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MARINE
ZOOPLANKTON
PRACTICAL GUIDE (Volume 2)
for the Northwestern Arabian Gulf
Faiza Yousif Al-Yamani

Valeriy Skryabin
Aleksandra Gubanova
Sergey Khvorov
Irina Prusova
MARINE ZOOPLANKTON
PRACTICAL GUIDE
Volume 2
Faiza Yousif Al-Yamani

Valeriy Skryabin
Aleksandra Gubanova
Sergey Khvorov
Irina Prusova
Kuwait Institute for Scientific Research, Kuwait
2011
MARINE ZOOPLANKTON PRACTICAL GUIDE
Published in Kuwait in 2011 by

Kuwait Institute for Scientific Research,
P.O. Box 24885, 13109 Safat, Kuwait
Copyright © Kuwait Institute for Scientific Research, 2011

All rights reserved
Design by
Mariposa Marketing & Advertising
Creative Director: Melad Helani
Printed and bound by

Waves Press
First Edition
Kuwait Institute for Scientific Research (Publisher)
ISBN 978-99966-95-07-0
No part of this work may be reproduced or utilized in any form or by any means
electronic or manual, including photocopying, recording or by any information
or retrieval system, without the prior written permission of the Kuwait Institute
for Scientific Research.
Salmiya beach - Photo by Dr. V. Skryabin - Kuwait Institute for Scientific Research
M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f

Table of CONTENTS
PREFACE i
ACKNOWLEDEGEMENTS iIi
2.13. Copepoda 1
General Characteristics, Morphology, Terminology and Abbreviations 2
List of Copepods Taxa 8
Systematic account of Copepod Species from the Arabian Gulf 10
Order CALANOIDA 10
Family CALANIDAE 10
Genus Canthocalanus 12
Canthocalanus pauper 12
Family PARACALANIDAE 14
Genus Acrocalanus 16
Acrocalanus gibber 16
Acrocalanus longicornis 18
Genus Bestiolina 20
Bestiolina arabica 20
Genus Paracalanus 22
Paracalanus indicus 22
Paracalanus sp. 24
Genus Parvocalanus 26
Parvocalanus crassirostris 26
Parvocalanus elegans 28
Family EUCALANIDAE 29
Genus Subeucalanus 29
Subeucalanus flemingeri 30
Subeucalanus subcrassus 32
Family CLAUSOCALANIDAE 33
Genus Clausocalanus 33
Clausocalanus minor 35
Family EUCHAETIDAE 36
Genus Euchaeta 36
Euchaeta concinna 38
Euchaeta rimana 40
K u wait I n s titute f or Scienti f ic R e s earc h

Family CENTROPAGIDAE 42
Genus Centropages 42
Centropages furcatus 44
Centropages orsinii 46
Centropages tenuiremis 46
Family PSEUDODIAPTOMIDAE 48
Genus Pseudodiaptomus 48
Pseudodiaptomus arabicus 49
Pseudodiaptomus ardjuna 50
Family TEMORIDAE 51
Genus Temora 51
Temora discaudata 52
Temora turbinata 54
Family CANDACIIDAE 56
Genus Candacia 56
Candacia bradyi 57
Family PONTELLIDAE 58
Genus Calanopia 58
Calanopia elliptica 58
Calanopia minor 60
Genus Labidocera 62
Labidocera acuta 62
Labidocera bengalensis 66
Labidocera kroyeri 68
Labidocera minuta 70
Labidocera sp. 72
Genus Pontella 74
Pontella danae 74
Pontella investigatoris 76
Genus Pontellopsis 78
Pontellopsis herdmani 78
Family ACARTIIDAE 80
Genus Acartia 80
Acartia (Acanthacartia) fossae 82
Acartia (Odontacartia) amboinensis 84
Acartia (Odontacartia) ohtsukai 86
Genus Acartiella 88
Acartiella faoensis 88

Family TORTANIDAE 90
Genus Tortanus 90
Tortanus barbatus 90
Tortanus forcipatus 92
Order CYCLOPOIDA 94
Family OITHONIDAE 94
Genus Oithona 94
Oithona attenuata 96
Oithona brevicornis 98
Oithona nana 98
Oithona plumifera 100
Oithona sp. 102
Family ONCAEIDAE 104
Genus Oncaea 104
Oncaea clevei 106
Family SAPPHIRINIDAE 106
Genus Copilia 108
Copilia mirabilis 108
Genus Sapphirina 110
Sapphirina nigromaculata 110
Family CORYCAEIDAE 112
Genus Corycaeus 112
Subgenus Dithrichocorycaeus 113
Corycaeus (Dithrichocorycaeus) andrewsi 113
Corycaeus (Dithrichocorycaeus) dahli 114
Corycaeus (Dithrichocorycaeus) lubbocki 116
Subgenus Onychocorycaeus 116
Corycaeus (Onychocorycaeus) agilis 117
Corycaeus (Onychocorycaeus) pacificus 118
Order HARPACTICOIDA 120
Family ECTINOSOMATIDAE 120
Genus Microsetella 120
Microsetella sp. 120
Family MIRACIIDAE 122
Genus Macrosetella 122
Macrosetella gracilis 122

Family EUTERPINIDAE 124
Genus Euterpina 124
Euterpina acutifrons 124
Family CLYTEMNESTRIDAE 124
Genus Clytemnestra 125
Clytemnestra scutellata 125
2.14. Mysida 126

Rhopalophthalmus sp. 126
2.15. Decapoda 128

Metapenaeus ensis, protozoea II 128
Parapenaeopsis stylifera, protozoea III 129
Penaeus semisulcatus, protozoea I 130
Penaeus semisulcatus, protozoea III 131
Solenocera crassicornis, protozoea III 132
Solenocera crassicornis, mysis 133
Solenocera hextii, protozoea III 134
Lucifer hanseni, protozoea III 135
Lucifer hanseni, mysis I 136
Lucifer hanseni, male 137
Lucifer typus, male 138
Acetes japonicas, female 140
Sergestes sp., protozoea II 142
Sergestes sp., mysis 143
Stenopus sp., zoea I 144
Thalassocaris obscura, zoea XI 145
Acanthephyra sp., zoea IV 146
Palaemon sp., zoea IV 147
Periclimenes sp., zoea IV 148
Alpheus sp. 149
Latreutes sp., zoea IX 150
Lysmata sp. 151
Callianassa sp., zoea I 152
Laomedia sp., zoea IV 153
Upogebia sp., zoea I 154
Thenus orientalis, phyllosoma I 155
Thenus orientalis, phyllosoma II 156
Thenus orientalis, phyllosoma III 157

Galathea sp., zoea III 158
Pachycheles sp., zoea I 159
Diogenes sp., zoea III 160
Dardanus sp., zoea I 161
Paguristes sp., zoea I 162
Pagurus sp., zoea III 163
Ebalia sp., zoea I 164
Leucosia sp., zoea I 165
Parthenope sp., zoea IV 166
Macrophthalmus sp., zoea III 167
Ilyoplax frater, zoea VI 168
2.16. Chaetognatha 169

Sagitta enflata 169
Sagitta neglecta 170
Sagitta pulchra 171
Sagitta regularis 172
2.17. Urochordata 173

Oikopleura dioica 173
Appendicularia sicula 174
Fritillaria sp. 175
Weelia cylindrica 176
Thalia sp. 177
Tadpole Larva 178
2.18. Cephalochordata 180

Branchiostoma sp. 180
Addendum 182
Catostylus mosaicus 182
Fritillaria pellucida 184
References 186
Index of Scientific Names 193

The marine zooplankton community of Kuwait
waters includes fascinating organisms, which are
abundant, diverse, and encompasses representatives
of all the major invertebrate phyla. Along with
PREFACE
phytoplankton, zooplankton are key components of
marine ecosystems forming the base of most marine
food webs. Plankton constitute the principal diet of
the early life stages of fish including commercially
important fish.
Zooplankton research is an essential component

in forming a more complete understanding of
the functioning of marine ecosystems. However,
identifying zooplankton can be a challenging and a
tedious task and requires a high level of expertise.
The samples used to produce this guide were

obtained during the 1999-2010 sea cruises covering
all Kuwait waters as well as some localities in
the Arabian Gulf waters. A total of 183 species of
zooplankton were recorded and identified. Species
diversity was highest for the groups of tintinnids,
copepods, and cnidarians (75, 54 and 14 species,
respectively).
This illustrated zooplankton guide is an

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

sample collection, sample processing and analysis
are included in Volume 1 of the zooplankton
guidebook (pages 2-8). Moreover, Volume 1 contains
descriptions and photographs of protozoan
zooplankton (Tintinnida), Cnidaria, Ctenophora,
Rotifera, Cladocera, Ostracoda and Larvae of marine
benthos.
i M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f

“We consider species to be like
a brick in the foundation of a
building. You can probably lose
one or two or a dozen bricks and
still have a standing house. But by
the time you’ve lost 20 per cent of
species, you’re going to destabilize
the entire structure. That’s the way
ecosystems work.”
Donald Falk
K u wait I n s titute f or Scienti f ic R e s earc h ii

Gratitude is extended to Mr. Alan Lennox (Kuwait
Institute for Scientific Research, Kuwait) for his
tremendous help in collecting the needed samples
for this study from Kuwait waters, and to ROPME for
Acknowledgements
providing Arabian Gulf samples. Our appreciation
to Mrs. Gracekutty Thomas Vargese for her hard
work in mesozooplankton sample analysis, and to
Mr. Vasiliy Prusov for his help in computerizing the
Copepod drawings. We are grateful to Dr. Inna
Drapun (IBSS, Sevastopol) for her great contribution
in identifying and description of ostracods and
photographing of zooplankton. We thank Mrs.
Linda Fernandes for compiling and formatting the
materials used for this guide.
The authors are grateful to the Kuwait Institute

for Scientific Research for the financial support of this
research project.
iii M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f

We share this planet with many species. It is our responsibility to
protect them, both for their sakes and our own.
Pamela A. Matson
K u wait I n s titute f or Scienti f ic R e s earc h iv

Bubiyan bridge off Khor Al-Sabbiya - Photo by Dr. V. Skryabin - Kuwait Institute for Scientific Research
12 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f

2.13. Copepoda
The copepods are crustaceans, which are abundant in Kuwait waters and often dominate the
plankton community, especially in Kuwait Bay. They are small in size (only a few species over 1
mm). They form a link in the food web between the primary-producing phytoplankton and the
plankton-feeding fish. Most of the economically important fish depend on copepods and other
zooplankters during their early life stages (larval fish stages) as well as adult stages of some fish
such as Zobaidy (Pampus argenteus), Suboor (Tenualosa illisha), and Beyah (Liza subviridis).
There are about 2,000 known marine pelagic species of copepods (Razouls and Bovée, 1998). The
copepod females lay eggs freely into the water, or produce external paired or single egg sacs. The
eggs hatch into copepod larvae. The first larvae are called nauplii. Larval copepods usually pass
five or six naupliar stages, which are separated by a moult. The 6th naupliar stage moults into the
first copepodite stage, which resembles the adult copepod. After molting through 5 copepodite
stages, copepods attain adulthood and cease molting.
Copepods comprise about 70% of the zooplankton community of Kuwait waters (Al-Yamani et al.,
1997 a, b), and have a variable distribution both spatially and temporally. The first comprehensive
zooplankton study in the northwestern Arabian Gulf with emphasis on copepod species was
conducted in 1979-1980 (Michel et al., 1986 a, b). The first guidebook on the copepods of Kuwait
waters was by Al-Yamani and Prusova, 2003.
This guidebook includes an updated version on copepods (54 species), with descriptions,
drawings and photographs of the adult stages of the common copepod species encountered
mainly in the waters of the NW Arabian Gulf. Relevant identification keys are also included in the
guide.
K u wait I n s titute f or Scienti f ic R e s earc h 1

General Characteristics, Morphology, Terminology and Abbreviations
Copepods are typically small metazoan organisms. In the marine planktonic forms, total body
length of copepods is usually between 0.5 and 5.0 mm, however, considering the full range of
species, total body length is from about 0.2 mm to about 10 mm.
Copepods belong to the subclass Copepoda according to the following hierarchy:
Phylum Arthropoda
Class Crustacea
Subclass Copepoda
Currently, there are 8 orders of marine planktonic copepods. Poecilostomatoida is no longer
considered as a group phylogenetically separated from Cyclopoida, and the order of Gelyelloida
is defined from species in subterranean karstic systems (Boxshall and Halsey, 2004). In this guide,
three orders of copepods were addressed as follows:
Order Calanoida
Order Cyclopoida
Order Harpacticoida
The classification of the copepods is largely based on body tagmosis (division into functional
regions) and segmentation and armature of the various limbs (Fig. 114). The major body
articulation subdivides the body into two major parts – the prosome (P) and the urosome
(Ur) (Fig. 115). The anterior region of the prosome, covered by the dorsal cephalic shield, is the
cephalosome (Ce), which comprises the five cephalic somites bearing the antennules (A1),
antennae (A2), mandibles (Md), maxillules (Mx1) and maxillae (Mx2), and the first thoracic
somite bearing the maxillipeds (Mxp). In many copepods the second thoracic somite is also
fused to the cephalosome, this somite is called the first pedigerous somite. The pedigerous
somites (Pd1-5) bear a pair of swimming legs (P1-5) each. The swimming legs usually have two
basal segments: coxa (C) and the basis (B). Two rami are ariculated to the basis: an outer exopod
(Exp) and inner endopod (Enp).
Within the Copepoda there are two major plans of body organization (tagmosis) into an anterior
prosome and a posterior urosome, separated by the major body articulation (asterisks in Fig.
116). In the gymnoplean tagmosis (in this document, Calanoida, Fig. 116a) the major body
articulation is located between the fifth pedigerous somite (Pd5), primitively bearing the P5, and
the anterior urosomal somite – genital somite (Gns). The abdomen is the limbless postgenital
region of the body. The last somite of the urosome is the anal somite bearing a pair of caudal
rami (CR).
In the podoplean type of tagmosis (in this document, Cyclopoida, Harpacticoida,

Poecilostomatoida) the major articulation is primitively positioned between Pd4 and Pd5 (Fig.
116b), i.e., in podopleans, the urosome includes the thoracic somite bearing the P5.

