the taxonomy of Pseudo-nitzschia species complex
in Kuwait.
Harmful Effect:
Blooms of Pseudo-nitzschia occur regularly in the neritic habitats worldwide and produce a potent
neurotoxin, domoic acid, which can accumulate in shellfish, other invertebrates, and sometimes fish,
leading to illness and death in a variety of seabirds and marine mammals. Human consumption of
shellfish contaminated with domoic acid can result in Amnesic Shellfish Poisoning (ASP), which can be
life threatening. The first incident of ASP caused by the toxin (domoic acid) was documented in 1987 on
Prince Edward Island, Canada (Bates et al., 1989) coincidently with a bloom of Pseudo-nitzschia multiseries
(syn. Nitzschia pungens f. multiseries). During this outbreak, over 100 people were affected after consuming
contaminated mussels. Three people died, and others lost their short-term memory permanently.
Numerous blooms of Pseudo-nitzschia have since been characterized in coastal waters worldwide and
have been linked to a harmful effect to marine environment and human health (Hallegraeff et al., 1995;
Hasle & Syvertsen, 1997; Orsini et al., 2002; Larsen & Nguyen, 2004; Throndsen et al., 2007).
Pseudo-nitzschia americana (Hasle) Fryxell

Plate 150
Hasle, 1964: Pls. 1: 4, 14: 13-19; Hasle & Syvertsen, 1997: Pl. 73; Lundholm et al., 2002: Figs. 1-20; Ajani et al., 2013:
Fig. 2.
Synonymized names:
Nitzschia americana Hasle 1974
Cells are small, weakly silicified, and solitary or often occur epiphytically on Chaetoceros setae. Frustules are
nearly linear-rectangular in girdle view, with straight or slightly convex margins and truncated apices, 19-32
μm long and 2-4 μm broad. Valves are linear to slightly lanceolate, with obtusely rounded apices. Raphe is
eccentric, marginal. Fibulae are evenly spaced, 20-22 in 10 μm. There is no larger interspace between the
median two fibulae and no central nodule. Two plate-like chloroplasts extend on either side of the central
nucleus toward the apices.
Distribution:
Pseudo-nitzschia americana is a marine plankton species widely distributed from temperate to tropical
regions (Hasle & Syvertsen, 1997; Lundholm et al., 2002), often epiphytically on the setae of Chaetoceros
and Bacteriastrum, as well on some other diatoms.
Local occurrence:
Pseudo-nitzschia americana frequently occurred across Kuwait’s waters, being recorded from 38% of all
collected samples. The species was present all year round, more abundantly (0.9-1.3∙106 cells/l) in the
inshore waters of Kuwait Bay during the warm season from June to September, sometimes in bloom
proportions. The species is recorded for the first time from Kuwait.

BACILLARIOPHYCEAE
Plate 150. Pseudo-nitzschia americana.

a-e – live cells (arrowheads) attached to setae and chains of Chaetoceros spp.; f – detached cells; g-i – detailed
single live cells.
LM, BF illumination (a-g) and phase contrast (h, i); scale bars: a-c, e, f – 25 μm; d – 10 μm; g-i – 5 μm.
Pseudo-nitzschia cf. calliantha Lundholm, Moestrup et Hasle
Plate 151
Lundholm et al., 2003: Fig. 2 A-G; Larsen & Nguyen, 2004: Pl. VII: 5-8; Orlova et al., 2008: Figs. 3-9; Stonik et al.,
2011: Figs. 13-17; Ajani et al., 2013: Fig. 4; Fernandes et al., 2013: Figs. 10-13.
Cells are weakly silicified, united into stepped chains by overlapping the ends of sibling cells, with cells
overlap is roughly 1/5-1/6 of the total cell length. Valves are symmetrical in outline, linear with tapering
apices, 51-69 μm long and 1.9-2.2 μm broad. Fibulae are discernible in LM, regularly spaced, 18-20 in 10
μm. Central interspace is present, corresponds to 4-5 striae. Intersriae are barely discernible with LM, 32-36
in 10 μm. Striae contain one row of poroids. Poroids are round to squarish, 4.5-5 in 1 μm, split into several
sectors with characteristic flower-like pattern. Two elongate plate-like chloroplasts extend on either side of
the central nucleus toward the cell apices.
Remarks:
Despite the high morphological similarity among Pseudo-nitzschia species belonging to Pseudo-nitzschia
pseudodelicatissima complex, there are several features that distinguish species from each other, including
the width and shape of valves, density of fibulae and striae, structure of the poroid hymen and the
cingular bands (Lundholm et al., 2003). Morphology and morphometrics of Pseudo-nitzshia isolated
from Kuwait’s waters are consistent with Pseudo-nitzschia calliantha (Lundholm et al., 2003). The flower-
like pattern of poroid hymen observed in Kuwait’s specimens with SEM distinguishes this Pseudo-
nitzschia species from P. pseudodelicatissima and P. cuspidata with the poroid hymen divided into two parts.
Additionally, Kuwaiti Pseudo-nitzschia differs from P. caciantha in frustule shape (linear versus lanceolate)
and in striae density (32-36 versus 28-31 in 10 μm). These taxonomic characters allow to discriminate
the Kuwait’s material as P. calliantha, however, further study of the poroid structure by TEM is required
to verify the identification.
Distribution:
Pseudo-nitzschia calliantha is a recently described species from Ejby Harbor (Denmark) belonging to Pseudo-
nitzschia pseudodelicatissima complex (Lundholm et al., 2003). The species may have been misidentified
as P. pseudodelicatissima worldwide. Based on taxonomically and genetically verified records, P. calliantha
can be considered as a marine cosmopolitan and geographically widespread species (Lundholm et al.,
2003).
Local occurrence:
The occurrence of Pseudo-nitzschia cf. calliantha in Kuwait’s waters could not be defined due to high
morphological similarity among species within the Pseudo-nitzschia complex and the lack of reliable
differentiating criteria between these taxa during routine phytoplankton examination. The illustrated
specimens were isolated from the Kuwait’s southern inshore waters during the winter of 2008. Pseudo-
nitzschia cf. calliantha is reported for the first time from Kuwait’s waters.
Harmful Effect:
Pseudo-nitzschia calliantha is a toxigenic bloom-forming species. Production of domoic acid has been
detected in cultures isolated from the north Chile, Black Sea, and Mediterranean Sea, Tunisia (Besiktepe
at al., 2008; Álvares et al., 2009; Sahraoui et al., 2009). The species has been implicated in occurrence
of shellfish contamination in the Mediterranean (Quiroga, 2006), and found in bloom conditions in
southern Brazil (Fernandes et al., 2013).

BACILLARIOPHYCEAE
Plate 151. Pseudo-nitzschia cf. calliantha.

a – live cells united to chain showing contiguous areas of the sibling cells; b – single cell; c – acid
cleaned frustule; d – detailed frustule showing fibulae; e – collapsed frustule; f, g – detail of valve
showing central nodule and larger fibulae interspace; h – detail of valve surface showing flower-like
poroid pattern.
LM, BF illumination (a, b) and phase contrast (c, d); SEM (e-h); scale bars: a – 10 μm; b-f – 5 μm;
g – 2 μm; h – 1 μm.
Pseudo-nitzschia pungens (Grunow ex Cleve) Hasle
Plate 152
Cupp, 1943: Fig. 156 (as Nitzschia pungens); Simonsen, 1974: p. 54 (as Nitzschia pungens); Ricard, 1987: Figs. 1105, 1106
(as Nitzschia pungens); Hasle & Syvertsen, 1997: Pl. 69; Skov et al., 1999: Fig. 11 C-F, H; Larsen & Nguyen, 2004: Pl.
VIII: 1-3; Sar et al., 2006: Figs. 2, 3; Throndsen et al., 2007: p. 197; Hoppenrath et al., 2009: Fig. 47 e, f; Ajani et al.,
2013: Fig. 9.
Synonymized names:
Nitzschia pungens Grunow ex Cleve 1897
Cells are long, slender, heavily silicified, united into stepped chains by overlapping the ends of sibling cells,
with cells overlap is roughly 1/4 to 1/3 of the total cell length. Frustules are spindle-shaped in girdle view.
Valves are symmetrical in outline, linear to narrowly lanceolate, with distinctly pointed apices, 95-124 µm
long and 2.9-3.7 µm broad. Interstriae and fibulae are discernible in LM. Number of fibulae is almost the
same as of interstriae; there are 16-18 fibulae and 14-16 interstriae in 10 μm. Striae are biseriate, central
interspace is absent. Two elongated plate-like chloroplasts extend on either side of the central nucleus toward
the apices.
Distribution:
Pseudo-nitzschia pungens is a neritic cosmopolitan species (Hasle & Syvertsen, 1997). It has been previously
reported from the Arabian Gulf (Simonsen, 1974, as Nitzschia pungens) and from Kuwait’s waters (Al-
Yamani et al., 2004).
Local occurrence:
Pseudo-nitzschia pungens has common occurrence in Kuwait’s waters, being recorded from 50% of all
collected samples. The species was present all year round in low to moderate abundance with higher
concentrations (5.1-7.8∙104 cells/l) recorded in the inshore waters during the period from November to
February.
Harmful Effect:
Pseudo-nitzschia pungens is regarded to be a potentially toxic bloom-forming species (e.g., Bates et al.,
1989; Lundholm, 2017). Production of domoic acid has been detected in cultures isolated from the
west coast of USA and New Zealand (Bates & Douglas, 1993; Rhodes et al., 1998; Trainer et al., 1998;
Lundholm, 2017). Some strains of this species were reported as nontoxic or mildly toxic (Lelong et al.,
2012).