Fig. 114. The different parts of an adult calanoid
copepod. Abbreviations- A1: antennule; A2: antenna;
Abd 1: abdominal somite 1; Abd 2: abdominal somite
2; Ans: anal somite; CR: caudal ramus; Gns: genital
somite; Md: mandible; Mx1: maxillule, Mx2: maxilla;
Mxp: maxilliped (with coxa, basis and endopod); P1-
5: swimming legs 1-5; and Pd 1-5: pedigerous somites
1-5 (Source: Owre and Foyo, 1967).

A total of 50 species of copepods from Kuwait waters have been identified, with additional 4
species that were identified only to the genus level. These identified copepod species belong to 3
orders: Calanoida, Cyclopoida and Harpacticoida. In calanoids the major movable articulation
of the body occurs immediately posterior to the last free thoracic somite, i.e., between the
last prosome somite and the genital somite, so that the urosome consists only of genital and
abdominal segments (Fig. 117A). In cyclopoids and harpacticoids a major articulation in the
trunk occurs immediately anterior to the last free thoracic segment, so the last body region
(urosome) consists of one segment, bearing a usually rudimentary pair of swimming legs, the
genital somite, and the abdominal segment ending in caudal rami (Fig. 117B – 117D). In most
cyclopoids, as in calanoids, the urosome is much narrower than the prosome. In harpacticoids,
there is much less of difference in segment width between the prosome and urosome (Fig. 117D).
The terminology adopted here for the description of the external morphology of copepods
follows that of Huys and Boxshall (1991). The term “somite” is used in describing certain parts of
the copepod body (the prosome and urosome), the term “segment” is used in descriptions of the
parts of the antennules, mouth appendages and legs. The list below identifies each abbreviation
used in the descriptions, figures and keys (Figs. 114 and 115).
A1: antennule
A2: antenna
Ans: anal somite
B: basis
C: coxa
Ce: cephalosome
CR: caudal ramus
Enp: endopod; Enp1-3 – endopod segments 1-3
Exp: exopod; Exp1-3 - exopod segments 1-3
Gn: gnathobase
Gns: genital somite
Md: mandible
Mdp: mandibular palp
Mx1: maxillule
Le: external lobe
Li: internal lobe
Mx2: maxilla
Mxp: maxilliped
P1-5: swimming legs 1-5
Pd1-5: pedigerous somites 1-5
Pr: prosome
R: rostrum
Ur1-5: urosomites 1-5
Total length – the length from the top of a copepod head up to the end of CR, excluding caudal
setae.
There are typically six copepodite stages (abbreviated C1-C6), the sixth being either the adult
male and female. Growing from stage to stage is carried through moulting. Lebour (1916) was
first to describe the copepodite stages. Later the tables to determine these stages were made.
Morphological features of stages of different species of Copepoda are presented in the following
table.

Identification of Copepodite Stages 1-6 (Source: Sazhina, 1987)
Segments Number
Stage Number of Swimming Legs
Prosome Urosome
C1 4 2 2+ underdeveloped P3
C5 female 6 3-4 4 or 5
C5 male 6 4-5 4 or 5
C6 female 6 3-4 (Gns swollen) 4 or 5
C6 male 6 4-5 (Gns elongated) 5
Fig. 115. External morphology of copepods: a. the different parts of an adult copepod, b. A1
(antennule), c. A2 (antenna), d. Md (mandible), e. Mx1 (maxillule), f. Mx2 (maxilla), and g. Mxp
(maxilliped) (Source: Bradford-Grieve et al., 1999).

b c
d e
f g
Fig. 115. Cont’d.

Conway et al. (2003) suggested that most of the literature on identification only give descriptions
of the adult stage of copepods. In most female copepods, swollen genital somite indicates that it
is mature. But genital somite of Acartia start to swell in pre-adult copepodite 5 stage (C5), while
the body segmentation is often identical between C5 and C6. However, in both female and male
pre-adult copepods, the segmentation of the P5 and urosome is not distinct as in the adult stage,
which becomes obvious when the identifier examines them together (Conway, et al., 2003). I n
other genera, e.g., Pseudodiaptomus and Centropages, pre-adult females (C5) have a bit swollen
genital somite similar to Acartia.
a b
Fig. 116. Plan of body organization in a. gymnopleans

and b. podopleans (Source: Bradford-Grieve et al.,
1999).
Fig. 117. Differences between copepod Orders:

a. Calanoida (Canthocalanus pauper); b, c. Cyclopoida (b.
Oithona brevicornis, c. Oncaea clevei); d. Harpacticoida
(Euterpina acutifrons). Arrows indicate the genital
a b c d somites. (Source: Al-Yamani and Prusova, 2003).

List of Copepods Taxa
Phylum Class Susclass Order Family Genus / Species
Arthropoda Crustacea Copepoda Calanoida Calanidae Canthocalanus pauper

Paracalanidae Acrocalanus gibber
Acrocalanus longicornis
Bestiolina arabica
Paracalanus indicus
Paracalanus sp.
Parvocalanus crassirostris
Parvocalanus elegans
Eucalanidae Subeucalanus flemingeri

Subeucalanus subcrassus
Clausocalanidae Clausocalanus minor
Euchaetidae Euchaeta concinna

Euchaeta rimana
Centropagidae Centropages furcatus

Centropages orsinii
Centropages tenuiremis
Pseudodiaptomidae Pseudodiaptomus arabicus

Pseudodiaptomus ardjuna
Temoridae Temora discaudata

Temora turbinata
Candaciidae Candacia bradyi
Pontellidae Calanopia elliptica

Calanopia minor
Labidocera acuta
Labidocera bengalensis
Labidocera kroyeri
Labidocera minuta
Labidocera sp.
Pontella danae
Pontella investigatoris
Pontellopsis herdmani
Acartiidae Acartia (Acanthacartia) fossae

Acartia (Odontacartia) amboinensis
Acartia (Odontacartia) ohtsukai
Acartiella faoensis

Phylum Class Susclass Order Family Genus / Species
Tortanidae Tortanus barbatus

Tortanus forcipatus
Cyclopoida Oithonidae Oithona attenuata

Oithona brevicornis
Oithona nana
Oithona plumifera
Oithona sp.
Oncaeidae Oncaea clevei
Sapphirinidae Copilia mirabilis

Sapphirina nigromaculata
Corycaeidae Corycaeus (Dithrichocorycaeus) andrewsi

Corycaeus (Dithrichocorycaeus) dahli
Corycaeus (Dithrichocorycaeus) lubbocki
Corycaeus (Onychocorycaeus) agilis
Corycaeus (Onychocorycaeus) pacificus
Harpacticoida Ectinosomatidae Microsetella sp.
Miraciidae Macrosetella gracilis
Euterpinidae Euterpina acutifrons
Clytemnestridae Clytemnestra scutellata

Systematic Account of Copepod Species from the
Northwestern Arabian Gulf
Phylum Arthropoda
Class Crustacea Brunnich, 1772
Subclass Copepoda Milne-Edwards, 1840
Order Calanoida Sars, 1903
The Calanoida are defined by the combination of the gymnoplean tagmosis, the presence of
only 1 spine on the outer margin of P2-P5 Exp1, the presence of a coxal epipodite (Le1), but
not a lobate basal exite on Mx1, and the presence of a seta on the inner margin of A2 coxa. The
presence of a maximum of 2 setae on the terminal segment of Mx2 (Enp4) is an apomorphy
(derived character) of the Calanoida (Huys and Boxshall, 1991; Bradford-Grieve et al., 1999).
Family Calanidae Dana, 1849

Reference: Bradford-Grieve et al., 1999.
Females
Cephalosome and Pd1 may be fused or separate, Pd4 and 5 always separate. Rostrum of 2
filaments. Caudal rami with 4 subequal, terminal setae and 1 outer seta. Urosome of 4 somites.
A1 25-segmented, generally segments 8-9 partly fused; segments 23-24 with elongate seta. A2
with 1 seta, B with 2 setae; A2 Exp 7-segmented; Exp1 and Exp2 with 2 setae each, Exp3-6 with 1
seta each, Exp7 usually with 1 seta nearly at midlength and 3 terminal setae; Enp1 with 2 setae,
Enp2 with 14-16 setae. Mdp B with 4 setae; Enp1 with 4 setae and prominent lobe; Enp2 totally
with 10-11 (8 or 9 terminal and 2 posterior) setae. Exopod and endopod of approximately equal
length. Mx1 Li1 with 9 terminal spines, 4 posterior and 1 anterior setae; Li2 and Li3 with 4 setae
each; Li4 with 4 setae, endopod with 14-15 setae; exopod with 11 setae; Le1 with 9 setae; Le 2
with 1 seta. Mx2 Li1-Li5 with 4-6, 3, 3, 3 and 4 setae, respectively. Terminal part with about 10
setae. Mxp coxa with 1 proximal seta, then with groups of 2, 4 and 4 setae from proximal to distal;
Enp2-6 with 4, 4, 3/4, 4, 4/5 setae respectively. P2-P3 B with an external articulated spine, on P4
and P5 this may be reduced to a small seta. P2 Exp1 sometimes with recurved spine (Neocalanus),
or Exp2 with deep invagination (Undinula). P5 similar to P2-4; sometimes with inner edge teeth
(Calanus, Nannocalanus, Cosmocalanus).
Males
Urosome of 5 somites. A1 25-segmented, with segments 1 and 2 always fused, segments 3-5,
7 and 8, 9 and 10, and 24 and 25 may also be fused. A2-Mx2 – either exactly as in female, or
reduced in size and setation. Mxp with reduced inner setae but enlarged outer setae on terminal
segments. P1-P4 as in female. P5 with both rami usually 3-segmented, right leg similar to other
legs, left leg variously modified; endopod sometimes reduced and devoid of segmentation on
one or both sides.

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

Genus Canthocalanus A. Scott, 1909
References: Bradford and Jillet, 1974; Bradford-Grieve et al., 1999.
Females
Cephalosome and Pd1 fused. Pd4 and Pd5 separate. Rostrum of 2 filaments. Caudal rami with
4 subequal, terminal setae and 1 outer seta. Urosome of 4 somites. A1 25-segmented. A2 Exp
7-segmented. Mdp B with 4 setae, Exp and Enp of approximately equal length. Mx2 with 4 setae
on Li1. P1: B1 anterior margin terminates in a well defined projection; B2 with distal seta on
anterior surface modified into a proximally thickened spine. P2-P3 B with an external articulated
spine. P2-P4 without modification or ornamentation. P5 with the inner border of B1 naked, Enp
with 7 setae.
Males
Urosome of 5 somites. A1 25-segmented. Cephalosome and Pd1 fused. Mouthparts as in female.
P5 with the inner border of B1 naked; both rami 3-segmented, hardly modified on right; left
endopod with only 2 terminal setae.
Canthocalanus pauper Giesbtecht, 1888
Female. Length: 1.4-1.7 mm (1.3-1.6 mm). (Fig. 118 A, B, D; Fig. 119 a, b)

Head and pedigerous segment 1 fused. Anterior cephalosome and posterior metasome rounded.
Rostrum of 2 filaments. Urosome 4-segmented. Strong setae on caudal rami. Mx2 Li1 with 4
setae. P1 C anterior margin terminates in well-defined projection; basis with inner marginal seta
modified into proximally thickened spine. P5 C inner edge without teeth (naked). P1-P5 Enp with
7 setae.
Male. Length 1.2-1.5 mm (1.3-1.5 mm). (Fig. 118 C, E; Fig. 120 a, b)
Right P5 asymmetric; endopod with 8 setae; exopod without inner marginal spines; left endopod
with 2 terminal setae; left exopod with elongated segments, with long outer distal setae on
segments 2 and 3. Inner margins of the P5 coxa are not denticulate.

a b
Fig. 119. Canthocalanus pauper. Female: a. dorsal view;

b. right lateral view.
Fig. 118. Canthocalanus pauper.

Female: A. dorsal view, B. right lateral view, D. P5; Male:
C. dorsal view, E - P5.
a b
Fig. 120. Canthocalanus pauper. Male: a. dorsal view; b. right lateral view.