BACILLARIOPHYCEAE
Plate 152. Pseudo-nitzschia pungens.

a – live cells united to chain; b-d – detailed parts of chains showing contiguous areas of the sibling cells; e –
acid cleaned frustule; f – detail of apex; g-i – detailed central part of frustule.
LM, BF illumination (a-c) and phase contrast (e-g); SEM (d, h, i); scale bars: a – 50 μm; b-g –5 μm; h, i –2
μm.
Pseudo-nitzschia sp. 1
Plate 153
overlap is roughly 1/5 of the total cell length. Valves are symmetrical in outline, lanceolate, with pointed
apices, 73-91 µm long and 5.7-6.9 µm broad. Interstriae and fibulae are discernible in LM. There are 16-
17 fibulae in 10 µm, central interspace is present. Intersriae are 24-26 in 10 µm. Two elongated plate-like
chloroplasts extend on either side of the central nucleus toward the apices.
Remarks:
Based on LM observations of frustule morphology and morphometrics, Pseudo-nitzschia sp. 1 isolated
from Kuwait’s waters has affinity to Pseudo-nitzschia subfraudulenta (Hasle) Hasle (Hasle & Syvertsen,
1997: Pl. 70; Larsen & Nguyen, 2004: Pl. IX: 6-8; Rivera-Vilarelle et al., 2013: Figs. 21, 25; Teng et
al., 2013: Figs. 89-94). Pseudo-nitzschia subfraudulenta is characterized by a warm to temperate water
distribution (Hasle & Syvertsen, 1997) and was reported as a weakly toxic species and producer of
domoic acid in low concentrations (Teng et al., 2016). Pseudo-nitzschia sp. 1 is included as belonging to
Pseudo-nitzschia subfraudulenta with caution, until the morphology of this species is examined by SEM
and TEM in details, to verify their identity.
Local occurrence:
The occurrence of Pseudo-nitzschia sp. 1 aff. Pseudo-nitzschia subfraudulenta in Kuwait’s waters could not be
defined due to high morphological similarity among species within the Pseudo-nitzschia complex and the
lack of reliable differentiating criteria between these taxa during routine phytoplankton analysis.
Pseudo-nitzschia sp. 2
Plate 154
overlap is roughly 1/6 to 1/7 of the total cell length. Valves are symmetrical in outline, lanceolate, with
pointed and somewhat sigmoid apices, 65-83 µm long and 5.2-7.7 µm broad. There are 16-18 fibulae in 10
µm, central interspace is absent. Intersriae are 26-32 in 10 µm. Two elongate a plate-like chloroplasts extend
on either side of the central nucleus toward the apices.
Remarks:
Based on LM observations of frustule morphology and morphometrics, Pseudo-nitzschia sp. 2 isolated
from Kuwait’s waters has affinity to Pseudo-nitzschia multistriata (Takano) Takano (Larsen & Nguyen,
2004: Pl. 7: 1-4; Ajani et al., 2013: Fig. 8; Rivera-Vilarelle et al., 2013: Figs. 19, 22; Teng et al., 2013: Figs.
78-81). Pseudo-nitzschia multistriata is distributed from warm temperate to tropical waters, being reported
from the Mediterranean, East Asia, Australia and New Zealand (Guiry & Guiry, 2017; Karthick, 2017)
and has been reported as a bloom-forming toxigenic species. Strains of this species isolated from the
coastal Australian waters were found to be producers of domoic acid in considerable concentrations
(Ajani et al., 2013). Pseudo-nitzschia sp. 2 is included as belonging to Pseudo-nitzschia multistriata with
caution, until the morphology of this species is examined by SEM and TEM in details to verify their
identity.
Local occurrence:
The occurrence of Pseudo-nitzschia sp. 2 aff. Pseudo-nitzschia multistriata in Kuwait’s waters could not be
defined due to high morphological similarity among species within the Pseudo-nitzschia complex and the
lack of reliable differentiating criteria between these taxa during routine phytoplankton examination.

BACILLARIOPHYCEAE
Plate 153. Pseudo-nitzschia sp. 1 aff. Pseudo-nitzschia pseudofraundulenta.

a-c – live cells united to chains showing contiguous areas of the sibling cells; d, e – single cells; f, g – acid
cleaned frustules; h – detailed central part of frustule, arrowhead points to central interspace.
LM, BF illumination (a-e) and phase contrast (f-h); scale bars: a – 20 μm; b-g – 10 μm; h – 5 μm.
Plate 154. Pseudo-nitzschia sp. 2 aff. Pseudo-nitzschia multistriata.
a – two live cells united to chain showing contiguous areas of the sibling cells; b-d – single cells; e, f – acid
cleaned frustules; g – detailed end of frustule; h – detailed central part of frustule.
LM, BF illumination (a-d) and phase contrast (e-h); scale bars: a, b – 20 μm; c, d – 10 μm; e-h – 5 μm.

Order Surirellales Mann
Family Entomoneidaceae Reimer
Genus Entomoneis Ehrenberg
The species belonging to the genus Entomoneis are typically benthic and distinguished by the presence of
highly arched bilobate wings, numerous intercalary bands, some torsion along the apical axis, keeled raphe
system, and juncture of the keel with the valve body in form of a junction line. The size, shape of valve
and valve apices, the type and density of striae and areolae can be considered as important morphological
characters for species identification within the genus (Patrick & Reimer, 1975).
Entomoneis alata (Ehrenberg) Ehrenberg

Plate 155: a-c
Hendey, 1964: Pl. XXXIX: 14-16; Witkowski et al., 2000: Pl. 109: 21, 22; Bahls, 2012.
Synonymized names:
Navicula alata Ehrenberg 1840
Amphiprora alata (Ehrenberg) Kützing 1844
Cells are solitary. Frustules are rectangular in girdle view, constricted in the middle and twisted along the
apical axis, with four distinct marginal wings. Highly arched bilobate keels dominate the view in girdle
aspect. The keels are strongly torsioned so that only one lobe is in focus at one time. Valves are linear-elliptic
to linear lanceolate, with slightly concave sides and acute apices, 96-105 μm long and 15-18 μm broad. Axial
axis is raised to form S-shaped keel enclosing the raphe. Valve surface is striated. Striae on the valve surface
continue onto the keel. Transapical striae are coarsely areolated, parallel in the valve surface and radiate on
the keel, 14-16 in 10 µm. Areolae on the valve face are very fine, 26-30 in 10 µm within a stria. Areolae on
the keel are much more prominent, 12-15 in 10 µm. Junction line between the valve and keel is barely wavy,
marked by an irregular series of subequal dots and swellings. Girdle is composed of numerous narrow, finely
punctate intercalary bands.
Distribution:
Entomoneis alata is a widespread marine to brackish water species inhabiting the shallow coastal areas
worldwide (Hendey, 1964; Witkowski et al., 2000).
Local occurrence:
Frustules of Entomoneis alata were found in plankton net hauls from the inshore waters around Bubiyan
Island, and observed in low numbers. The species is reported for the first time from Kuwait’s waters.
BACILLARIOPHYCEAE
Entomoneis paludosa (Smith) Reimer
Plate 155: d, e
Péragallo & Péragallo, 1897-1908: Pl. XXXVIII: 12-20 (as Amphiprora paludosa); Hustedt, 1985: Fig. 624; Witkowski et
al., 2000: Pls. 109: 26, 27, 173: 8; Al-Yamani & Saburova, 2011: Pl. 169: a-c; Bahls, 2012.
Synonymized names:
Amphiprora paludosa Smith 1853
Cells are solitary. Frustules are rectangular in girdle view, strongly constricted in the middle, with biarcuated
sides, 43-61 μm long, 24-34 μm broad, with four distinct marginal wings. Valves are linear-lanceolate with
acute apices. Winged keel is elevated upon the valve surface, appearing slightly to strongly sigmoid in valve
view. Valve surface is finely striated, transapical striae punctate, parallel, 24-26 in 10 μm. Junction line between
the valve and keel is weakly biarcuated, contains a single swelling. Girdle is composed of numerous narrow,
finely punctate intercalary bands in form a crossing, sigmoid pattern. Single large elongate chloroplast with
wavy edges lies along the girdle.
Distribution:
Entomoneis paludosa is a cosmopolitan species of marine coasts (Witkowski et al., 2000). It has been
previously recorded in Kuwait’s coastal waters (Al-Yamani et al., 2004) and intertidal sediments (Al-
Local occurrence:
Entomoneis paludosa was found as an abundant epipelic form widely distributed in Kuwait’s marine
sediments (Al-Yamani & Saburova, 2011), but the species sporadically occurred in neritic habitats along
Kuwait’s coast as well, and observed in low numbers.
Entomoneis punctulata (Grunow) Osada et Kobayasi

Plate 155: f-k
Witkowski et al., 2000: Pl. 173: 4-7; Bahls, 2012.
Synonymized names:
Amphiprora paludosa var. punctulata Grunow 1880
Entomoneis paludosa var. punctulata (Grunow) Czarnecki et Reinke 1982
Cells are solitary, weakly silicified. Frustules are roughly rectangular in girdle view, slightly constricted in
the middle, with rounded corners and biarcuated sides, 32-41 μm long, 25-32 μm broad. Winged keel lobes
are broadly rounded and the keels are only slightly torsioned so that both lobes are more or less in focus at
the same time. Junction line between the valve and keel is short, curved and restricted to a terminal portion
of each lobe of the keel. Transapical striae are very delicate, finely punctate, 35-38 in 10 μm. Girdle is
composed of several narrow, slightly sigmoid, intercalary bands crossing at the cell poles.
Distribution:
Entomoneis punctulata is a marine to brackish water species inhabiting the littoral of temperate coastal
areas (Witkowski et al., 2000).
Local occurrence:
Frustules of Entomoneis punctulata were found in plankton net hauls from the inshore waters around
Bubiyan Island, and observed in low numbers. The species is reported for the first time from Kuwait’s
waters.
Family Surirellaceae Kützing

Genus Campylodiscus Ehrenberg ex Kützing
The species of the genus Campylodiscus are characterized by circular valves, often appearing elliptical or
dorsiventral due to the saddle-shaped frustules bent in girdle view, pennate structure in form of a median
rib or hyaline area subtending lateral striations, and raphe system that opens inwardly into a tubular canal
(canal-raphe) raised on a ridge or keel and running around the whole perimeter of the valve (Hendey, 1964).
Campylodiscus cf. fastuosus Ehrenberg

Plate 157: g
Péragallo & Péragallo 1897-1908: Pl. 57: 4-9 (as Campylodiscus thuretii); Stidolph et al., 2012: Pl. 21: 14, 6: 132 (the
latter as Campylodiscus thuretii); Lobban et al., 2012: Pl. 68: 4 (as Surirella scalaris); Lobban, 2015: Figs. 13, 14, 16, 18-20.
Synonymized names:
Campylodiscus parvulus Smith 1851
Campylodiscus thuretii Brébisson 1854
Campylodiscus simulans Gregory 1857
Cells are small, solitary, weakly silicified. Valves are nearly circular in outline, 23 μm in diameter. Median
area is narrow, almost linear, furnished with distinct transapical striae, 16 in 10 μm. Marginal infundibula are
faintly striated, dilated towards the margins, tapering towards the center, extending over half of the radius.

BACILLARIOPHYCEAE
Plate 155. Entomoneis spp. and Coronia daemeliana.

a-c – Entomoneis alata: acid cleaned frustules in valve view; d, e – Entomoneis paludosa: d – live cell
in girdle view; e – acid cleaned frustules in girdle view; f-k – Entomoneis punctulata: f, g – live cell in
girdle view; h – acid cleaned frustules in valve view; i – detail of collapsed frustule in girdle view; j, k –
frustules in girdle view; l – Coronia daemeliana: acid cleaned frustule in valve view;.
LM, BF illumination (f, g), DIC optic (d), and phase contrast (a-c, j-l); SEM (e, h, i); scale bars: l – 50
μm; a-h, j, k – 10 μm; i – 5 μm.
Remarks:
Valve view of Campylodiscus fastuosus is morphologically similar to Surirella scalaris Giffen, but differ in
more tapering shape of infundibula with a single prominent rib reaching the central area (Lobban, 2015),
however, this difference is hardly visible in small specimens in LM. The presented here isolated valve (Pl.
157: g) is consistent with the small specimen of Campylodiscus fastuosus (Lobban, 2015: Fig. 14) rather
than with Surirella scalaris (Lobban, 2015: Fig. 15). However, the complete frustules need to be examined
to determine the positional relationship of valves before being sure about Campylodiscus fastuosus from
Kuwait’s material since valves in Surirella scalaris are oriented in parallel while in Campylodiscus they are
arranged at right angles to one another (Lobban et al., 2012; Lobban, 2015).
Distribution:
Campylodiscus fastuosus is a marine littoral species distributed from warm temperate to tropical areas
(Guiry & Guiry, 2017). The species has been previously recorded in Kuwait’s intertidal sediments (Al-
Yamani & Saburova, 2011, as Surirella scalaris).
Local occurrence:
Frustules of Campylodiscus cf. fastuosus were found in low numbers in plankton net hauls collected from
Kuwait’s coastal waters.
Campylodiscus iyengarii Subrahmanyan

Plate 156: a
Desikachary & Prema, 1987: Pls. 253: 1, 352: 4.
Valves are circular in outline, slightly saddle-shaped, 72-75 μm in diameter. Valve surface is costate. Costae
are curved, 4.5-5 in 10 μm, radiating from the narrow lanceolate hyaline median area.
Distribution:
Campylodiscus iyengarii is reported from the coastal Indian waters (Desikachary & Prema, 1987).
Local occurrence:
Frustules of Campylodiscus iyengarii were found in low numbers in plankton net hauls collected from
Kuwait’s coastal waters. The species is reported for the first time from Kuwait.
Campylodiscus sp.
Plate 156: b
Cells are solitary. Frustules are bent in girdle view, saddle-shaped, 28 µm long and 22 µm broad. Valve
surface is furnished with costae. Costae are radiate, 6-7 in 10 µm, terminating in rhomboid internal openings
of the alar canal alongside the margin. The median area is hyaline, broadly lanceolate. Ridges inscribe the
median area, continuous from the central line to the valve margin. Valve surface is furnished with radiating
rows coarse areolae between costae, 24 in 10 µm.
Local occurrence:
Frustules of Campylodiscus sp. were found in low numbers in plankton net hauls collected from Kuwait’s
coastal waters.

BACILLARIOPHYCEAE
Plate 156. Campylodiscus spp., Petrodictyon gemma and Surirella fastuosa.

a – Campylodiscus iyengarii: acid cleaned frustule in valve view; b – Campylodiscus sp.: acid cleaned frustule
in valve view; c-g – Petrodictyon gemma: c – live cell in valve view; d – live cell in girdle view; e – acid
cleaned frustule in valve view; f, g – detailed parts of acid cleaned valves; h-j – Surirella fastuosa: h – live
cell in valve view; i – acid cleaned frustule in girdle view; j – acid cleaned frustule in valve view.
LM, BF illumination (c, d, h) and phase contrast (a, c, f, g, i, j); SEM (b); scale bars: a – 20 μm; c-f, h-j – 10
μm; b, g – 5 μm.
Genus Coronia (Ehrenberg ex Grunow) Ehrenberg
Coronia daemeliana (Grunow) Ruck et Guiry

Plate 155: l
Al-Handal, 2009: Fig. 217 (as Campylodiscus daemelianus); Ruck & Guiry, 2016.
Synonymized names:
Campylodiscus daemelianus Grunow 1874
Cells are large, solitary, with raised edges, 150-180 µm in diameter. Valves are suborbicular. Valve surface
is furnished with rows of coarse elongated puncta radiating from the median area to the valve margin and
interrupted by almost circular hyaline ring along the margin. Valve margin is striated with short marginal
costae. Median area is large, roughly rectangular, with irregularly scattered puncta.
Distribution:
Coronia daemeliana is a marine littoral species distributed from warm temperate to tropical areas (Guiry
& Guiry, 2017). It has been previously reported from the bottom sediments of the Shatt Al-Arab Estuary,
Iraq (Tynni, 1983; Al-Handal, 2009, as Campylodiscus daemelianus).
Local occurrence:
Frustules of Coronia daemeliana were found in low numbers in plankton net hauls from the inshore waters
around Bubiyan Island. The species is reported for the first time from Kuwait’s waters.
Genus Petrodictyon Mann
Petrodictyon gemma (Ehrenberg) Mann