Family Paracalanidae Giesbrecht, 1892
References: Giesbrecht, 1892; Vervoort, 1963; Andronov, 1971, 1977; Bradford-Grieve, 1994 (cited
by Bradford-Grieve et. al., 1999).
Females (Fig. 121)

Cephalon and Pd1 usually fused. Pd4 and 5 fused, or separated. Urosome of 2 to 4 somites.
Anal somite usually longer than any somite between it and genital somite. Rostrum of 2
filaments (Acrocalanus, Calocalanus, Paracalanus); 2-pointed, solid (Delius, Bestiola); or massive
(Parvocalanus). A1 usually 25-segmented, generally with segments 1 and 2, also 8 and 9, partially
fused; in Delius first 8 segments fused. A2 Exp 7-segmented with Exp1 and Exp2 each bearing 2
setae, segment 7 elongate. Md Enp1 without prominent lobe. Mx1 Li1 with 14 setal elements (4
posterior and 1 anterior setae, and 9 terminal spines); Li2 and Li3 with 3-4 setae each; Li4 with
4 setae; endopod with 13-14 setae; exopod with 11 setae; Le2 with 1 reduced seta; Le1 with
7-9 setae. Mx2 Li1-Li5 with 6, 3, 3, 3 and 3-4 setae respectively; terminal part with 8 setae, 1 of
them may belong to developed Li6. Mxp C with 1 proximal seta, then 3 groups of 2, 3 and 4
setae from proximal to distal; Enp2-Enp6 with 3, 4, 3, 3+1 and 4 setae respectively. P1-P4 exopod
usually 3-segmented but P1 exopod 2-segmented in some Calocalanus species. P2-P4 endopods
usually 3-segmented but 2-segmented in Calocalanus minor, posterior surfaces of some
segments ornamented with spines. P2-P4 Exp3 external borders serrated, or smooth in Delius
and Calocalanus. Terminal spines of P2-P4 exopods smooth. P5 uniramous, absent, or vestigial
(Acrocalanus), present on left only (Delius). If both P5 present, then symmetrical, presented by
coxae (Bestiolina), 2-segmented (or 3-segmented) (Paracalanus, Parvocalanus), or 3-4 segmented
(Calocalanus). Females of four genera of family Paracalanidae common for the Kuwait waters,
Acrocalanus, Bestiolina, Paracalanus and Parvocalanus, are shown in Fig. 121.
Differences between Genera Acrocalanus, Bestiolina, Paracalanus and Parvocalanus are displayed
in the table below.
Males (Fig. 122)

Urosome of 5 somites. Genital opening on left (in Calocalanus on left or right). Cephalic hump
present (Acrocalanus, Bestiolina and Paracalanus), or absent. A1 25-segmented, with some or all
segments 1-6 fused and swollen, segments 9-10 may also be fused. A2 Exp7 very short without
3 terminal setae, which are present in female. Exp1 and Exp2 without setae. Other oral parts
reduced compared with female. Right P5 may be absent (Acrocalanus, Delius), or present as C
(Bestiolina), 2-3-segmented (Paracalanus, Parvocalanus), or 3-4-segmented (Calocalanus).
There are four common genera of Paracalanidae in Kuwait water: Acrocalanus, Giesbrecht, 1888;
Bestiolina, Andronov, 1991; Paracalanus, Boeck, 1864; Parvocalanus, Andronov, 1970.
Construction of swimming legs is also an important feature, which allows to distinguish one
Paracalanidae genus from another (Fig. 122). Acrocalanus P2-P4 Exp2 and Exp3 external edges
serrated; Bestiolina P2-P4 Exp2 and Exp3 external borders smooth. Paracalanus P2-P4 Exp3
external borders serrated; Parvocalanus P2-P4 Exp3 external borders serrated.
Differences between Genera Acrocalanus, Bestiolina, Paracalanus and Parvocalanus
Genus Size Rostrum P5
Acrocalanus 0,8 – 1,25 of 2 filaments absent

Bestiolina 0,6 – 0,75 of 2-pointed, solid, long extensions presented by coxa
Paracalanus 0.7 –1,1 of 2 filaments symmetrical, uniramous
2 segmented
Parvocalanus 0,45 – 0,65 of 2-pointed, solid, short extensions symmetrical, uniramous
2 segmented

1 2 3 4
Fig. 121. Females of four genera of Paracalanidae family common for the Kuwait waters: 1. Acrocalanus;
2. Bestiolina; 3. Paracalanus; 4. Parvocalanus. Arrows indicate the rostrum and leg 5.
(Sources: 1, 4 – from Al-Yamani and Prusova, 2003; 2 – from Ali et al. 2007, 3 – from Bradford-Grieve et al.,1999).
1 2 3 4
Fig. 122. Fourth swimming leg (P4) exopods of Paracalanidae females: 1. Acrocalanus; 2. Bestiolina;
3. Paracalanus; 4. Parvocalanus.
(Sources: 1, 3, 4 – from Al-Yamani and Prusova, 2003; 2 – from Ali et al., 2007).

Genus Acrocalanus Giesbrecht, 1888
Reference: Bradford-Grieve et al. (1999).
Females
Cephalosome and Pd1 fused. Pd4 and Pd5 partly fused. Urosome of 4 somites, anal somite longer
than any somite between it and genital somite. Rostrum of 2 filaments. A2 Exp 7-segmented
with Exp1 and Exp2 each baring 2 setae, segment 7 elongate. Md Enp1 without prominent lobe.
P1-P4 exopods 3-segmented; P1 B with inner marginal seta; P2-P4 endopods 3-segmented, Enp2
posterior surface ornamented with spines; P2-P4 Exp2-3 external edges serrated. Terminal spines
of P2-P4 Exp smooth. P5 absent or vestigial.
Males
Urosome of 5 somites. Genital opening on left. Cephalic hump present (lateral view). A1 several
first segments fused and swollen. Right P5 absent.
Acrocalanus gibber Giesbrecht, 1888
Female. Length: 0.9-1.2 mm (0.93-1.28 mm). (Fig. 123 A, B, C D, F; Fig. 124 a, b)

Deep body in lateral view, with a pronounced hump on the dorsal cephalosome; distinct
though partial line of separation of the cephalosome from the fist pedigerous segment is
obvious; dorsal outline of cephalosome significantly humped in lateral view. A1 exceeds beyond
end of caudal rami by 2 terminal segments. 4 pair of swimming legs. P4 E xp3 external edge
teeth on distal end equal in size to those on proximal end. Genital somite longer than any of
others, with a prominent ventral swelling.
Male. Length: 0.9-1.2 mm (0.94-1.24). (Fig. 123 E; Fig. 124 c)

P4 Exp3 with about 10-12 teeth o n distal part of segment. P5 is only present on the left and
extends to the end of the urosome segment 3, or the middle of urosome segment 4 when this
leg is fully extended; very simple limb, composed of four segments with two tiny spines on
the last segment.

Fig. 123. Acrocalanus gibber. Female.
A. dorsal view; B. left lateral view; C. Ur, dorsal; D. Ur, left lateral view; E. P4, male; F. P4, female.
(Sources: A-D - from Al-Yamani, Prusova, 2003; E, F - from Sewell (1929) cited by Conway, 2003).
a b c
Fig. 124. Acrocalanus gibber: a. female, dorsal view; b. female, lateral view; c. male, lateral view.

Acrocalanus longicornis Giesbrecht, 1888
Reference: Bradford-Grieve, 1994.
Female. Length 1.00 – 1.26 mm. (Fig. 125 1 – 7; Fig 126 a – d)

Dorsal surface of the body moderately strongly arched, anterior head rounded. A1 extends
beyond the caudal rami by its last 5 segments. Outer distal border of exopod segment 3 of P4
with very small numerous teeth.
Male. Length 0.95 – 1.25 mm. (Fig. 127 a, b)

Exp 3 of P4 similar to that of the female. P5 - 5 segmented on the left.
Fig. 125. Acrocalanus longicornis. Female.

1. dorsal view; 2. lateral view; 3. P1; 4. P2; 5. P3; 6. P4; 7. P5.
(Source: Corral Estrada, 1970).

c d
a b
Fig. 126. Acrocalanus longicornis. Female. a. dorsal view; b. lateral view; c. exopod of P4; d. distal part of exopod
of P4.
a b
Fig. 127. Acrocalanus longicornis. Male. a. lateral view; b. P5.

(Source: Chen et al., 1974).

Genus Bestiolina, Giesbrecht, 1888
References: Ali et al., 2007
Bestiolina arabica Ali et al., 2007
Female. Length 0.79-0.92 mm. (Fig. 128 a – g; Fig. 130 a, d)

Cephalosome is fused to the first pedigerous segment and slightly gibbous dorsally. Rostrum 2-
pointed and solid. The fourth and fifth pedigerous segments are separated. Urosome consists of
four segments having genital somite as the widest region in the middle part of the dorsal view.
Anal segment longer than any other segments except the genital somite. The antennule is long
in a way that almost reaches the end of the anal segment. Marginal spinules are absent from
exopods and endopods of legs 2-4. All legs are not serrated. The fifth leg is reduced (basipodites).
Male. Length: 0.7-0.90 mm. (Fig. 129 a – k; Fig. 130 b)

A cephalic hump observed from the lateral view of the prosome. Rostrum 2-pointed, solid, more
slender than in female. The shape of the prosome is narrower if compared to female. Urosome is
composed of five segments. Genital opening on the left. Antennule is long in a way that almost
reaches the middle of urosome. The description of legs 1-4 almost same as those of females but
the main difference lies in the number of dorsal spines of the second endopods of legs 2-3, which
are four dorsal spines. The second main difference is the asymmetric shape of the fifth leg,
which has a rudimentary right part with a basipodite only. The left part is composed of five
segments, and the last one carries two terminal spines; one is obviously longer than the other.
Remarks: Some individuals of B. arabica have features both of female and male (Fig. 130 c, e).
This androgynes have swollen genital somite same as typical female. However, Leg 5 of these
individuals has male construction, although underdeveloped.
Fig. 128. Bestiolina arabica. Female.

a. dorsal view; b. lateral view; c. R; d-e. Ur (dorsal and lateral, respectively); f. A1; g, A2. Scale - 0.01 mm (a, b,. d-g);
0.005 mm (c). (Source: Ali et al., 2007).

c
Fig. 129. Bestiolina arabica. Male.

a. dorsal view; b. lateral view; c. R; d. Ur (dorsal);
e. A2; f. Mxp; g-j. P1 to P4; k. P5. Scale bars: 0.01
mm (a, b, d-k); 0.05 mm (c). (Source: Ali et al., 2007).
a b c d e
Fig. 130. Bestiolina arabica. a. female, lateral view; b. male, lateral view; c. androgyne, lateral view;
d. female Gns and P5, e. androgyne, Gns and P5. Arrows indicate leg 5.

Genus Paracalanus Boeck, 1864
Females
Cephalosome and Pd1 fused. Pd4 and Pd5 partly fused. Urosome of 4 somites, anal somite longer
than any somite between it and genital somite. Rostrum of 2 filaments. A2 exopod 7-segmented
with Exp1 and Exp2 each baring 2 setae, segment 7 elongate. Md Enp1 without prominent
lobe. P1-P4 Exp 3-segmented; P2-P4 Exp3 external borders serrated; P2-P4 Enp 3-segmented,
posterior surfaces of some segments ornamented with spines; P1 B with inner marginal seta.
Both P5 developed, symmetrical 2-3-segmented.
Males
Urosome of 5 somites. Genital opening on left. Cephalic hump present (lateral view). A1 several
first segments fused and swollen. Right P5 2-3-segmented
Paracalanus indicus Wolfenden, 1905
Female. Length 0.85-0.95 mm. (Fig. 131 A – L; Fig. 133 a)

Prosome about 3.2 times as long as urosome; dorsal hump only slightly or not at all developed.
Genital somite viewed dorsally, narrower than in Paracalanus sp., lateral parts of posterior margin
with row of minute spinules, without cluster of spinules above spermatheca.
Male. Length 0.85-1.02 mm. (Fig. 132 A – L; Fig. 133 b)

P2 – P4 basis without posterior surface spinules.
Fig. 131. Paracalanus indicus. Female.

A. dorsal view; B. right lateral view; C. A1; D. A2;
E. Mdp; F. mandibular blade; G. Mx1; H. Mx2; I.
Mxp. (Source: Bradford-Grieve, 1994).

Fig. 132. Paracalanus indicus. Male.
A. dorsal view; B. left lateral view; C. A1; D. A2; E. Mdp; F. Mx1; G. Mxp; H. P1; I – P2; J. P3; K. P4; L. P5.
(Source: Bradford-Grieve, 1994).
a b
Fig. 133. Paracalanus indicus: a. female, lateral view; b. male, lateral view. Arrow indicates genital somite.

Paracalanus sp.
Female. Length: 0.75-1.05 mm. (Fig. 134 A – L; Fig. 136 a, b)

Body stout with wide, smoothly rounded head. Prosome 2.5 times longer than wide. Rostral
filaments relatively long. A1 exceeds beyond end of caudal rami by 2 terminal segments. P1-P4
coxa naked , without hairs and setules both on anterior and posterior surface. P2 – P4 Enp2 with
long spines on posterior surface; P4 endopod 3 with 1 pair of spines at proximal and 1 pair at
distal end. P5 terminal segment and terminal inner spine of about equal lengths. Genital somite
large.
Male. Length: 0.8-1.2 mm.

Fig. 135 M – R; Fig. 137 a, b
A1 not reaching caudal rami. P2-P4 exopods 2-3 with teeth on outer edge. P2-P4 endopods 2-3
with many long spines and spinules. P5 right leg 3-segmented, left 5-segmented.
Remarks: Paracalanus sp. female resembles P. aculeatus Giesbrecht, 1888 in size and body
proportions, however, it differs from the latter in having more rounded Ce in lateral view, absence
of finger-like processes on P4 Exp2 and shorter distal spines of P5.
Paracalanus sp. differs from another large species of Paracalanus – P.campaneri Bjornberg, 1982,
by absence of tufts of long hairs on Prosome posterolateral corners.
Fig 134. Paracalanus sp. Female.

A. dorsal view; B. left lateral view; C. Ur, dorsal; D. Ur, left lateral view; E. Gns, ventral; F – R; G. P5; H – Mxp; I. P1;
J. P2; K. P3; L. P4.

Fig. 135. Paracalanus sp. Male.
M. right lateral view; N. P1; O. P2, with abnormal Enp; P. P3; Q. P4; R. P5.
a b a b
Fig. 136. Paracalanus sp. Female. Fig. 137. Paracalanus sp. Male.
a. dorsal view; b. left lateral view. a. dorsal view; b. right lateral view.

Genus Parvocalanus Andronov, 1970
Females
Cephalosome and Pd1 fused. Pd4 and Pd5 partly fused. Urosome of 4 somites, anal somite
longer than any somite between it and genital somite. Rostrum bifurcate, short, solid. P1 Enp
1- or 2-segmented, P1 B1 usually without inner marginal seta. P1-P4 Exp 3-segmented; P2-P4
Exp3 external borders serrated; P2-P4 Enp 3-segmented. Both P5 developed, small, symmetrical,
uniramous, 2-segmented.
Males
Urosome of 5 somites. Genital opening on left. Cephalic hump absent (lateral view). Right P5
2- or 3-segmented.
Parvocalanus crassirostris (F. Dahl, 1894)
Female. Length: 0.48-0.65mm (0.47-0.55 mm). (Fig. 138 A – J; Fig. 140 a – d)

Body stout with head narrowing anteriorly both in dorsal and lateral view. Prosome about 2 times
as long as wide and 3 times as urosome. Dorsal outline of cephalosome slightly humped in lateral
view. Genital somite wider than long. First swimming leg (P1) basis without inner margin seta;
endopod 2-segmented. Legs 2-4 (P2-P4) exopod 3 proximal outer borders serrated, distal outer
margin naked. Leg 5 (P5) 2-segmented, distal segment with 2 apical spines of wish the inner one
is about twice as long as outer marginal.
Male. Length: 0.4-0.4 mm (0.35-0.39). (Fig. 139 A – C; Fig. 141 a – b)

Head narrowly rounded both in dorsal and lateral view. Prosome 1.7 times as long as wide,
3 times as urosome. P5 right leg 2-segmented. Left leg longer than urosome, right leg less than
left leg basal segment length.
Fig. 138. Parvocalanus crassirostris. Female.

A. dorsal view; B. right lateral view; C. urosome,
dorsal; D. rostrum (lateral); E. rostrum (ventral);
F. P1; G. P2; H. P3; I. P4; J. P5.

Fig. 139. Parvocalanus crassirostris. Male.
A. dorsal view; B. right lateral view; C. P5.
Fig. 140. Parvocalanus crassirostris. Female.

a. dorsal view; b. right lateral view; c. rostrum (ventral
view); d. rostrum (right lateral view).
a b
Fig. 141. Parvocalanus crassirostris. Male.

a. dorsal view; b. right lateral view.
a b

Parvocalanus elegans Andronov, 1972
Female. Length: 0.45-0.58 mm (0.46-0.51). (Fig. 142 1, 2, 3-7, 8, 10, 11; Fig. 143)
Body quite slender. Prosome 2.75 times as long as wide. Cephalosome and first pedigerous somite fused.
Urosome 4-segmented, genital somite longer than any of others. P1 endopod 1-segmented. P4 Enp2
with a row of leaf-like spines.
Male. Length: 0.4-0.4 mm. (Fig. 142 9, 12-15, 16, 17)

Prosome is oval, head smoothly rounded. Р5 asymmetrical: left leg 5-segmented, right 3-segmented.
Fig. 142. Parvocalanus elegans.

Female: 1. ventral view; 2. lateral view;
3-7. P1 to P5; 8. R; 10-11. Gns (dorsal
and left lateral side, respectively). P.
elegans, male: 9 – R; 12-15 – P2 to P5;
16 – ventral view; 17 – lateral view.
(Source: Andronov, 1972).
Fig. 143. Parvocalanus elegans. Female, left lateral view.

Family Eucalanidae Giesbrecht, 1892
References: Bragdord-Grieve, 1994; Brodsky et al., 1983; Fleminger, 1973 (cited by Bradford-
Grieve et al., 1999).
Females
Body elongate, especially anterior to A2, more or less transparent, often with triangular anterior
part of cephalosome. Cephalosome and pedigerous somite 1 fused, pedigerous somites 4 and
5 partly fused. Caudal ramus sometimes asymmetrical; usually left ramus slightly and second
seta much longer than others. Rostrum of 2 long, slender filaments. Urosome of 3 or 4 somites
with caudal ramus usually completely fused to anal somite. A1 23-segmented, generally with
segments 1-2 and 8-9 fused. A2 Exp shorter than Enp; Exp1 and Exp2 with 2 setae each, and
either fused or separate. Md Enp reduced and B and Exp elongate (except in Rhincalanus). Mx1
Li1 with 14 terminal spines and setae including 4 posterior and 1 anterior setae; Li2 may be
absent or without setae, basis and Enp elongate; Exp with reduced number of setae; Le2 with 1
seta; Le1 with 9 setae. Mx2 Li1-Li5 with 6, 3, 3, 3 and 3-4 setae respectively. Mxp Enp2-Enp6 with
3, 4, 3, 4 and 4 setae respectively. Terminal Exp spines of swimming legs smooth. P5 absent or
present (Rhincalanus), 3-segmented, symmetrical.
Males
Urosome of 5 somites, with caudal ramus usually completely fused to anal somite. A1
24-segmented. Oral parts and legs similar to females. P5 uniramous on both sides (Eucalanus,
Pareucalanus), biramous on left (Rhincalanus), or absent on right (Subeucalanus).
Genus Subeucalanus Geletin, 1976

References: Geletin, 1976, Bradford-Grieve, 1994.
Females
Body elongate, especially anterior to A2; cephalosome and Pd1 fused, Pd4 and 5 partly fused.
Caudal rami sometimes asymmetrical; left ramus slightly and second seta much longer than
others. R of 2 long, slender filaments. Urosome of 3 somites with caudal rami fused to anal somite.
A1 23-segmented, with segments 1-2 and 8-9 fused. A2 Exp shorter than Enp, Enp1 elongate;
Exp1 and Exp2 fused. Mdp B2 and Exp elongate; Enp small, inserted on the terminal one third or
quarter of B2, which also bears 2 or 3 setae. Mx1 with 2 inner lobes. P1 Enp 2-segmented and Exp
3-segmented. P2-P4 Exp and Enp 3-segmented. P5 absent.
Males
Urosome of 5 somites, with caudal rami fused to anal somite. A1 24-segmented. Oral parts and
legs similar to females. P5 uniramous, right leg absent.
Key to Subeucalanus species

Females
Anterior medial part of cephalosome in dorsal view eminent and narrowing anteriorly, with small
depressions at both sides of anterior outline; prosome nearly 3.4 times as long as wide. Total
length 2.12-2.54 mm. …………………………………………………………………S. subcrassus
Anterior part of cephalosome smoothly rounded both in dorsal and in lateral views, without
depressions at anterior ouline in dorsal view; prosome nearly 3.1 times as long as wide. Total
length 2.01-2.14 mm. …………………………………………………………………S. flemingeri

Subeucalanus flemingeri Prusova, et al., 2001.
Female. Length 2.01-2.14 mm. (Fig. 144 A – G; Fig.146 a – b)

Prosome nearly 3.1 times as long as wide and 5.7 times urosome. Forehead rounded. Rostrum
robust, elongated, and ending in fine filaments slightly swollen at their bases. Genital somite
broader than long with widest part in dorsal view on the posterior half of the somite. A1 of 23
articulated segments, extending beyond tips of caudal rami by last 5-6 segments. Mdp B2 with
3 setae on inner margin, endopod reaches the distal border of B2, Enp1 with 2 setae Enp2 with 4
setae. Medial part of outer margin of P2-P4 Enp2 with acute projection.
Male. Length 1.97-2.14 mm. (Fig. 145 H – M; Fig.147 a – b)

Prosome nearly 3.1 times as long as wide and 4.1 times as urosome. Forehead rounded. Genital
somite slightly asymmetrical, with genital aperture located on left side; second somite the largest
and cask-shaped. A1 of 23 articulated segments, extending beyond end of urosome by last 5-6
segments.Mdp more robust than in female, with 3 seta on inner margin of B2. P1-P4 resemble
those of female, but the acute projection on P2-P4 Enp2 more developed than in female. P5
uniramous, 4-segmented, ending with distal spine nearly as long as terminal segment; the latter
with fine hairs on outer edge.
Fig. 144. Subeucalanus flemingeri. Female.

A. dorsal view; B. right lateral view; C. head and R, ventral; D. head and rostrum, lateral; E. Ur, dorsal; F. Ur, right
lateral view; G. Mdp.

Fig. 145. Subeucalanus flemingeri. Male.
H. dorsal view; I. right lateral view; J. Ur and P5, right lateral view; K. Ur, dorsal; L. P5; M. Mdp.
a b
a b
Fig. 146. Subeucalanus flemingeri. Female. Fig. 147. Subeucalanus flemingeri. Male.
a. dorsal view; b. left lateral view. a. dorsal view; b. right lateral view.

Subeucalanus subcrassus (Giesbrecht, 1888)
Female. Length 2.12-2.54 mm. (Fig. 148 A – F; Fig. 149 a, b)

Anterior medial part of cephalosome in dorsal view eminent and narrowing anteriorly, with small
depressions at both sides of anterior outline. Prosome nearly 3.4 times as long as wide and 5.7
times urosome. Genital somite broader than long with widest part in dorsal view on the posterior
half of the somite. Mdp B2 with3 setae, Enp reaches the distal border of B2, Enp1 with 2 setae
Enp2 with 4 setae.
Fig. 148. Subeucalanus subcrassus. Female.

A. dorsal view; B. right lateral view; C. anterior head and rostrum, right lateral view; D. Ur, dorsal; E. Ur, right lateral
view; F – Mdp.
a b
Fig. 149. Subeucalanus subcrassus. Female. a. dorsal view; b. left lateral view.

Family Clausocalanidae Giesbrecht, 1892
References: Frost and Fleminger, 1968; Bradford-Grieve, 1994; Brodsky et al., 1983 (cited by
Bradford-Grieve, 1994).
Females
Small copepods slightly over 1-1.5 mm. Pd1 and cephalosome usually fused, sometimes
separate, Pd4 and 5 usually fused. Rostrum usually with 2 filaments, two-pointed, or absent.
Posterior corners of prosome usually symmetrical, rounded. Genital somite usually symmetrical.
A1 generally with segments 8-9 fused; extremely long setae absent. A2 Exp 6- or 7-segmented.
Mdp B with 4 setae. P1 Enp with external lobe, or may be absent. Posterodistal border of B of P2
and P3 may be ornamented with spines, Enp of P2-P4 usually narrow. P5 absent, or present.
Males
P5 present on both sides, but of unequal length, uniramous; longer leg nearly always on left,
5-segmented, shorter leg 1-3-segmented, less than half length of coxa of other leg (Calusocalanus,
Ctenocalanus), or longer (Microcalanus); or both legs of similar length (Preudocalanus).
Genus Clausocalanus Giesbrecht, 1888

References: Frost and Fleminger, 1968; Bradford-Grieve, 1994.
Females
Small copepods, with total length less than 2.0 mm. Cephalosome and Pd1, Pd4 and 5 fused.
Urosome of 4 somites. Rostrum of 2 short, rigid spiniform processes. A1 23-segmented with
segments 8-9 and 24-25 fused. Posterior corners of prosome symmetrical, rounded. Genital
somite symmetrical. Mdp B with 4 setae. P1-4 Exp 3-segmented; Enp1 1-segmented, Enp2
2-segmented, Enp3 and Enp4 3-segmented. P2-3 B2 broadening distally to about 1.5 or more
times their width in region of attachment to B1; distal posterior margin with 3 or more spiniform
processes. Right and left P5 present, uniramous, 3-segmented, symmetrical; P5 third segment
produced distally into a short, pointed, bifid process.
Males
Urosome of 5 somites, anal somite very short. Rostrum reduced to a single, median, ventrally
protruding knob or not well developed. Right and left P5 present, uniramous, rami of very
unequal length, longer ramus somewhat styliform, 5-segmented, fifth segment short and
attached subapically to fourth segment; first segment more than twice as long as shorter ramus
of opposing leg.

Clausocalanus minor Sewell, 1929
Female. Length: 1.1 – 1.3 mm (1.08-1.26). (Fig. 150 A – F; Fig. 152 a – d)

Prosome posterior margins somewhat angular in lateral view. Ventral profile of Genital somite
in lateral view with a prominent step posterior to the genital pores. Genital somite about
twice as long as second urosome segment. Dorsal lobe of seminal receptacle in lateral view
conspicuous. P5 terminal points divergent and spinulate in inner margins.
Male. Length: 0.8-1.0 mm (0.79-1.04). (Fig. 151 G – K; Fig. 153 a, b)

P5 longer ramus and genital pore on left side. Left P5 longer than urosome; armed distally with
slender, straight setae. Right P5 2-segmented (according to Frost and Fleminger, 1968, right P5 in
C. minor may be 2 or 3-segmented).
Fig. 150. Clausocalanus minor. Female.

A. dorsal view; B. left lateral view; C. Ur, dorsal;
D. R; E. Gns, left lateral view; F. P5.
Fig. 151. Clausocalanus minor. Male.

G. dorsal view; H. left lateral view; I. Ur (dorsal); J. B2 of
P3; K. P5.

a b
c d
Fig. 152. Clausocalanus minor. Female. a. dorsal view; b. left lateral view; c. Ur; d. P3.
a b
Fig. 153. Clausocalanus minor. Male. a. dorsal view; b. left lateral view.

Family Euchaetidae Giesbrecht, 1892
References: Bradford, 1974; Bradford et al., 1983; Park, 1995 (cited by Bradford-Grieve et al., 1999).
Females
Pd1 separated from cephalosome by faint line representing partially fused joint, Pd4 and 5
completely fused. Rostrum single. spiniform. Urosome of 4 somites. Genital somite symmetrical,
or asymmetrical. Appendicular caudal setae highly developed, straight, smoothly curved, or
geniculated, and usually longer than marginal caudal setae. A1 with extremely long seta on each
of segments 3, 7, 9, 14, 18, 21 and 24; segments 8 and 9 fused. Mx2 very strongly built; Li1-Li5 with
1 short and 2 long setae each; terminal setae of Li1-3 or Li-4 armed with long spinules in addition
to dense rows of short spinules; terminal part of Mx2 with 6 long setae. Mxp very strongly built;
coxa with 1 seta (this seta may be absent), then with groups of 2, 3 and 3 setae from proximal to
distal; Enp2-6 with 4, 3, 2, 3 and 4 setae; medial edge of basis armed either with short spinules
only, or with both short and long spinules. P1 Enp with usually well-developed outer lobe; two
first P1 Exp segments usually unseparated, Exp1 with outer spine that is usually very small or
missing, Exp2 with outer spine and Exp3 with outer spine.
Males
Similar to female, but appendicular caudal setae much less developed than in female. A1 with
8-10 segments completely fused; segments 12 and 13 partially fused; long seta found on each
of segments 3, 9, 14, 18, 21 and 24. Mx2 reduced compared with female. P5 large and complex.
Male left P5 Exp2 extends distally into serrated lamella and with digitiform process; Exp3 with
hairy tubercle proximally and distal groups of hairs; segment extended beyond these spines into
spine-like process of various lengths or vestigial.
Genus Euchaeta Philippi, 1843

Reference: Park, 1995.
Females
Pd1 separated from cephalosome by faint line representing partially fused joint, Pd4 and
5 completely fused. Urosome of 4 somites. Genital somite symmetrical, or asymmetrical.
Appendicular caudal seta greatly developed, straight, and much thicker and longer than distal
marginal setae of caudal rami. Forehead in lateral view produced anteriorly into frontal eminence
varying from tall conical process to low ridge, bearing suprafrontal sensilla. Rostrum single,
1-pointed. Mx2 very strongly built, 1 or 2 endopodal setae armed with long spinules in addition
to short spinules. Mxp very strongly built; coxa with a posterior seta close to proximal end (this
seta may be absent); medial edge of basis armed either with short spinules only, or with both
short and long spinules. P1-P4 Exp 3-segmented, two first P1 Exp segments usually unseparated,
P1 Exp1 usually without an outer spine; P1-P2 Enp 2-segmented, P3-P4 Enp 3-segmented. P5
absent.
Males
Similar in habitus to female, except that prosome in lateral view usually with a pointed, varying
in form, process. Rostrum single, 1-pointed. Urosome of 5 somites, appendicular caudal setae
much less developed than in female. A1 with 8-10 segments completely fused; segments 12
and 13 partially fused; long seta found on each of segments 3, 9, 14, 18, 21 and 24. Mx2 reduced
compared with female. Coxal seta of Mxp reduced in both number and size, basis marginal
spinules absent. P1-P4 similar to those of female, but their Exp outer spines poorly developed
and Exp1 and Exp2 of P1 fully separated. P5 large and complex. Left P5 Exp2 extends distally into
serrated lamella and with digitiform process; Exp3 hollow, with hairy tubercle proximally and
distal groups of hairs; distal Exp segment on both right and left P5 tapering into long spine. Right
P5 Exp2 longer than Exp1.