Plate 156: c-g
Hendey, 1964: Pls. XL: 5, XLII: 4 (as Surirella gemma); Hendey, 1970: p. 158 (as Surirella gemma); Ricard, 1977: Pl. 7:
2 (as Surirella gemma var. ovata); Podzorski & Håkansson, 1987: Pl. 48: 3 (as Surirella gemma); Ricard, 1987: Figs. 963,
964 (as Surirella gemma); Round et al., 1990: p. 674; Witkowski et al., 2000: Pl. 216: 8, 9; Al-Handal, 2009: Fig. 209;
Al-Yamani & Saburova, 2011: Pl. 170: a-e.
Synonymized names:
Navicula gemma Ehrenberg 1840
Surirella gemma (Ehrenberg) Kützing 1844
Cells are solitary. Frustules are heteropolar in girdle view, wedge-shaped with truncate ends. Valves are almost
isopolar, elliptic with broadly rounded apices, 59-114 μm long and 28-51 μm broad. Narrow longitudinal
sternum lies along the apical axis. Valve surface is furnished with transapical costae and striated in between.
Costae are alternate and unevenly spaced, almost parallel in the middle, becoming radiate towards apices,
4-5.5 in 10 μm. Transapical striae are finely punctate, 19-20 in 10 μm. Two large plate-shaped chloroplasts
with crenulated edges lie against the valves.
Distribution:
Petrodictyon gemma is a marine littoral species, found sporadically in the plankton collections. It has been
previously reported from the bottom sediments of the Shatt Al-Arab Estuary, Iraq (Al-Handal, 2009),
and from Kuwait’s shores (Hendey, 1970, as Surirella gemma) and intertidal sediments (Al-Yamani &
Saburova, 2011).
Local occurrence:
Petrodictyon gemma occurred occasionally in Kuwait’s waters, being recorded from 8% of all collected
samples. The species was present throughout the year in low abundance with higher concentrations (78-
467 cells/l) recorded during the period from October to January.

Genus Surirella Turpin
The species belonging to the genus Surirella are common in benthic epipelic or epiphytic habitats, and found
occasionally in the plankton. The species are distinguished by isopolar or heteropolar frustules and the raphe
system positioned along entire valve margin within a canal, which may be raised above the valve surface in
some species (Cupp, 1943; Hendey, 1964).
Surirella comis Schmidt

Plate 157: e, f
Hendey, 1964: Pl. XL: 1.
Cells are solitary. Valves are slightly heteropolar, broadly elliptical in outline, with rounded apices, 48-54 μm
long and 31-34 μm broad. Raphe system is marginal, occupying the whole perimeter of the valve. Median
area is lanceolate, not reaching the apices, enclosed by two longitudinal bands of short striae. Marginal
infundibula are dilated towards the margins, tapering towards the center, extending over half of the radius,
ending into sparse prominent ribs converging towards the median area, about 2 in 10 μm. Two large plate-
like deeply lobed chloroplasts lie against the valves.
Distribution:
Surirella comis is a marine littoral mostly warm water species (Hendey, 1964).
Local occurrence:
Surirella comis is a typically benthic species, sporadically found in Kuwait’s coastal waters in low numbers.
The species is reported for the first time from Kuwait.
Surirella fastuosa (Ehrenberg) Kützing

Plates 156: h-j; 157: a-d
Péragallo & Péragallo, 1897-1908: Pls. LVIII: 1-7, LIX: 1-3; Hendey, 1964: Pl. XL: 4; Hendey, 1970: Pl. 5: 45 (as
Surirella recedens); Simonsen, 1974: p. 56; Tynni, 1983: Pl. 10: 103-105 (as Surirella recedens); Foged, 1984: Pl. LX: 3, 4;
Podzorski & Håkansson, 1987: Pls. 46: 10, 47: 2, 3, 48: 1, 55: 2, 3; Ricard, 1987: Figs. 951-961; Witkowski et al., 2000:
Pls. 215: 1-3, 216: 4; Hein et al., 2008: Figs. 62: 3, 64: 1; Al-Handal, 2009: Fig. 213; Al-Kandari et al., 2009: p. 121, Pl.
79: A-I; Al-Yamani & Saburova, 2011: Pl. 171: a-h.
Synonymized names:
Navicula fastuosa Ehrenberg 1840
Novilla fastuosa (Ehrenberg) Cleve 1868
Suriraya fastuosa (Ehrenberg) Schaarschmidt 1880
Surirella fastuosa var. genuina Cleve-Euler 1952
Campylodiscus neofastuosus Ruck, Nakov, Alverson et Theriot 2016
BACILLARIOPHYCEAE
Cells are solitary, cuneate. Frustules are wedge-shaped in girdle view. Valves are heteropolar, broadly ovate,
with one obtusely rounded apex and cuneately rounded opposite apex, 64-81 μm long and 37-47 μm broad.
Median area is hyaline, lanceolate. Raphe system is marginal, occupying the whole perimeter of the valve.
Marginal infundibula are dilated towards the margins, strongly narrowed towards the center, 2 in 10 μm.
Two large plate-like chloroplasts with lobed edges lie against the valves.
Distribution:
Surirella fastuosa is one of the most commonly reported marine species of Surirella worldwide, found
occasionally in plankton collections. The species has been previously reported from the bottom
sediments of the Shatt Al-Arab Estuary, Iraq (Tynni, 1983, as Surirella recedens; Al-Handal, 2009), and
from Kuwait’s shores (Hendey, 1970, as Surirella recedens), coastal waters (Al-Yamani et al., 2004; Al-
Kandari et al., 2009), and intertidal sediments (Al-Yamani & Saburova, 2011).
Local occurrence:
Surirella fastuosa has common occurrence in Kuwait’s waters, being recorded from 61% of all collected
samples. The species was present all year round, with more abundance (2.0-4.6∙103 cells/l) in the inshore
waters of Kuwait Bay during the warm season from May to October.
Plate 157. Surirella spp. and Campylodiscus cf. fastuosus.
a-d – Surirella fastuosa: a – external valve view; b – internal valve view; c - frustule in girdle
view; d – detail of valve in internal view; e, f – Surirella comis: live cells in valve view at different
focal planes; g – Campylodiscus cf. fastuosus: acid cleaned valve.
LM, BF illumination (e, f) and phase contrast (g); SEM (a-d); scale bars: e, f – 25 μm; a-d – 10
μm; g – 5 μm.

Genus Amphora sensu lato
The species belonging to the genus Amphora sensu lato are distinguished by asymmetric frustules shaped like
a segment of sphere (or wedge-shaped) so that their valves are not parallel with each other, but with surfaces
in planes at an angle one with another. The presence of an eccentric raphe displaced to the ventral side of the
valve face is a characteristic feature of Amphora species as well. The species within the genus are identified by
the size and shape of valve and valve apices, the shape of raphe, axial and central areas, the type and density
of striae and areolae (Hendey, 1964; Spaulding, 2011).
The historical approach to definition of the genus Amphora by overall frustule symmetry alone resulted
in a nonmonophyletic group that persisted for 150 years. With modern imaging and molecular techniques
employed over the past decades, many ‘amphoroid’ taxa, formerly classified within Amphora, have been
transferred to other newly created or existing genera.
Recent re-examination of the genus Tetramphora Mereschkowsky 1903 based on modern microscopic
techniques and molecular analysis has revealed that taxa with four chloroplasts, arranged in pairs linked by
large pyrenoids, aligning with Tetramphora are monophyletic and distinct from Amphora sensu stricto (Stepanek
& Kociolek, 2016). Based on newly obtained results, the genus Tetramphora Mereschkowsky was emended
with designation of Mereschkowsky’s T. ostrearia as the generitype, and consequently a range of Amphora
taxa were transfered to this genus. Moreover, based on the phylogenetic, chloroplast and structural evidence
available, the genus Tetramphora was replaced within the order Mastogloiales (Stepanek & Kociolek, 2016).
Order Mastogloiales Mann

Family Mastogloiaceae Mereschkowsky
Genus Tetramphora Mereschkowsky emend. Stepanek et Kociolek
The ‘amphoroid’ taxa currently included in Tetramphora are distinguished from other Amphora species by
strong ‘amphoroid’ frustule symmetry, moderately to strongly dorsiventral valve with a biarcuate raphe,
central hyaline area that creates undulate proximal raphe ends, slit-like external areola openings, axial costae,
protruding dorsal central thickening, and Mereschkowsky’s type 8 chloroplast (Mereschkowsky, 1903) with
two pairs of linked plastids (Stepanek & Kociolek, 2016).
Tetramphora decussata (Grunow) Stepanek et Kociolek