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

Euchaeta concinna Dana, 1849
Female. Length 3.10-3.15 mm. (Fig. 154 A – G; Fig. 156 a – d)

Laterally, dorsal margin of forehead slightly ached. Frontal eminence large and conical. Rostrum
small, slightly curved backward. Prosome posterolateral corners angular, asymmetrical, left side
slightly longer. Genital somite strongly asymmetrical dorsally as well as ventrally, with a large
toothlike outgrowth midway along right side and a concpicuous notch on left side about 2/5
length of somite from proximal end. Only 1 seta of Mx2 Enp armed with long spinules. P2 Exp2
outer spine large, overreaching following outer spine; marginal lobe bearing outer spine also
large, separated from segment by a deep incision. All other outer spines of similarly small size.
Male. Length 2.61-2.68 mm. (Fig. 155 I – J; Fig. 157 a, b)

Laterally, dorsal margin of forehead slightly ached. Frontal eminence conical, highly pronounced.
Rostrum relatively short, pointing straight downward. Prosome posterolateral corners in lateral
view produced into a rounded lappet. A large toothlike process on posterodorsal margin of
prosome. Serrated lamella of left P5 Exp elongate, hollow, with uniformly small teeth on medial
and lateral margins and long teeth on distal margin.
Fig. 154. Euchaeta concinna. Female.

A. Ur, dorsal; B. Gns, right lateral view; C. Gns, left lateral view; D. Gns, ventral view; E. P2; F.
forehead, left lateral view; G. Mx2 Enp.

Fig. 155. Euchaeta concinna. Male.
I. Ur; left lateral view; H. forehead right lateral view; J. distal end of serrated lamella of left P5 Exp.
b
a
a b
c d
Fig. 156. Euchaeta concinna. Female. a. dorsal view Fig. 157. Euchaeta concinna. Male. a. dorsal view; b.
(with eggs sac); b. dorsal view; c. right lateral view; left lateral view.
d. Gns dorsal view.

Euchaeta rimana Bradford, 1974.
Female. Length 2.80-4.30 mm. (Fig. 158 a – j)

Posterior thoracic corners slightly asymmetrical, more produced on right. Left side of genital
somite in dorsal view without prominent projection, right side with moderate projection that
obscures more ventral projection. Genital field in ventral view with genital pads almost meeting
in midline, right pad triangular.
Male. Length 3.11-4.10 mm. (Fig. 159 a – d)

Serrate lamella of left P5 Exp segment 2 without distal smooth excavation separating inner distal
fine teeth and coarser outer teeth. Thick spine arising at base of serrate lamella less than half
length of lamella.
Fig. 158. Euchaeta rimana. Female.

A. lateral right view; B-E. Gns (dorsal, right lateral, left lateral, ventral, respectively); F. Exp of P1; G. Exp3
of P2. E. rimana, male H. lateral left view; I. P5; J – terminal part of left P5 Exp.
(Source: Bradford et al., 1983).

a
Fig. 159. Euchaeta rimana. Male. a. dorsal view; b. anterior part of cephalosome; c. P5; d. terminal part of left P5 Exp.

Family Centropagidae Giesbrecht, 1892
References: Gurney, 1931; Bayly and Arnott, 1969 (cited by Bradford-Grieve et. al., 1999).
Females
Rostrum always present, with 2 filaments. Ce abd Pd1. Pd4 and 5 separate, latter somite rounded or
often with asymmetrical spiniform posterior corners. Urosome usually of 3 somites, Genital somite
often with asymmetrical spine armature; Ur2 may also bear processes. A1 24-25-segmented.
A2 Exp and Enp almost equal in size; coxa with 1 seta, basis with 2 setae; Enp 2-segmented; Exp
7-segmented. Mx2 well-developed. Mxp relatively small. P1-5 Exp and Enp 3-segmented; apical
spine of Exp strongly serrate. P5 biramous, natatory, coxa and basis without setae, Exp segments
1 and 2 with or without 1 outer edge spine, segment 2 with strong inner spine-like process, Exp
segment 3 with or without 2 outer edge spines, 1 terminal serrate spine and 2 or 4 inner edge setae;
Enp segments 1 and 2 with 0-1, 1 inner edge seta respectively, Enp segment 3 with 4-6 setae.
Males
Mouthparts and P1-4 identical to female apart from male A1, and sometimes P2. Urosome of 4-5
somites. A1 prehensile on right. P5 asymmetrical, usually resembling other swimming legs, rami
2 or 3-segmented, right exopod variously transformed into grasping organ, sometimes endopod
atrophied.
Genus Centropages Kroyer, 1848

Reference: Vervoort, 1964 (cited by Bradford-Grieve, 1999).
Females
Rostrum always present, with 2 filaments. Cephalosome and Pd1, Pd4 and Pd5 separate, latter
somite rounded or often with asymmetrical spiniform posterior corners. Urosome of 3 somites,
Genital somite usually asymmetrical, often with asymmetrical spine armature; Ur2 may also bear
processes. A1 24-25-segmented. A2 Exp and Enp almost equal in size; coxa with 1 seta, basis with
2 setae; Enp 2-segmented; Exp 7-segmented. Mx2 well-developed. Mxp relatively small. P1-5 Exp
and Enp 3-segmented. P2-P4 Exp3 with 3 outer-edge spines. P5 coxa and basis without setae;
Exp2 with strong inner spine-like process.
Males
Urosome of 5 somites. A1 prehensile on right. Mouthparts and P1-4 identical to female. P5
asymmetrical, usually resembling other swimming legs, basis not ornamented. Left Exp
2-segmented; Exp1 with 1 outer edge spine; Exp2 with 3 outer edge spines and 1 small terminal
spine; both endopods as in female. Right Exp 3-segmented, in the form of a chela.
Key to Centropages species

Females
1. Posterior borders of prosome pointed and have an accessory spine on the interior border of
this spine; A1 with spines on segments 1, 2, and 5 …………… …………………… C. furcatus
Posterior borders of prosome pointed, without an accessory spine on the interior border of this
spine; A1 without spines on anterior margin …………………2
2. Posterior border of prosome weakly pointed. Genital somite symmetrical, with spine on ventral
surface …………………………………………………………….………………… C. orsinii
Prosome posterior corners prolonged into long asymmetrical spines. Genital somite without
ventral spine; right P5 Exp 2-segmented, Exp1 inner edge projection large, swollen at its base, with
tubercles along inner edge …..………………………………………………………C. tenuiremis
Males
Posterior borders of prosome pointed and have an accessory spine on the interior border of this
spine; A1 with spines on segments 1, 2, and 5 …..…………………………………C. furcatus
Posterior corners of prosome pointed, without an accessory spine on the interior border, slightly
asymmetrical; right P5 Exp2 rounded protrusion with small tubercles along outer edge …..…
……………………………..………………………………..………...……………… C. tenuiremis

Kuwait City from Ras Al-Ardh - Photo by Dr. V. Skryabin - Kuwait Institute for Scientific Research

Centropages furcatus (Dana, 1849)
Female. Length 1.68-1.78 mm. (Fig. 160 A – H; Fig. 161 a, b)

Head with a ventral ball-like eye. Posterior borders of prosome pointed and have an accessory
spine on the interior border of this spine. Genital somite without spines on ventral surface; anal
segment is not quite symmetrical and more than twice the length of Ur2; caudal rami relatively
slender. A1 with spines on segments1, 2, and 5. P5 Exp2 inner edge spine does not reach distal
border of Exp3, being larger in right side.
Male. Length 1.54-1.66 mm. (Fig. 160 I – J; Fig. 162 a, b)

Prosome posterior border slightly asymmetrical, pointed, with an accessory spine on the interior
border; left side more protruding than right. Caudal rami slender. A1 with spines on segments1,
2, and 5. Right P5 Exp2 extension with rounded protrusion on proximal portion; Exp3 claw stout
and with spine on inner margin and 2 spines on outer margin; left Exp2+3 with long terminal
fixed appendages.
Fig. 160. Centropages furcatus.

Female: A. dorsal view; B. right lateral view; C. Ur, right lateral view; D. Ur, ventral; E. anterior
part of head and A1, dorsal view; G. leftP5 Exp; H. right P5 Exp.
Male: I. dorsal view, F. anterior part of head and A1, dorsal view; K. right P5 Exp; J. left P5 Exp.

a b
Fig. 161. Centropages furcatus. Female. a. dorsal view;

b. left lateral view.
a b
Fig. 162. Centropages furcatus. Male. a. dorsal view;


Centropages orsinii Giesbrecht, 1889
Female. Length 1.61 mm. (Fig. 163 A – G; Fig. 164)

Posterior border of prosome weakly pointed. Caudal rami about twice as long as wide. Genital
somite symmetrical, with spine on ventral surface. A1 without spines on segments 1, 2, and
5 and when held straight back does not extend beyond the caudal rami. P5 Exp2 inner spine
curved, usually bordered by small spines.
Centropages tenuiremis Thompson and Scott, 1903
Female. Length 1.74-1.75 mm. (Fig. 165 A – G; Fig. 166 a, b)

Prosome posterior corners prolonged into long asymmetrical spines, left directed backward
and laterally, right – backward; their points extend beyond distal border of Genital somite. A1
without teeth on anterior margin. Genital somite, excluding the ventral hook, unarmed, slightly
swollen laterally in both left and right anterior parts. Caudal rami symmetrical, 2.5 times as long
as Ur3. P5 asymmetrical. Right Exp 2-segmented, Exp1 inner edge projection large, swollen at its
base, with tubercles along inner edge; distal outer spine long, extending beyond midlength of
Exp2. Left Exp 3-segmented, Exp2 inner spine-like process shorter than that of right leg.
Male. Length 1.60 mm. (Fig. 167)

Prosome resembles that of female, posterior corners slightly asymmetrical, left side more
protruding than right. Caudal rami slender. Right P5 Exp2 extension with rounded protrusion
on proximal portion, with small tubercles along outer edge; Exp3 claw stout and with 1 spine
on inner margin and 1 pointed process on outer margin.
Fig. 163. Centropages orsinii. Female.

A. dorsal view; B. right lateral view; C. Ur,
dorsal; D. Ur, right lateral view; E. Ur, ventral,
F. forehead, right lateral view; G. P5.

a b c
Fig. 164. Centropages orsinii. Female. a.dorsal view; b. left lateral view.
Male. c. dorsal view.
a b
Fig. 165. Centropages tenuiremis. Female. Fig. 166. Centropages tenuiremis. Female. a. dorsal
A. dorsal view; B. A1; C. last thoracic somite and Ur, view; b. Ur (right lateral view).
dorsal; D. last thoracic somite and Ur, right lateral
view; E. P5, F. right P5 Exp; G. Gns, dorsal.
Fig. 167. Centropages tenuiremis. Male. dorsal view.

Family Pseudodiaptomidae Sars, 1902
References: Madhupratab and Haridas, 1978; Walter, 1986 (cited by Bradford-Grieve et. al., 1999).
Females
Small copepods slightly over 1 mm. Usually with characteristic eye composed of pigmented
spot surrounded by refractile lens. Cephalosome and Pd1 fused or separate, Pd4 and 5 fused or
separate; posterior metasomal cornes may be rounded or variously decorated with large posterior
spines or rows of spinules. Cephalon with pair of rostral filaments. Urosome of 2-4 somites, may
be asymmetrical; genital somite very variable (symmetrical, asymmetrical, ornamented with spines
and/or spinules, or undecorated), caudal rami tend to be elongate and may be asymmetrical. A1
symmetrical, usually 22-segmented. A2 exopod slightly longer than endopod, Endopod with fusion
between segments and with basis. Md gnatobase with numerous small teeth. P1-4 biramous, with
3-segmented rami; coxa and basis often decorated with spinules on anterior, posterior and lateral
surfaces, also sometimes on anterior surfaces of endopod and exopod. P5 uniramous or biramous;
not natatory; with 3 Exp segments (the terminal 2 may be fused); may be slightly asymmetrical.
Ovisacs present.
Males
Body of similar to female, urosome of 5 somites. A1 asymmetrical; left 22- or 24-segmented; right
with 20-21 segments, geniculate between segments 18 and 19; terminal part beyond joint with 2-4
segments. P5 uniramous or biramous, very asymmetrical; Exp 2- or 3-segmented; spines, exopod
segments, and endopod, if present, variously modified into a form which appears to be adapted for
clasping.
Genus Pseudodiaptomus Herrick, 1884

Reference: Walter, 1986 (cited by Bradford-Grieve, 1999).
Females
Small copepods slightly over 1 mm. Usually with characteristic eye composed of pigmented spot
surrounded by refractile lens. Cephalosome and Pd1 fused or separate, Pd4 and 5 fused; Prosome
posterior corners often extended into points. Cephalosome with a pair of rostral filaments. Urosome
of 2-4 somites, Genital somite variously ornamented, may be asymmetrical; caudal rami tend to be
elongate and may be asymmetrical. A1 symmetrical, usually 22-segmented. A2 Exp slightly longer
than Enp, Endopod with fusion between segments and with B2. Md gnatobase with numerous
small teeth. P1-4 biramous, with 3-segmented rami; coxa and basis often decorated with spinules
on anterior, posterior and lateral surfaces, also sometimes on anterior surfaces of endopod and
exopod. P5 uniramous, with 3-segmented exopod; Exp3 spiniform, distally produced, usually equal
in length or longer than Exp2 spiniform process, with both margins hairy, also with a proximomedial,
spiniform process.
Males
Body of similar form to female, urosome of 5 somites. A1 asymmetrical; left 22- or 24-segmented;
right with 20-21 segments, geniculate between segments 18 and 19; terminal part beyond joint with
2-4 segments. P5 uniramous or biramous, very asymmetrical; Eopodite 2- or 3-segmented.

Pseudodiaptomus arabicus Walter, 1998
Female. Length: 1.2-1.7 mm. (Fig. 168 A, B, E, G; 169 a, b)

Head separate from pedigerous segment 1; pedigerous segment 5 with pointed posterior
corners. Urosome 4-segmented. Urosome segment 1 with paired genital flaps, egg sac single.
Dorsal side of Genital somite with pointed protrusion. Leg 5 symmetrical, exopod segments 1
and 2 with distolateral outer spine. Exopod 3 elongate with small proximomedial process and
plumose.
Male. Length: 1.1-1.45 mm. (Fig. 168 C, D, F; Fig. 170 a – c)

Head separate from pedigerous segment 1; pedigerous segment 5 with pointed posterior
corners. Urosome of 5 segments; urosome segment 2 with ventral paired spinule rows. Leg 5
asymmetrical, exopod of right leg 3-segmented; left leg 2-segmented; both legs with endopods.
Right leg endopod bifid, both branches simple, medial with terminal seta longer than lateral,
lateral branch distally rounded and heavily spinulated. Left leg endopod elongate, slightly
swollen after mid-length.
a b
Fig. 168. Pseudodiaptomus arabicus. Fig. 169. Pseudodiaptomus arabicus. Female.

Female: A. dorsal view; B. right lateral view; E. Ur, right a. dorsal view; b. left lateral view.
lateral view; G. P5. Male: C. dorsal view, D. left lateral
view, F. P5.
Fig. 170. Pseudodiaptomus arabicus. Male.

a b c a. dorsal view; b. right lateral view; c. P5.

Pseudodiaptomus ardjuna Brehm, 1953
Female. Length: 1.1-1.57 mm. (Fig. 171 a, b)
Male. Length: 1.08-1.32 mm. (Fig. 172 a–c)

Comparison of figures 168 and 171-172 indicates the differences between P. ardjuna and P.
arabicus. Head of P. ardjuna (both female and male) is narrower than head of P. arabicus. The
genital somite of P. ardjuna female is without pointed protrusion. P5 of male asymmetrical as in P.
arabicus but left leg endopod broad and rounded in terminal part; right leg Exp1 with bifid seta.
a b
Fig. 171. Pseudodiaptomus ardjuna. Female. a. left lateral view; b. Gns. Arrow indicates genital somite.
a b c
Fig. 172. Pseudodiaptomus ardjuna. Male. a. left lateral view; b. P5, c. right P5 terminal part.

Family Temoridae Giesbrecht, 1892
Females
Body of varying form, in some cases rather short and stout, in other cases comparatively slender;
anterior cephalosome tends to be widest part of body. Cephalosome separate from Pd1, anterior
cephalosome unarmed or proved with 2 soft rostral filaments or R 1-2-pointed (Temoropia); Pd4
and 5 fused, partly fused or separate. Urosome of 3-4 somites, caudal rami of different structure
in different genera, sometimes elongate. A1 24- or 25-segmented. A2 Enp more or less equal in
length to Exp, Exp 6-7-segmented. Md with broad gnatobase with 1 large tooth set slightly apart
from remaining teeth, endopod segment 1 often fused to basis. Mxp of moderately size, may be
slightly modified with recurved endopod or with fusion between endopod segments. P1-4 with
the endopod 1-, 2- or 3-segmented; exopod segments 1 and 2 may be fused.
Males
Segmentation of body and legs as in female except urosome of 5 somites. Right A1 distinctly
geniculate. P5 not natatory, usually without endopod; larger than those of female and prehensile,
often pincer-like on one side, 2-4-segmented with common basal segment.
Genus Temora Baird, 1850

Females
Body short and compact. Head vaulted dorsally. Head remarkably dilated with a posterodorsal
prominence, anterior head with 2 slender rostral filaments. Pd4 and 5 fused. Urosome of 3 somites;
genital somite comparatively short and hardly protuberant ventrally; caudal rami narrow and
elongate, sometimes asymmetrical, setae comparatively short and of the usual number, one is on
the outer border some distance from others. A1 slender, 24-segmented. A2 Exopod 7-segmented,
scarcely longer than endopod. Swimming legs with endopod small and 2-segmented; P2-4 Exp1
and Exp2 partly fused, Exp3 with 3 outer edge spines and a terminal coarsely toothed spine. P5
small, 3-segmented, first two simple, last segment dentate terminally. No ovisac present.
Males
Segmentation of body and legs as in female except urosome of 5 somites. Right A1 distinctly
geniculate. P5 asymmetrical, left leg much larger, 4-segmented, segment 2 produced on inner
edge into a long curved thumb-like process, which opposes the 2 terminal segments; right leg
3-segmented; terminal segment incurved, claw-like.
Key to Temora species

Females
Posterior corners of prosome pointed; Ur3 and caudal rami hardly asymmetrical
…………………………………………………………….… ……………………… T. discaudata
Posterior corners of prosome rounded; one terminal caudal seta conspicuously thickened
…………………………………………………………….… ………………………..T. turbinata
Males
Posterior corners of prosome pointed; right P5 terminal hook very long
…………………………………………………………….… ……………………….T. discaudata
Posterior corners of prosome rounded; right P5 terminal segment short
…………………………………………………………….… ……………………….T. turbinata

Temora discaudata Giesbrecht, 1889
Female. Length: 1.6-2.0 mm (1.7-2.0 mm). (Fig. 173 A – D; Fig. 174 a – c)

Quite a large robust species; posterior metasome segments produced into spines. Posterior
corners of prosome pointed; Ur3 and caudal rami hardly asymmetrical; right side of anal segment
elongated, right caudal rami slightly bent and thickened. Long slender caudal rami asymmetrical.
P5 uniramous and symmetrical, inner spine of segment 3 longer than the 2 terminal spines,
which are almost equal in length.
Male. Length: 1.7-1.85 mm (1.7-1.9 mm). (Fig. 173 E, F; Fig. 175 a, b)

Lateral angles of the last metasome segment pointed and slightly asymmetrical; right Al
geniculate. Urosome of 5 segments, almost symmetrical, with long slender caudal rami. The
thumb-like process on the basis of the left P5 is wide, the terminal segment is flattened and has
4 marginal spines. The terminal segment of the right P5 is hook-like and sharply bent backwards.
Fig. 173. Temora discaudata.

Female: A. dorsal view; B. left lateral view; C. Ur and last thoracic somite, dorsal view; D. P5.
Male: E. dorsal view; F. P5.

a b c
Fig. 174. Temora discaudata. Female. a. dorsal view; b. left lateral view; c. Ur dorsal view.
a b
Fig. 175. Temora discaudata. Male. a. dorsal view; b. right lateral view.

Temora turbinata (Dana, 1849)
Female. Length: 0.9-1.6 mm (1.05-1.61 mm). (Fig. 176 A – E; Fig. 177 a, b)

Body widest at cephalosome, tapering anteriorly to urosome; last metasome segment rounded.
Urosome 3-segmented; anal segment symmetrical, shorter in length than previous segment;
caudal rami almost symmetrical, very long and slender. P5 symmetrical, inner marginal spine of
the 3 terminal spines shorter than the other two.
Male. Length: 0.9-1.4 mm (0.9-1.56 mm). (Fig. 176 F – H; Fig. 178 a, b)

Body shape similar to female. Urosome quite symmetrical; caudal rami long and slender; P5
asymmetrical, the left with a claw; thumb of claw is slender and gradually curved.
Fig. 176. Temora turbinata.

Female: A. dorsal view; B. left lateral view; C. Ur, dorsal view; D. Ur, left lateral view; E. P5. Male: F.
dorsal view; G. Ur, dorsal view; H. P5.

a b
Fig. 177. Temora turbinata. Female. a. dorsal view; b. left lateral view.
a b
Fig. 178. Temora turbinata. Male. a. dorsal view; b. left lateral view.

Family Candaciidae Giesbrecht, 1892
Females
Total length 2-4 mm. Body relatively robust, anterior cephalosome rectangular in dorsal view
with conspicuous lateral construction anteriorly. Many species possess dark pigment which
gives dark brown or black color to some parts of body. Cephalosome and Pd1 separated, Pd4
and 5 fused and extended into pointed, often symmetrical, processes, rarely rounded. Rostrum
atrophied. Urosome of 3 somites; genital somite often spinose or asymmetrically swollen; Ur2
is sometimes asymmetrical. Caudal rami short with 6 setae. A1 symmetrical 24-segmented,
segments 24 and 25 fused. Md gnatobase narrow with few teeth, basal tooth usually divided
into one or more pointed cusps or may be simple. Mx2 very large; Li1 and 2 rudimentary with 3
and 2 setae respectively; Li3 and 4 with 2 setae each; Li5 with 2 stout spines of variable length
and thickness and very small spinule or proximal spine may be one-half length and considerably
thinner than distal spine; endopod with 3 stout spines and 4 small setae. Mxp small.P1-4 with
3-segmented exopods with serrated outer margins, especially Exp2 and 3, and 2-segmented
endopod (segments 1 and 2 fused). P5 uniramous, not natatory, usually symmetrical.
Males
Body similar to female, but urosome of 5 somites; posterior corners of prosome always
asymmetrical; Genital somite is almost always asymmetrical, bearing process on right side
sometimes with complex apex. Right A1 geniculate, 23-segmented, segments 17-18 and 19-20
fused, with or without denticulate segments at bend. Mouthparts and P1-4 similar to the female.
P5 not natatory, 4-segmented on left and 3-segmented on right; may be chelate on right or
ending in a long feather-like seta.
Genus Candacia Dana, 1846

Reference: Grice, 1963; Bradford-Grieve, 1999.
Females
Total length 2-4 mm. Body relatively robust, anterior cephalosome rectangular in dorsal view
with conspicuous lateral construction anteriorly. Cephalosome and Pd1 separated, Pd4 and
5 fused and extended into pointed, often symmetrical, processes, rarely rounded. Rostrum
atrophied. Urosome of 3 somites; Genital somite often spinose or asymmetrically swollen; Ur2
is sometimes asymmetrical. Caudal rami short with 6 setae. A1 symmetrical 24-segmented,
segments 24 and 25 fused. Md gnatobase narrow with few teeth, basal tooth usually divided
into one or more pointed cusps or may be simple. Mx2 very large; Li1 and 2 rudimentary with 3
and 2 setae respectively; Li3 and 4 with 2 setae each; Li5 with 2 stout spines of variable length
and thickness and very small spinule or proximal spine may be one-half length and considerably
thinner than distal spine; endopod with 3 stout spines and 4 small setae. Mxp small. P1-4 with
3-segmented exopod with serrated outer margins, especially Exp2 and 3, and 2-segmented
endopod (segments 1 and 2 fused). P5 uniramous, usually symmetrical; terminal segments may
end in one or more spine-like processes, or a single long seta; setae may or may not be present
on the inner lateral margins.
Males
Body similar to female, but urosome of 5 somites; posterior corners of prosome always
asymmetrical; genital somite is almost always asymmetrical, bearing process on right side
sometimes with complex apex. Right A1 geniculate, 23-segmented, segments 17-18 and 19-20
fused; with teeth present on one or more segments in the geniculate region. Mouthparts and
P1-4 similar to the female. P5 4-segmented on left and 3-segmented on right; right leg chelate.

Candacia bradyi (Scott, 1902)
Female. Length 1.86-1.94 mm. (Fig. 179 A – C; Fig. 180 a, b)

Prosome posterior corners each terminate in a short spine. Genital somite symmetrical in dorsal
view and globular in shape, with a well-marked genital swelling. Ur2 produced ventrally in
midline into short spine. Caudal rami about twice as long as broad, slightly asymmetrical, that
on right being slightly broader than on left. A1 23-segmented. P5 terminal segment ending in a
spine-like process, with 3 outer spines and 2 inner seta.
Male. Length 1.74-1.88 mm. (Fig. 179 D – F; Fig. 181)

Prosome posterior corners symmetrical, tip of right process does not reach beyond midpoint of
genital somite. Genital somite produced into a toothed tubercle on right side. Left P5 segment 4
elongate and narrow, with terminal small spines.
a b
Fig. 180. Candacia bradyi. Female. a. dorsal view;

b. right lateral view.
Fig. 179. Candacia bradyi.

Female: A. Ur, dorsal; B. Ur, right lateral view; C. P5.
Male: D. Ur, dorsal, E. Ur, right lateral view, F. P5.
Fig. 181. Candacia bradyi. Male, dorsal view.