Plate 158: a-f
Péragallo & Péragallo, 1897-1908: Pl. XLIX: 23-25; Hendey, 1964: Pl. XXXVII: 9; Simonsen, 1974: p. 45; Al-Kandari
et al., 2009: p. 111, Pl. 63: D, E; Al-Yamani & Saburova, 2011: Pl. 128: a-g; Lobban et al., 2012: Pls. 1: 7-9, 54: 5, 55:
1-3 (all as Amphora decussata); Stepanek & Kociolek, 2016: p. 16.
BACILLARIOPHYCEAE
Synonymized names:
Amphora decussata Grunow 1877
Cells are large, solitary. Frustules are broadly lanceolate with truncate ends. Valves are flat, semilanceolate,
with convex dorsal margin, straight ventral margin and subacute apices, 55-102 μm long and 13-16 μm
broad. Raphe is biarcuated, eccentric, lying closer to ventral margin. Axial area is narrow, central area on the
dorsal valve side is in form of narrow transverse fascia reaching valve margin. Dorsal striae are composed of
elongated areolae arranged in decussating oblique rows, 16-22 in 10 μm. Ventral striae are short, composed
of single row of areolae. Girdle is composed of numerous striated intercalary bands. Two pairs of large,
often deeply lobed, plate-like chloroplasts with distinct bar-like pyrenoids extend from the centrally located
nucleus towards the cell apices. One pair of chloroplasts is dorsally appressed, the other one is ventrally
appressed (Mereschkowsky’s type 8 amphoroid chloroplast).
Distribution:
Tetramphora decussata is a marine benthic species distributed predominantly in warmer latitudes. It has
been previously recorded as Amphora decussata from the neritic habitats of the Arabian Gulf (Simonsen,
1974), and from Kuwait’s coastal waters (Al-Kandari et al., 2009) and intertidal sediments (Al-Yamani
& Saburova, 2011).
Local occurrence:
Tetramphora decussata occurred occasionally across Kuwait’s waters, being recorded from 13% of all
period from November to January.
Tetramphora lineolata (Ehrenberg) Mereschkowsky

Plate 158: g-l
Witkowski et al., 2000: Pl. 167: 17-19; Al-Yamani & Saburova, 2011: Pl. 132: a-f (as Amphora lineolata); Stepanek &
Kociolek, 2016: Figs. 9-21.
Synonymized names:
Amphora lineolata Ehrenberg 1838
Navicula lineolata Ehrenberg 1838
Amphora tenera Smith 1853
Frustulia rhomboides var. lineolata (Ehrenberg) Cleve 1894
Cells are solitary. Frustules are rectangular with broad truncate ends. Valves are narrow, semielliptical,
strongly dorsiventral, with almost straight to slightly convex ventral margin, smoothly arched dorsal margin,
and acutely rounded, slightly ventrally deflected apices, 49-58 μm long and 9-10 μm broad, frustule width
is 25-27 μm. Raphe is weakly biarcuated, eccentric. Axial area is narrow, with slight dilation at the central
area. Dorsal transapical striae are parallel in the middle, becoming weakly radiate towards the apices, 17-19
in 10 μm. Ventral striae are short, 17 in 10 μm. Girdle is composed of numerous striated intercalary bands,
striae are 16-18 in 10 μm. Two pairs of large, often with undulated edges, plate-like chloroplasts with distinct
bar-like pyrenoids extend from the centrally located nucleus towards the cell apices. One pair of chloroplasts
is dorsally appressed, the other one is ventrally appressed (Mereschkowsky’s type 8 amphoroid chloroplast).
Distribution:
Tetramphora lineolata is a widely distributed marine to brackish water littoral species (Witkowski et al.,
2000). It has been previously recorded as Amphora lineolata from Kuwait’s intertidal sediments (Al-
Local occurrence:
Tetramphora lineolata was commonly recorded along Kuwait’s shores in the composition of epiphytic
assemblages on macrophytes (Al-Yamani & Saburova, 2011), and rarely occurred in Kuwait’s waters,
being recorded from 1.5% of all collected samples. The species was present in low concentrations (100-
320 cells/l) in the neritic habitats during the fall season from September to November.
Tetramphora lineolatoides Spepanek et Kociolek

Plate 159: e, f
Stepanek & Kociolek, 2016: Figs. 68-82.
Cells are solitary. Frustules are rectangular with broad truncate ends. Valves are semielliptical, strongly
dorsiventral, with almost straight ventral margin, smoothly arched dorsal margin, and narrowly rounded,
slightly ventrally deflected apices, 38-46 μm long and 9-11 μm broad. Raphe is weakly biarcuated, eccentric,
close to ventral margin, with dorsally deflected distal raphe ends and proximal ends dipping towards the
ventral margin before terminating in simple endings. Axial area is narrow, with slight dilation at the central
area. Externally, areola openings consist of narrow longitudinally oriented slits arranged in striae. Dorsal
transapical striae are parallel througout, 17-18 in 10 μm. Ventral striae are very short, with 1-2 rows of
areolae. Girdle is composed of numerous smooth intercalary bands.
Distribution:
Tetramphora lineolatoides is a recently described species from Wakasa Bay, Japan (Stepanek & Kociolek,
2016).

BACILLARIOPHYCEAE
Plate 158. Tetramphora spp.

a-f – Tetramphora decussata: a-c – live cell in ventral (a), oblique (b) and dorsal (c) views; d
– acid cleaned frustule in valve view; e – detailed part of live cell in valve view; f – detailed
central part of acid cleaned valve; g-l – Tetramphora lineolata: g, j – cells in ventral view; h,
i – cell in dorsal view at different focal planes; k – acid cleaned frustule in ventral view; l –
detailed part of acid cleaned valve.
LM, BF illumination (a-c, e, g-j) and phase contrast (d, f, k, l); scale bars: a-e, g-k – 10 μm; f,
l – 5 μm.
Local occurrence:
Frustules of Tetramphora lineolatoides were found in low numbers in plankton net hauls from Kuwait’s
southern inshore locality in May 2008. The species is reported for the first time from Kuwait’s waters.
Tetramphora ostrearia (Brébisson) Mereschkowsky

Plate 159: a-d
Péragallo & Péragallo, 1897-1908: Pl. XLIX: 13-20; Hendey, 1964: Pls. XXXVII: 18, XXXVIII: 5, 11; Foged, 1984: Pls.
LII: 8, LIII: 4; Podzorski & Håkansson, 1987: Pl. 16: 1, 2; Ricard, 1987: Fig. 922; Witkowski et al., 2000: Pl. 170: 23;
Hein et al., 2008: Figs. 21: 2, 7, 22: 1; Al-Yamani & Saburova, 2011: Pl. 137: a-f (all as Amphora ostrearia var.); Stepanek
& Kociolek, 2016: Figs. 1-8.
Synonymized names:
Amphora ostrearia Brebisson ex Kützing 1849
Cells are solitary. Frustules are rectangular with rounded corner and truncate ends. Valves are semielliptical
and dorsiventral, with smoothly arched dorsal margin, straight to slightly convex in the middle ventral
margin, and narrowly rounded, deflected ventrally apices, 42-61 μm long and 8-11 μm broad, frustule
breadth is 24-29 μm. Raphe is biarcuated, eccentric, lying closer to ventral margin. Axial area is very narrow.
Central transverse thickening of internal virgae appears as narrow fascia in LM. Dorsal transapical striae are
distinctly punctate, parallel throughout or slightly radiate towards the apices, 13-16 in 10 μm. Ventral striae
short, 17-19 in 10 μm. Girdle is composed of numerous striated intercalary bands. Two pairs of large plate-
like chloroplasts with distinct bar-like pyrenoids extend from the centrally located nucleus towards the cell
apices. One pair of chloroplasts is dorsally appressed, the other one is ventrally appressed (Mereschkowsky’s
type 8 amphoroid chloroplast).
Distribution:
Tetramphora ostrearia is mainly a littoral species, but is sporadically found in the plankton. The species
has been previously reported from Kuwait’s waters (Al-Kandari et al., 2009) and intertidal sediments
Local occurrence:
Tetramphora ostrearia occasionally occurred in Kuwait’s waters, being recorded from 9.6% of all collected
concentrations (0.4-1.3∙103 cells/l) during the period from March to July.
Order Thalassiophysales Mann

Family Catenulaceae Mereschkowsky
Genus Amphora Ehrenberg ex Kützing
Amphora hyalina Kützing