Family Pontellidae Dana, 1953
Reference: Silas and Pillai, 1973 (cited by Bradford-Grieve et. al., 1999).
Females and males

Cephalosome separated from edigerous somite 1, often with lateral cephalic hooks; rostrum
bifurcate, ending in two prongs with thickened base, often bearing a lens; eyes usually prominent
with one or two pairs of dorsal subcuticular eye lenses and a medioventral eye lens. Pd4 and
5 fused or separate; last thoracic segment posterior lateral margin often produced posteriad.
Urosome usually asymmetrical, of 1-3 somites in female and 5 somites in male. Female A1
16-24-segmented, with last two segments usually fused; right A1 of male geniculate and strongly
modified. A2 with B2 and Enp1 fused together; exopod small with 5 segments, terminal segment
of exopod shortened, second segment longest. Md blade with 5-7 teeth. Mx1 with B1 large,
B2, exopod and endopod relatively small. Mx2 with distal seta long and robustly developed in
most of the genera, with long setae; B2 and Exp relatively small. P1-4 Exp 3-segmented; P1-4
Enp 2-3-segmented. Female P5 small; Exp 1-2-segmented. Male P5 uniramous, each leg 3- or
4-segmented; right leg chelate.
Genus Calanopia Dana, 1853

Reference: Silas and Pillai, 1973.
Females
Cephalosome with or without lateral cephalic hooks, dorsal cuticular lenses and ventromedial
eye absent; rostrum long, without eye lenses; last thoracic segment posterolateral corners
produced; urosome of 2 somites; A1 17-19-segmented; Mxp with 6 distinct segments; P1 Enp
2-segmented; P5 uniramous, 3-4-segmented.
Males
A1 geniculate with 4 distinct segments distal to hinge; Urosome of 5 somites; P5 forming on right
side an ill-developed chela.
Calanopia elliptica (Dana, 1846, 1849)
Female. Length: 1.6-1.9 mm (1.4-1.9 mm). (Fig. 182 A – D; Fig. 183 a, b)

Cephalosome enlarged, conically rounded. Rostrum bifid, slightly bulged at ending in acute
points. The prosome nearly twice as long as the combined length of the urosome. Long points on
posterior metasome. Urosome segment 2 as long as genital somite. Caudal rami nearly 3 times
as long as broad. P5 asymmetrical, each limb uniramous of 4 segments: left leg longest, exopod
of 2 segments. Exp1 with 1 median and 1 distolateral spine; Exp2 ending in acute spine and carry
2 outer marginal spines; the spines with fine lateral serrations.
Male. Length: 1.4-1.8 mm (1.8-1.8 mm). (Fig. 182 E – G; Fig. 184)

Urosome segment 2 right side distal border produced into a well defined tooth. P5 terminal
segment of the left exopod with a pad of fine hairs, pointed at the distal end with three seta on
the outer margin and a seta on the posterior side. The flattened margin of the right exopod
segment 1 with 3 blunt teeth, while the claw-like third segment has 3 small pointed teeth.

a b
Fig. 183. Calanopia elliptica. Female. a. dorsal view;

Fig. 182. Calanopia elliptica.

Female: A. dorsal view; B. lateral view; C. P5; D. R. Male:
E. dorsal view; F. Ur dorsal view; G. P5.
Fig. 184. Calanopia elliptica. Male, dorsal view. Arrow

indicates the tooth on the urosome.

Calanopia minor A. Scott, 1902
Female. Length: 1.2-1.4 mm (1.4 mm). (Fig. 185 a – c; Fig 186 a, b)

Superficially resembles a small C. elliptica. The last metasome segment produced on each side
into an acute spine; the rostrum pointed. Urosome of 2-somites, the second somite distinctly
longer than the genital somite. The P5 is quite symmetrical with 4 segments; the distal segment
terminates in long inner spine (which is not stout as in Calanopia elliptica) and one short spine,
with a further spine on outside mid segment.
Male. Length: 1.1-1.2 mm (1.2 mm). (Fig. 185 d – f )

The second urosome somite has no spiny process. Antennule extends to the end of the third
thoracic somite; the right Al is geniculate. The basis of the left P5 is swollen toward the proximal
end of the inner margin, the swollen part produced into a small tooth-like spine. The basis of the
right exopod is also swollen. Flattened joint of the first exopod segment, with a seta on the inner
margin; outer thumb is short and naked. The claw-like second segment is spoon- shaped and
without any teeth, but with 2 inner marginal seta.

Fig. 185. Calanopia minor.
Female: A. cephalothorax, dorsal; B. Ur, dorsal; C. P5. Male: d. right A1; E. last thoracic somite and Ur, dorsal; F. P5.
(Source: Tanaka, 1964).
a c b
Fig. 186. Calanopia minor: a. female, lateral view; b. female, dorsal view. Calanopia elliptica: c. female, lateral view.
a and c indicate differences in size of C. minor (a) and C. elliptica (c).

Genus Labidocera Lubbock, 1853
Reference: Giesbrecht, 1892 (cited by Bradford-Grieve, 1999).
Females
Cephalosome separated from Pd1, with or without lateral cephalic hooks and with one pair of
dorsal cuticular lenses and a protuberant ventral eye which extends anteroventrally between
the rostral prongs. Rostrum deeply bifurcate with 2 relatively fine filaments. Pd4 and 5 fused with
corners produced into pointed lobes. Urosome of 2 or 3 somites, Genital somite and caudal rami
usually asymmetrical. A1 23-segmented. A2 with B2 and Enp1
fused together; exopod small with 5 segments, terminal segment of exopod shortened, second
segment longest. Md blade with 3-4 small teeth. Mx1 Exp relatively well-developed. Mxp with
6 distinct segments. P1-4 Exp 3-segmented; P1-4 Enp 2-segmented. P5 biramous, each ramus
1-segmented.
Males
Urosome of 4 or 5 somites, Genital somite and caudal rami symmetrical. Right A1 of male
geniculate, strongly modified, with at least 4 separate segments distal to the hinge between
segments 18 and fused segments 19-21, the middle section expanded. P5 right leg uniramous,
with a chela; left leg sometimes with rudimentary endopod.
Labidocera acuta (Dana, 1849)

Reference: Othman and Toda, 2006
Female. Length 2.88 mm, length ratio prosome to urosome 2.94:1. (Fig. 187 A – E; Fig. 189 a – d)
Body elongated ending in a single spiniform process, cephalon rounded with a median anterior
hook, dorsal eye lenses moderate, separated. Lateral cephalic hook absent.
Thoracic process reaching beyond middle of genital somite. Urosome of 3 somite, genital somite
with a stout distolateral conical process on right margin. P5 asymmetrical, right leg being stouter
and longer than left, Re with three outer, one inner and three apical spines of which medial one
longest; Ri bifurcated at apex.
Male. Length 2.65 mm, length ratio prosome to urosome 2.96:1. (Fig. 188 A – E; Fig. 190 a – d)
Body similar to female, anterior hook more pronounced, dorsal lenses slightly larger and closer
together than in female. Thoracic process asymmetrical, left process similar to females, right
produced into a curved process turned distolaterally and reaching distal end of Ur2. Urosome
of 5 somites, genital somite widest, asymmetrical, left side convex posteriorly, right side armed
on posterior end with pointed process. Right A1 geniculate, segment 17 naked, anterior margin
of segment 18 with row of prominent denticles, extends proximally to almost whole length of
segment 17, fused segments 19-21 with toothed plate extending to 2/3 length of the segment,
segment 22 prolonged distally into spur-like process which is as long as its own segment. P5
asymmetrical, right leg chelate, Re1 orbicular, Re2 short, broader medially, with two inner and
two apical setae. Left leg, Re1 with distolateral spine, Re2 ending in three finger-like processes,
one small cresented basal process and one spine near distal end, inner margin of segment hirsute.
Remarks: In the Arabian Gulf this species was previously recorded only near the Strait of Hormuz.

Fig. 187. Labidocera acuta. Female. Fig. 188. Labidocera acuta. Male.
A. dorsal view; B. lateral view; C. last thoracic somite A. dorsal view; B. lateral view; C. last thoracic somite
and Ur, lateral view; D. last thoracic somite and Ur, and Ur, lateral view; D. last thoracic somite and Ur,
dorsal view; E. P5. dorsal view; E. P5.
(Source: Othman and Toda, 2006). (Source: Othman and Toda, 2006).

a b
c d
Fig. 189. Labidocera acuta. Female. a. dorsal view; b. lateral view; c. P5; d. Ur.

a b
c d
Fig. 190. Labidocera acuta. Male. a. lateral view; b. P5; c. Ur; d. R.

Labidocera bengalensis Krishnaswamy, 1952
References: Silas and Pillai, 1973.
Female. Length: 1.5-2.0 mm (1.41-1.68 mm). (Fig. 191 A – E; Fig. 193 a – c)

Cephalosome anteriorly squarly rounded; rostrum pointed and bent ventrally; dorsal eye lenses
prominant. Urosome of 3 somites. Genital somite elongated and longer than combined lengths
of the second and third urosome somites and caudal rami; genital somite slightly swollen on
its right margin and bears a number of ventral papillae; second urosome somite produced
posteriorly slightly on right margin. Caudal rami asymmetrical, left ramus slightly longer and
broader. P5: with exopod long, slender and bifid at its tip; exopod about 4 times longer than
endopod, the latter being short, stout and with a pointed tip.
Male. Length 1.3-1.6 mm (1.09-1.26 mm). (Fig. 192 A – F; Fig. 194 a – c)

Cephalosome resembles that of female. Dorsal eye lenses well developed, closely packed..
Urosome of 5 somites, caudal rami symmetrical. Right antennule geniculate. Right P5 with a
well-developed chela; palm of hand with an inner marginal blunt process and with a spinous
seta, claw bent inwards at its distal half and with an inner marginal transparent flap; a seta
present proximally along inner margin and second curved seta present distally. Left leg three-
segmented; subterminal segment with a disto-lateral spine; terminal segment with a pair of
stout processes distally and a seta toward outer margin of inner process; inner proximal margin
of segment hirsute.
Fig. 191. Labidocera bengalensis. Female. Fig. 192. Labidocera bengalensis. Male.
A. dorsal view; B. lateral view; C. last thoracic somite A. dorsal view; B. lateral view; C. A1; D. last thoracic
and Ur, lateral view; D. last thoracic somite and Ur, somite and Ur, dorsal view; E. last thoracic somite and
dorsal view; E. P5. Ur, lateral view; F. P5.
(Source: Othman and Toda, 2006). (Source: Othman and Toda, 2006).

a b c
Fig. 193. Labidocera bengalensis. Female. a – dorsal view; b – lateral view; c – P5. Arrow indicates genital somite.
a b c
Fig. 194. Labidocera bengalensis. Male. a. dorsal view; b. lateral view; c. P5.
Arrow indicates the male left P5 terminal segments.

Labidocera kroyeri (Brady, 1883)
Female. Length: 2.2-2.75 mm (2.4-2.75 mm). (Fig. 195 a – d; Fig. 196 a – e)

Cephalosome with lateral cephalic hooks, lateral wings of last metasome somite pointed and
symmetrical. Urosome of 3 somites; anal somite short, genital somite and Ur2 asymmetrical, with
many spines and hooks (variable in appearance); caudal rami symmetrical. P5 endopod reduced
and bifurcate at the end.
Male. Length: 2.0-2.35 mm (1.95-2.36 mm). (Fig. 195 e – g; Fig. 197 a – c)

Lateral cephalic hooks on cephalosome. Lateral corners of the last pedigerous somite
asymmetrical, left one pointed the other one bifurcated. Urosome of 5 somites, genital somite
swollen on left side. Right antennule (A1) geniculate. Right P5 exopod segment 1 claw-like, with
curved thumb, middle portion of the external margin with 2 processes.
Fig. 195. Labidocera kroyeri.

Female: a. dorsal view; b. forehead (right lateral side); c. last thoracic somite and Ur, left lateral side; d. P5.
Male: e. right P5; f. left P5. g. dorsal view.
(Source: a-e. Tanaka, 1964; g. Greenwood, 1979, cited by Conway et al., 2003).

c d
a b
Fig. 196. Labidocera kroyeri. Female. a. dorsal view; b. lateral view; c. Ur, dorsal view; d. Ur lateral view;
e. P5. Arrow indicates urosome.
a b c
Fig. 197. Labidocera kroyeri. Male. a. dorsal view; b. lateral view; c. urosome, dorsal view. Arrows indicate P5 and
corners of the last pedigerous somite.

Labidocera minuta Giesbrecht, 1889
References: Al-Yamani and Prusova, 2003.
Female. Length: 1.6-1.9 mm (1.95-2.05). (Fig. 198 A – G; Fig. 200 a – d)

Cephalosome narrow and rounded, small dorsal eye lenses; small lateral hooks present. Lateral
posterior margin of metasome rounded, but extended on right side with a ventrally pointing
spine; left side without any spine; urosome of 3 somites, genital somite elongated and half
the length of urosome; right anterior and also posterior corner of genital somite with lateral
swellings, in the proximal center of the ventral surface is a small chitinous outgrowth.
Right ventral surface of urosome and second urosome somite with chitinous knobs along right
margin; caudal rami asymmetrical, right side larger. P5 symmetrical, exopod bifurcated at tip; the
endopod is normally bent outwards over the exopod, short, broad-based and ends in an apical
spine plus a rather shorter spine on the inner margin. Two minute spines on outside margin of
exopod (Greenwood, 1979) or one spine (Silas and Pillai, 1973).
Male. Length: 1.4-1.6 mm (1.65 mm). (Fig. 199 A – F; Fig. 201 a – c)

Cephalosome similar to female, but dorsal eye lenses are very large and close together. Last
metasome segment asymmetrical, shorter point on left, extended into a longer flat curved tip
on right. Urosome of 5 somites, genital somite broader than long, with a small lobe at the right
posterior margin. Right A1 geniculate. Right P5 with a claw on the first exopod segment, thumb
short with a seta at the base, terminal segment curved, with 3 setae along internal margin and
two seta distally. Left leg terminal segment with at least 3 lobe-like processes.
Fig. 198. Labidocera minuta. Female. Fig. 199. Labidocera minuta. Male.
A. dorsal view; B. right lateral view, C. Ur, dorsal; D. Ur, A. dorsal view; B. lateral view, C. A1, segments 21-
right lateral view; E. Ur, ventral; F. R; G. P5. 24; D. last thoracic somite and Ur, lateral view; E. last
thoracic somite and Ur, dorsal view; F. P5.
(Source: Othman and Toda, 2006)

a c b
d
Fig. 200. Labidocera minuta. Female. a. dorsal view; b. left lateral view; c. Ur left lateral view, d. P5.
a b c
Fig. 201. Labidocera minuta. Male. a. dorsal view; b. right lateral view; c. male P5.