Plate 159: g-i
Hendey, 1964: Pl. XXXVII: 10; Witkowski et al., 2000: Pl. 163: 22; Al-Yamani & Saburova, 2011: Pl. 180: h-k.
Synonymized names:
Amphora hemisphaerica Grunow 1867
Cells are solitary, weakly silicified. Frustules are broadly elliptical, with subtruncate ends. Valves are
semicircular, with convex dorsal margin and nearly straight ventral margin, and narrowly rounded, deflected
ventrally apices 77-89 μm long and 16-19 μm broad. Raphe is straight, closely appressed to ventral margin.
Axial and central areas are absent. Transapical striae are delicate, parallel, hardly visible with LM, 22-25 in
10 μm. Two large irregularly shaped plate-like chloroplasts lie against the valves.
Distribution:
Amphora hyalina is a marine littoral widespread species, frequently found in estuarine and neritic habitats
(Hendey, 1964; Witkowski et al., 2000). The species has been previously reported from Kuwait’s intertidal

BACILLARIOPHYCEAE
Plate 159. Tetramphora spp. and Amphora hyalina.

a-d – Tetramphora ostrearia: a – live cell in dorsal view; b – acid cleaned frustule in ventral
view; c – acid cleaned valve; d – internal valve view; e, f – Tetramphora lineolatoides: e –
external oblique valve view; f – detail of valve center; g-i – Amphora hyalina: g, h – live cells in
ventral view; i – live cell in dorsal view.
LM, BF illumination (a, g-i) and phase contrast (b, c); SEM (d-f); scale bars: g-i – 20 μm; a-e
– 10 μm; f – 5 μm.
Local occurrence:
Amphora hyalina sporadically occurred in the coastal Kuwait’s waters, and was observed in low numbers.
Amphora laevis Gregory

Plates 160, 161: a, b
Gregory, 1857: Pl. XII: 74; Hendey, 1964: p. 267; Witkowski et al., 2000: Pl. 166: 23.
Synonymized names:
Amphora nobilis Flogel 1873
Cells are solitary, weakly silicified, occur epiphytically on Chaetoceros or loricae of living tintinnid ciliates.
Frustules are broadly elliptical, with subtruncate ends. Valves are semilanceolate, with convex dorsal margin,
nearly straight ventral margin, and with subrostrate, deflected ventrally apices, 30-38 μm long and 9-11 μm
broad. Raphe is biarcuate, eccentric, lying closer to ventral margin. Axial area is absent, central area is
transversely expanded to form a distinct narrow fascia. Transapical striae are delicate, parallel, hardly visible
with LM, 20-22 in 10 μm. Girdle is composed of numerous narrow intercalary bands. Two large deeply
lobed plate-like chloroplasts expand on either side of the central nucleus toward the apices, lying against the
girdle.
Distribution:
Amphora laevis is a marine species distributed from temperate to warm water areas (Guiry & Guiry,
2017).
Local occurrence:
Amphora laevis occurred sporadically in the phytoplankton of Kuwait’s waters, being attached to the setae
of large-sized chain-forming diatom Chaetoceros (Pl. 160: a, b) or to the outer wall of lorica of tintinnid
ciliates (Pl. 160: c-h). The species is reported for the first time in Kuwait.
Family Thalassiophysaceae Mann

Genus Thalassiophysa Conger
Thalassiophysa hyalina (Greville) Paddock et Sims

Plate 161: c-g
Round et al., 1990: p. 606; Hein et al., 2008: Pls. 60: 6, 61: 1; Al-Yamani & Saburova, 2011: Pl. 180: c, d; Lobban et
al., 2012: Pls. 2: 1-8, 58: 1.
Synonymized names:
Amphiprora hyalina Greville 1865
Thalassiophysa rhipidis Conger 1954
Proboscidea insecta (Grunow ex Schmidt) Paddock et Sims 1980
Cells are solitary, weakly silicified, and strongly dorsiventral. Frustules are broadly elliptical, with truncate
ends, 79-90 μm long and 47-53 μm broad. Valves are compressed, with arched dorsal margin and nearly
straight ventral margin. Raphe system is fibulate. Raphe is keeled, strongly biarcuate, curving into the ventral
side near the center to form a characteristic funnel-shaped central notch. Girdle is composed of numerous
narrow bands. Valve striation is not resolved with LM. A single large plate-like chloroplast lies against the
ventral side of the girdle, connecting with the cell edges by numerous cytoplasmic threads.
Distribution:
Thalassiophysa hyalina is a marine littoral species distributed in temperate to tropical waters (Round et
al., 1990). The species has been previously reported from Kuwait’s intertidal sediments (Al-Yamani &
Saburova, 2011).
Local occurrence:
Thalassiophysa hyalina is of rare occurrence in Kuwait’s waters, being recorded from 3% of all collected
samples. The species was associated mostly with the northern waters, and usually was present in low
abundance with higher concentration (3.1∙103 cells/l) in Kuwait Bay during the summer season.

BACILLARIOPHYCEAE
Plate 160. Amphora laevis.

a – live cells attached to chain of Chaetoceros teres; b – live cells attached to seta of Chaetoceros peruvianus; c-f
– live cells attached to loricae of tintinnid ciliates; g, h – detailed tintinnid lorica with attached cells.
LM, BF illumination (a, b) and DIC optic (c-h); scale bars: a – 50 μm; b-h – 10 μm.
Plate 161. Amphora laevis, Thalassiophysa hyalina, and Seminavis robusta.
a, b – Amphora laevis: a – group of live cells detached from tintinnid lorica; b – two detached cells in
ventral (bottom) and valve (top) views, arrowheads point to eccentric biarcuate raphe; c-g – Thalassiophysa
hyalina: live cells at different focal planes; h, i – Seminavis robusta: h – live cell in valve view; i – acid
cleaned valve.
LM, BF illumination (c-h) and DIC optic (a, b, i); scale bars: a, c-g – 20 μm; b, h, i – 10 μm.

BACILLARIOPHYCEAE
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Index to taxa
A
Actinocyclus curvatulus – p. 64, Pl. 27: b-h Chaetoceros peruvianus f. gracilis – p. 120, Pl. 59: a-c
Actinocyclus exiguus – p. 66, Pl. 29: a-d Chaetoceros protuberans – p. 123, Pl. 50: f-i
Actinocyclus octonarius – p. 66, Pl. 28: a-j Chaetoceros pseudobrevis – p. 123, Pl. 57: d, e
Actinocyclus octonarius var. crassus – p. 68, Pl. 28: e, f Chaetoceros pseudocurvisetus – p. 124, Pl. 60
Actinocyclus octonarius var. ralfsii – p. 68, Pl. 28: a-d Chaetoceros seiracanthus – p. 124, Pl. 59: e-g
Actinocyclus octonarius var. tenellus – p. 69, Pl. 28: g-j Chaetoceros socialis – p. 126, Pl. 61
Actinocyclus subtilis – p. 69, Pl. 28: k-m Chaetoceros subtilis var. abnormis – p. 126, Pl. 39: a, b
Actinocyclus sp. – p. 70, Pl. 29: e-g Chaetoceros tenuissimus – p. 128, Pl. 59: d
Actinoptychus senarius – p. 62, Pl. 27: a Chaetoceros teres – p. 128, Pl. 62
Amphiprora angustata – p. 267, Pl. 137: j-l Chaetoceros tortissimus – p. 130, Pl. 63
Amphiprora gigantea var. sulcata – p. 267, Pl. 137: a-i Chaetoceros vanheurckii – p. 130, Pl. 54: h-k
Amphora hyalina – p. 316, Pl. 159: g-i Climacodium frauenfeldianum – p. 141, Pl. 69: a-d
Amphora laevis – p. 318, Pl. 160, Pl. 161: a, b Climaconeis lorenzii – p. 269, Pl. 138: a
Anorthoneis excentrica – p. 222, Pl. 118: h Climaconeis undulata – p. 270, Pl. 138: b-d, g-j
Asterionellopsis glacialis – p. 213, Pl. 112 Climaconeis sp. – p. 270, Pl. 138: e, f
Asteromphalus cleveanus – p. 40, Pl. 13 Cocconeis coronatoides – p. 222, Pl. 117: j
Asteromphalus flabellatus – p. 40, Pl. 14: a-c Cocconeis neothumensis var. marina – p. 224, Pl. 118: e
Asteromphalus heptactis – p. 42, Pl. 15 Cocconeis placentula var. euglypta – p. 226, Pl. 118: a-d
Asteromphalus sarcophagus – p. 42, Pl. 14: d-f Cocconeis scutellum var. parva – p. 226, Pl. 118: f, g
Attheya decora – p. 76, Pl. 32: f Cocconeis scutellum var. posidoniae – p. 226, Pl. 117: h
B Cocconeis scutellum var. scutellum – p. 226, Pl. 117: a-g, i
Bacillaria paxillifera – p. 271, Pl. 139 Corethron histrix – p. 38, Pl. 12
Bacteriastrum comosum var. hispidum – p. 86, Pl. 35 Coronia daemeliana – p. 310, Pl. 155: l
Bacteriastrum delicatulum – p. 86, Pl. 36 Coscinodiscopsis jonesiana – p. 59, Pl. 24; Pl. 25
Bacteriastrum furcatum – p. 89, Pl. 37 Coscinodiscus asteromphalus – p. 46, Pl. 16
Bacteriastrum hyalinum – p. 89, Pl. 38 Coscinodiscus gigas – p. 48, Pl. 17
Bacteriastrum hyalinum var. princeps – p. 89, Pl. 38: e-j Coscinodiscus granii – p. 48, Pl. 18
Bellerochea horologicalis – p. 76, Pl. 31: a-d Coscinodiscus janischii var. arafurensis – p. 51, Pl. 19
Bellerochea malleus – p. 77, Pl. 31: e-g Coscinodiscus marginatus – p. 51, Pl. 20: a-f
Bellerochea cf. yucatanensis – p. 77, Pl. 31: h-k Coscinodiscus nobilis – p. 54, Pl. 21
Biddulphia pulchella – p. 78, Pl. 32: a Coscinodiscus oculus-iridis – p. 54, Pl. 22
Biddulphia tuomeyi – p. 78, Pl. 32: b-e Coscinodiscus perforatus – p. 57, Pl. 20: g-k
C Coscinodiscus radiatus – p. 57, Pl. 23
Caloneis elongata – p. 192, Pl. 97: e, f Cyclotella atomus var. atomus – p. 156, Pl. 77: d-f
Campylodiscus cf. fastuosus – p. 306, Pl. 157: g Cyclotella atomus var. marina – p. 156, Pl. 77: g-i
Campylodiscus iyengarii – p. 308, Pl. 156: a Cyclotella choctawhatcheeana – p. 158, Pl. 78
Campylodiscus sp. – p. 3082, Pl. 156: b Cyclotella cubiculata – p. 161, Pl. 80
Cerataulina bicornis – p. 136, Pl. 66 Cyclotella litoralis – p. 158, Pl. 79
Cerataulina dentata – p. 136, Pl. 67 Cyclotella stylorum – p. 161, Pl. 81
Cerataulina pelagica – p. 137, Pl. 68 Cymatotheca minima – p. 170, Pl. 86
Chaetoceros affinis – p. 92, Pl. 39: c-g; Pl. 40: a-c Cylindrotheca closterium – p. 271, Pl. 140: a-e
Chaetoceros brevis – p. 94, Pl. 40: d-f; Pl. 41 Cymatonitzschia marina – p. 273, Pl. 140: f. g
Chaetoceros coarctatus – p. 94, Pl. 42 Cymatosira lorenziana – p. 144, Pl. 71: a-c
Chaetoceros compressus – p. 98, Pl. 43 D
Chaetoceros costatus – p. 98, Pl. 44 Dactyliosolen phuketensis – p. 19, Pl. 2: g, h
Chaetoceros curvisetus – p. 101, Pl. 45 Delphineis surirella – p. 213, Pl. 113: e
Chaetoceros danicus – p. 101, Pl. 46 Detonula pumila – p. 172, Pl. 87
Chaetoceros decipiens – p. 104, Pl. 47 Diplomenora cocconeiformis – p. 215, Pl. 113: a-c
Chaetoceros denticulatus – p. 104, Pl. 48; Pl. 49: a-c Diploneis weissflogii – p. 231, Pl. 119: g-i
Chaetoceros diadema – p. 108, Pl. 49: d-g Ditylim brightwellii – p. 153, Pl. 75
Chaetoceros didymus – p. 108, Pl. 50: a-e Ditylim sol – p. 153, Pl. 76
Chaetoceros distans – p. 110, Pl. 51 E
Chaetoceros diversus – p. 110, Pl. 52 Entomoneis alata – p. 305, Pl. 155: a-c
Chaetoceros laciniosus – p. 113, Pl. 53 Entomoneis paludosa – p. 305, Pl. 155: d, e
Chaetoceros lauderi – p. 113, Pl. 54: a-g Entomoneis punctulata – p. 306, Pl. 155: f-k
Chaetoceros lorenzianus – p. 116, Pl. 55; Pl. 56 Ephemera planamembranacea – p. 246, Pl. 125: a-d
Chaetoceros paradoxus – p. 116, Pl. 57: a-c Eucampia cornuta – p. 79, Pl. 32: g-l
Chaetoceros peruvianus – p. 120, Pl. 58 Eucampia zodiacus – p. 82, Pl. 33
G
Giffenia cocconeiformis – p. 274, Pl. 140: h-k Petroneis marina – p. 230, Pl. 119: l-n
Gomphotheca sinensis – p. 274, Pl. 147: j-m Plagiolemma confusa – p. 246, Pl. 125: e-i
Grammatophora oceanica – p. 218, Pl. 114 Plagiotropis lepidoptera – p. 248, Pl. 127: a-f
Guinardia delicatula – p. 19, Pl. 2: a-f Plagiotropis tayrecta – p. 250, Pl. 127: g-i
Guinardia flaccida – p. 21, Pl. 3 Planktoniella sol – p. 172, Pl. 88
Guinardia striata – p. 23, Pl. 4 Pleurosigma decorum – p. 250, Pl. 128: a-f; Pl. 130: a-c
Gyrosigma balticum – p. 232, Pl. 120 Pleurosigma diversestriatum – p. 252, Pl. 128: g-k; Pl. 129
Gyrosigma tenuissimum – p. 235, Pl. 123: a-d Pleurosigma elongatum – p. 256, Pl. 131: a-f
H Pleurosigma falx – p. 256, Pl. 131: g-k
Haslea gigantea – p. 235, Pl. 121 Pleurosigma formosum – p. 258, Pl. 132
Haslea howeana – p. 236, Pl. 122: a-d Pleurosigma inflatum – p. 258, Pl. 130: d-h; Pl. 133: a-d
Haslea cf. meteorou – p. 238, Pl. 122: f-h Pleurosigma inscriptura – p. 260, Pl. 133: e-h
Haslea wawrikae – p. 238, Pl. 122: i, j Pleurosigma intermedium – p. 260, Pl. 134: a-e
Helicotheca tamesis – p. 82, Pl. 34: a-f Pleurosigma javanicum – p. 262, Pl. 135
Hemiaulus hauckii – p. 142, Pl. 69: e-g Pleurosigma speciosum var. javanicum – p. 262, Pl. 134: f-j
Hemiaulus membranaceus – p. 142, Pl. 69: h-k Pleurosigma strigosum – p. 265, Pl. 136: a-f
Hemiaulus sinensis (chinensis) – p. 144, Pl. 70 Podosira stelligera – p. 74, Pl. 30: c-e
Hemidiscus cuneiformis – p. 72, Pl. 30: a Porosira pentaportula – p. 175, Pl. 89; Pl. 90
L Proboscia alata – p. 16, Pl. 1: a, b
Lauderia annulata – p. 164, Pl. 82; Pl. 83 Proboscia indica – p. 17, Pl. 1: c-i
Leptocylindrus danicus – p. 132, Pl. 64: a-f Psammodictyon panduriforme – p. 290, Pl. 148: a-i
Leptocylindrus mediterraneus – p. 132, Pl. 65 Psammodictyon roridum – p. 292, Pl. 148: j-l
Leptocylindrus minimus – p. 133, Pl. 64: g-l Pseudo-nitzschia americana – p. 296, Pl. 150
Licmophora abbreviata – p. 216, Pl. 113: f-l Pseudo-nitzschia cf. calliantha – p. 298, Pl. 151
Lyrella abrupta – p. 228, Pl. 119: a-d Pseudo-nitzschia pungens – p. 300, Pl. 152
Lyrella lyroides – p. 229, Pl. 118: i-k Pseudo-nitzschia sp. 1 aff. P. subfraudulenta – p. 302, Pl. 153
M Pseudo-nitzschia sp. 2 aff. P. multistriata – p. 302, Pl. 154
Mastogloia rostrata – p. 230, Pl. 118: l Pseudosolenia calcar-avis – p. 26, Pl. 5: e, f
Mastoneis biformis – p. 240, Pl. 123: e-h R
Meuniera membranacea – p. 248, Pl. 126 Rhizosolenia bergonii – p. 27, Pl. 6: a-d
N Rhizosolenia cochlea – p. 27, Pl. 6: e-g
Navicula directa – p. 240, Pl. 123: i-k Rhizosolenia hyalina – p. 29, Pl. 7
Navicula palpebralis – p. 241, Pl. 123: p-n Rhizosolenia imbricata – p. 29, Pl. 8; 9: e-h
Neocalyptrella robusta – p. 23, Pl. 5: a-d Rhizosolenia setigera – p. 32, Pl. 9: a-d
Nitzschia amabilis – p. 276, Pl. 141: i-o Roperia tesselata – p. 70, Pl. 29: h, i
Nitzschia bicapitata – p. 276, Pl. 141: a-e S
Nitzschia braarudii – p. 278, Pl. 141: f Schuettia annulata var. minor – p. 45, Pl. 10: f, g
Nitzschia dissipata – p. 278, Pl. 147: f Seminavis robusta – p. 241, Pl. 161: h, i
Nitzschia distans – p. 279, Pl. 142: a-c Shionodiscus oestrupii var. venrickiae – p. 175, Pl. 91
Nitzschia distans var. tumescens – p. 279, Pl. 142: d. e Skeletonema grevillei – p. 167, Pl. 84; Pl. 85
Nitzschia fluminensis – p. 279, Pl. 141: g, h Stellarima stellaris – p. 72, Pl. 30: b
Nitzschia hybrida – p. 280, Pl. 142: f-k Stephanopyxis palmeriana – p. 36, Pl. 11
Nitzschia lanceolata – p. 280, Pl. 143: a-c Streptotheca indica – p. 84, Pl. 34: g-i
Nitzschia longissima – p. 282, Pl. 143: g-i Surirella comis – p. 311, Pl. 157: e, f
Nitzschia lorenziana – p. 282, Pl. 144: a-i Surirella fastuosa – p. 3115, Pl. 156: h-j: Pl. 157: a-d
Nitzschia pellucida – p. 284, Pl. 144: j-l T
Nitzschia reversa – p. 284, Pl. 143: d-h Tetramphora decussata – p. 313, Pl. 158: a-f
Nitzschia sigma – p. 286, Pl. 145: a, b, e-g Tetramphora lineolata – p. 313, Pl. 158: g-l
Nitzschia sigma var. intercedens – p. 286, Pl. 145: c, d, h, i Tetramphora lineolatoides – p. 314, Pl. 159: e, f
Nitzschia sigmaformis – p. 286, Pl. 145: j-m Tetramphora ostrearia – p. 316, Pl. 159: a-d
Nitzschia spathulata – p. 288, Pl. 146: a-d Thalassionema frauenfeldii – p. 218, Pl. 115
Nitzschia ventricosa – p. 288, Pl. 147: a-e Thalassionema nitzschioides – p. 221, Pl. 116: e-i
Nitzschia spp. – p. 290, Pl. 146: e-j; Pl. 147: g-i Thalassionema pseudonitzschioides – p. 221, Pl. 116: j, k
O Thalassiophysa hyalina – p. 318, Pl. 161: c-g
Odontella aurita – p. 146, Pl. 71: d-g; Pl. 72 Thalassiosira andamanica – p. 179, Pl. 92
Odontella weissflogii – p. 149, Pl. 71: h-j Thalassiosira cathariensis – p. 181, Pl. 93
P Thalassiosira cf. concava – p. 181, Pl. 94
Palmerina hardmaniana – p. 62, Pl. 26 Thalassiosira concaviuscula – p. 184, Pl. 95
Paralia sulcata – p. 75, Pl. 30: f-j Thalassiosira decipiens – p. 186, Pl. 96: a-i
Perissonoë cruciata – p. 215, Pl. 113: d Thalassiosira delicatula – p. 188, Pl. 97
Petrodictyon gemma – p. 310, Pl. 156: c-g Thalassiosira diporocyclus – p. 190, Pl. 98
Petroneis granulata – p. 229, Pl. 119: j, k Thalassiosira eccentrica – p. 190, Pl. 99

Thalassiosira exigua – p. 192, Pl. 100
Thalassiosira hendeyi – p. 195, Pl. 101; Pl. 102
Thalassiosira lineata – p. 199, Pl. 103: a-c
Thalassiosira lundiana – p. 199, Pl. 103: d-k; Pl. 104
Thalassiosira mala – p. 199, Pl. 105
Thalassiosira minima – p. 202, Pl. 106: a-g
Thalassiosira minuscula – p. 202, Pl. 106: h-j
Thalassiosira nanolineata – p. 204, Pl. 96: j
Thalassiosira simonsenii – p. 204, Pl. 107
Thalassiosira sinica – p. 206, Pl. 108
Thalassiosira teleata – p. 208, Pl. 109
Thalassiosira tenera – p. 208, Pl. 110
Toxonidea insignis – p. 265, Pl. 136: g-j
Trachyneis antillarum – p. 243, Pl. 124: a-c, k
Trachyneis aspera – p. 243, Pl. 124: d-g, l
Trachyneis debyi – p. 244, Pl. 124: h-j, m
Triceratium dubium – p. 34, Pl. 10: h, i
Triceratium robertsianum – p. 34, Pl. 10: a-e
Trieres mobiliensis – p. 149, Pl. 73
Trieres sinensis (chinensis) – p. 151, Pl. 74
Tryblionella coarctata – p. 292, Pl. 149: a-c
Tryblionella compressa – p. 294, Pl. 149: d-f
Tryblioptychus cocconeiformis – p. 211, Pl. 111
Notes

Volume II
Diatoms
ISBN 978-99966-37-20-9
Kuwait Institute for Scientific Research
P.O. Box 24885, Safat - 13109, Kuwait
Tel: (965)24989000 Fax: (965)24989399
www.kisr.edu.kw

1b.marine Phytoplankton Vol2

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MARINE PHYTOPLANKTON

Faiza Y. AL-YAMANI and Maria A. SABUROVA

Faiza Y. AL-YAMANI and Maria A. SABUROVA

Kuwait Institute for Scientific Research

Faiza Y. AL-YAMANI and Maria A. SABUROVA

Published in Kuwait in 2019 by

Copyright © Kuwait Institute for Scientific Research, 2019

Printed and bound by

SYSTEMATIC ACCOUNT: Marine Planktonic Diatoms

References …………………………………………………………………………………………… 322

Coral reef surrounding Qaru Island

We acknowledge with thanks the assistance provided by Mr. Alan

The authors are grateful to the Kuwait Institute for Scientific

The presented phytoplankton taxonomic account of the diatom

As a whole, a pronounced numerical

Marina Crescent Beach, Salmiya

Empire EUKARYOTA Chatton

Proboscia alata (Brightwell) Sundström

Proboscia indica (Péragallo) Hernández-Becerril

Dactyliosolen phuketensis (Sundström) Hasle

Genus Guinardia Péragallo

Guinardia delicatula (Cleve) Hasle

Guinardia flaccida (Castracane) Péragallo

Genus Neocalyptrella Hernández-Becerril et Meave del Castillo

Neocalyptrella robusta (Norman ex Ralfs) Hernández-Becerril et Meave

Plate 5. Neocalyptrella robusta and Pseudosolenia calcar-avis.

Genus Pseudosolenia Sundström

Pseudosolenia calcar-avis (Schultze) Sundström

Rhizosolenia bergonii Péragallo

Rhizosolenia (Pseudosolenia?) cochlea Brun

Rhizosolenia hyalina Ostenfeld

Rhizosolenia imbricata Brightwell

Plate 7. Rhizosolenia hyalina.

Rhizosolenia setigera Brightwell

Plate 9. Rhizosolenia spp.

Triceratium dubium Brightwell

Triceratium robertsianum Greville

Plate 10. Triceratium spp. and Schuettia annulata var. minor.

Stephanopyxis palmeriana (Greville) Grunow

Plate 11. Stephanopyxis palmeriana.

Corethron histrix Hensen

Plate 12. Corethron histrix.

Asteromphalus cleveanus Grunow

Asteromphalus flabellatus (Brébisson) Greville

Plate 13. Asteromphalus cleveanus.

Asteromphalus heptactis (Brébisson) Ralfs

Asteromphalus sarcophagus Wallich

Schuettia annulata var. minor (Grunow) De Toni

Coscinodiscus asteromphalus Ehrenberg

Plate 16. Coscinodiscus asteromphalus.

Coscinodiscus granii Gough

Plate 17. Coscinodiscus gigas.

Coscinodiscus marginatus Ehrenberg

Plate 20. Coscinodiscus spp.

Coscinodiscus oculus-iridis Ehrenberg

Plate 21. Coscinodiscus nobilis.

Coscinodiscus radiatus Ehrenberg

Plate 22. Coscinodiscus oculus-iridis.

Genus Coscinodiscopsis Sar et Sunesen

Coscinodiscopsis jonesiana (Greville) Sar et Sunesen

Plate 23. Coscinodiscus radiatus.

Plate 25. Coscinodiscopsis jonesiana.

Palmerina hardmaniana Hasle

Family Heliopeltaceae Smith