Labidocera sp.
Reference: Al-Yamani and Prusova, 2003
Female. Length: 2.8-3.35 mm (3.02-3.51 mm). (Fig. 202 A – G; Fig. 204 a – e)

Cephalosome separated from prosome, without lateral hooks and with 1 pair of dorsal cuticular
lenses and a protuberant ventral eye, which extends anteroventrally between the rostral prongs.
Last thoracic somite posterior wings acutely pointed slightly asymmetrical, with right corner
more elongate. Urosome of 2 somites. Genital somite strongly asymmetrical, with large dorsal
projection on the right side. Caudal rami asymmetrical, left ramus inflated. P5 asymmetrical.
Male. Length: 2.55-3.2 mm. (Fig. 203 H – L; Fig. 205 a – c)

Urosome of 4 somites, genital somite and caudal ramus symmetrical. P5 right leg uniramous,
with a chela; left leg exopod 2-segmented, endopod rudimentary, in a form of curved seta, with
hairs on apical end.
Fig. 202. Labidocera sp. Female.

A. last thoracic somite and Ur, dorsal view; B. anterior
head, dorsal view; C. anterior head, left lateral view; D.
P1; E. R; F. P5; G. Gns, dorsal view (another specimen).

Fig. 203. Labidocera sp. Male.
H. last thoracic segment and Ur, dorsal view; I. anterior
head, dorsal view; J. anterior head, right lateral view; K.
A1; L. P5.
c
d
Fig. 204. Labidocera sp. Female.

a. dorsal view; b. right lateral view; c. Ur, dorsal view;
d. Ur, right lateral view; e. P5.
a e b
c
Fig. 205. Labidocera sp. Male.
a. dorsal view; b. right lateral view; c. P5.
a b

Genus Pontella Dana, 1846
Cephalosome with lateral hooks, usually without a crest; with one pair of cuticular lenses and
usually with an additional lens in rostrum; last metasome somite usually with pointed lateral
lobes; female urosome of 2 or 3 somites and asymmetrical, male urosome of 4 or 5 somites,
symmetrical. Right male antennule geniculate. Female P5 biramous with exopods and endopods
of 1 segment, male P5 uniramous.
Pontella danae var. ceylonica* Thompson and Scott, 1889

Female. Length: 2.7-3.2 mm. (Fig. 206 a – e)

Cephalosome with dorsal eye lenses and ventral lenses well developed; rostrum bifid, tapers
to tip; rostral lens feebly developed; prosome more or less of same width throughout. Last
metasome somite asymmetrically produced posteriorly into acuminate spines, left lobe longer
than right. Urosome of two somites, genital somite somewhat globular; a blunt projection
was observed on mid-dorsal margin of genital somite, perceptible in lateral view; caudal rami
asymmetrical, right ramus distinctly broader and longer and carries a marginal fold medio-
dorsally, middle three setae of right ramus bulbous at bases. A1 relatively short and consists of
23 segments. P5 asymmetrical, left ramus distinctly larger; stout and ends in acuminate tips; left
exopod provided with three outer marginal spinules of which distal most one is large; endopods
of both legs asymmetrically bifid at tip.
Remarks: Sewell (1932) pointed out that the specimens collected by A. Scott (1909) from the
Malay Archipelago was intermediate in structure between the Pacific form and var. ceylonica of
Indian waters, and that they form a series rather than separate varieties. Specimens from Malay
Archipelago were characterized by the structure of P5, which resembles that of var. ceylonica
but the caudal rami description agrees with Giesbrecht’s original description. In the shape of the
body the typical Pacific form tapers gradually from Pd2 to Pd5, while in specimen for the Indian
waters the prosome is nearly of the same length throughout. The shape of the genital somite is
also more quadrate in the Pacific form, while it is more globular in Indian specimens.
Silas and Pillai, (1973) remarked that specimens from Indian Ocean fit in with the description of
var. ceylonica, P5 also shows different spinulations for exopod, and the caudal setae are distinctly
bulbous on the right ramus. One additional feature noted in the material at hand is the presence
of short processes on the dorsal margin of genital somite.
Sewell (1932) remarked that although he considered the two forms, P. danae var. ceylonica and
P. investigatoris (male only) as being respectively the female and male of a single species (1914),
his studies on the growth-factor on these material proved that the males showed distinct growth
factors and P. investigatoris should be regarded as a separate species.
*In the present material females that agree with the description of P.danae var. ceylonica and
males that agree with the description of P. investigatoris were found.

c d
a b
Fig. 206. Pontella danae var. ceylonica. Female. a. dorsal view; b. lateral view; c. Ur, dorsal view; d. Ur, lateral view; e. P5.

Pontella investigatoris* Sewell, 1912
Male. Length 2.7 mm (2.9-3.2 mm). (Fig. 207 A – G; Fig. 208 a – d)

Body robust; dorsal eye lenses, ventral lenses and rostral lenses are well developed and
conspicuous; urosome of five somites; caudal rami asymmetrical, right ramus being stouter, rami
being nearly two times as long as broad. Right antennule geniculate; segments 18 and fusion
segments 19-21 carrying denticulated plates on their dorsal margins; segment 14 with a long
spire, carrying a small flagella at its tip; denticulated plate on segment 18 with sharp teeth and
extends proximally over segment 17; segment 19-21 with two toothed plates, both armed with
sharp, villiform teeth; segment ends in a falcate spur; segments 22-25 fused completely. Legs
P5: right P5 chelate; thumb well developed, in the form of a curved stout spine; inner margin
of hand with a quadrate process, dorsal margin of which is crescentic and this process carries a
seta toward its base; claw curved, elongated and with three inner marginal and one outer distal
spine; left leg: terminal segment short, and provided with one outer marginal spine, two terminal
spines and a flagelliform process; inner margin of segment provided with two patches of hairs;
subterminal segment with a distolateral spine.
Remarks: Sewell (1914) considered this species, which is described from male specimens only,
as the male of P. danae var. ceylonica. Later (1932) he drew attention to the differences, chiefly
in the growth factors of the two forms and opined that they were different. Almost throughout
the collections from where P. investigatoris were found, P. danae var. ceylonica was also collected.
*In the present material females that agree with the description of P.danae var. ceylonica and
males that agree with the description of P. investigatoris were found.
Fig. 207. Pontella investigatoris. Male.

A. dorsal view; B. forehead lateral; C.
right A1; D. A2; E – Md; F. Mxp; G. P5.
(Source: Fazal-Ur-Rehman, 1974).

a b c
Fig. 208. Pontella investigatoris. Male. a. dorsal view; b. lateral view; c. right P5; d. right A1.

Genus Pontellopsis Brady, 1883
References: Silas and Pillai; 1973; Al-Yamani and Prusova, 2003; Conway et al., 2003.
Head without lateral hooks, dorsal or rostral lenses. Last pedigerous somite with lateral points,
usually asymmetrical in the male. Urosome asymmetrical in both sexes, of 1 or 2 somites in the
female, of 5 somites in the male. The male right antennule is geniculate, the terminal section with
2 segments. Male urosome segment 3 with a projection on the right.
Pontellopsis herdmani Thompson and Scott, 1903
Female. Length: 1.7-2.1 mm (1.9-2.32 mm). (Fig. 209 a – d; Fig. 211 a – c)

Body stout, rostral base prominent on dorsal view. Urosome asymmetrical and of two segments.
Genital somite longer than segment 2 of urosome; segment dorsally towards mid-dorsal margin
on the left side with a small spine and with a well developed spine on its right distal corner;
a short indentation present on its left distal corner; a short indentation present on its left
distal corner. Caudal rami slightly asymmetrical, right one being perceptibly broader than left.
P5 symmetrical; exopod with two small outer marginal spines and with subequally bifid tip;
endopod bifid at its tip, both rami curved inwards.
Male. Length: 1.5-1.9 mm (1.76-2.0 mm). (Fig. 210 A – D; Fig. 212 a, b)

Cephalosome resembles that of female. Posterior corners of metasome 5 modified into a
rounded lobe on its left side and into an acuminate long spine on its right side, which is
curved inwards; tip of this process reaches mid-margin of urosome somite 3. Right antennule
geniculate; segment 14 with a moderately long spine with a flagellum at its tip; segment 17
with a dorsal plate, finely serrated; 18 with dorsal denticles more triangular and well packed;
segment 19 of fusion segment 19-21 with 5-7 flagellum like setae arranged radially. P5: right
leg with a chela; thumb of chela is long, exceeds length of claw and is serrated at its apex; left
leg with terminal segment carrying two subequal spins distally and one outer marginal spine;
inner margin with hand of setae; subterminal segment with a distolateral spine.
Fig. 209. Pontellopsis herdmani. Female. Fig. 210. Pontellopsis herdmani. Male.
a. dorsal view; b. Ur, dorsal; c. Ur, lateral; d. P5. A. dorsal view; B. Ur dorsal; C. A1; D. P5.
(Source: Greenwood, 1979). (Source: Othman and Toda, 2006)

a b c
Fig. 211. Pontellopsis herdmani. Female. a. dorsal view; b. lateral view; c. Ur right lateral view.
a b
Fig. 212. Pontellopsis herdmani. Male. a. dorsal view; b. lateral view.

Family Acartiidae Sars, 1903
References: Bradford, 1976; Huys and Boxshall, 1991 (cited by Bradford-Grieve, 1999).
Females
Small, more or less slender copepods. A single eye present. Cephalosome and Pd1 separate, Pd4
and 5 always fused, posterolateral corners of prosome rounded or pointed. Urosome of 3 somites,
anal operculum may be on anal segment or anus may open between the last two urosome
somites into a dorsal grove on anal segment; caudal rami generally slightly asymmetrical, short
or long, sometimes fused with anal segment, with 6 setae. Without a rostrum with or without
paired filaments. Upper lip large, prominent and trilobed. A1 17-22-segmented, many segments
with long setae. A2 Enp 3-segmented; B2 and Enp1 fused. Md with well-developed blade with a
large separate tooth on one border. Mx1 with reduced lobes and setation; Enp apparently absent.
Mxp reduced and highly modified, B1 with 5-6 long setae; B2 with 1 short thick seta; Enp with
fused segments with 4-5 short thick setae. P1-4 slender and delicate with long natatory setae;
distolateral borders of P2-4 Exp expanded into a conspicuous tooth. P1 B2 may have small outer
edge spine or seta; P2-4 Exp terminal spine with outer edge teeth. P5 usually uniramous, small,
symmetrical, 3-segmented, B1 and coupler may be fused or B1-2 and the coupler may be fused
(Paracartia); B2 with an outer seta; exopod in the form of a spine or elongate seta; small endopod
present in Acartiella.
Males
Urosome of 5 somites. A1 usually prehensile on right side only, with 12-18 segments; middle
segments may be greatly inflated. Mouthparts more or less the same in both sexes. P5 uniramous,
asymmetrical, attached to a plate formed from fusion of B1 and coupler; B2 usually asymmetrically
developed with outer distal seta; left leg with 2-segmented exopod (3-segmented in Paracartia
africana), Exp2 with variously decorated tip; right leg usually with 3-segmented exopod, Exp1-2
often with inner processes; Exp3 opposed to the process on Exp2 to form a type of subchela.
Genus Acartia Dana, 1846

Females
Small, slender copepods. Single eye present. Cephalosome and Pd1 separate, Pd4 and 5 always
fused, posterodorsal corners of prosome rounded or pointed. Urosome of 3 somites, anal
segment without an anal operculum, as the anus opens between the last 2 urosome segments
into a dorsal grove on the anal segment. Caudal rami short, separated from Ur3. Rostrum
absent or present as 2 filaments. A1 17-22 segmented, many segments with long setae. A2 Enp
3-segmented, Exp 4-segmented; B2 fused with Enp1 which is long and slender and bears 9 setae;
Exp shorter than Enp1. Mxp reduced and highly modified, coxa with 5-6 long setae, basis with
1 short thick seta, endopod with fused segments with 4-5 short thick setae. P1-4 slender and
delicate with long natatory setae. P1 Exp1-2 with a long slender outer distal spine each, and
with 2 spines on Exp3. P2-4 Exp terminal spine with outer edge teeth. P5 uniramous, very small,
symmetrical, 3-segmented; B with 1 outer seta; Exp3 modified into a long, slender spine; small
Enp present in Acartiella.
Males
Urosome of 5 somites. A1 prehensile on right side, middle segments may be greatly inflated. P5
uniramous, asymmetrical; basis usually asymmetrically developed with 1 outer distal seta; left P5
with 2-segmented exopod, Exp2 with variously decorated tip; right P5 usually with 3-segmented
exopod, Exp1 and 2 often with inner processes, Exp3 opposed to process on Exp2 to form type
of subchela.

Beach at the northern coast of Kuwait Bay - Photo by Dr. V. Skryabin - Kuwait Institute for Scientific Research

Acartia (Acanthacartia) fossae Gurney, 1927
Female. Length: 0.9-1.2 mm (1.03-1.4).

Fig. 213 a, b; Fig. 214 A, B
Posterior metasome segment with a row of 4-5 tiny spinules on the dorsoposterior margin. A1
extends to the posterior border of urosome segment 2. Genital somite with 2 pairs of lateral
tufts of hair on anterior part. No spines on this or succeeding segments. Terminal segment of P5
almost square, terminal spine swollen near its base, length about half that of outer seta.
Male. Length: 0.9-1.1 mm (0.91-1.3 mm). (Fig. 213 c, d; Fig. 215 a, b)

Posterior borders of Pd5 with row of 4-5 tiny spinules on dorsoposterior margin, as in the female.
A1 extends to the posterior border of Ur2. Ur1 with lateral tufts of hair, Ur2 with dorsolateral
rows of tiny spinules arranged across and on the posterior edge of the somite, Ur3 and Ur4 also
have tiny spinules on posterior dorsal edge. Genital somite with two pairs of lateral tufts of hair
on anterior part. No spines on this or succeeding segments. Terminal segment of left P5 almost
square, with thick naked terminal spine, one modified spine and one seta on the midanterior
surface. Terminal spine swollen near its base, length about half that of outer seta. The inner
margin has a row of hairs.
Fig. 213. Acartia (Acanthacartia) fossae.

Female: a. Pd5 and Ur, dorsal; b. P5. Male: c. Pd5 and Ur, dorsal; d. P5.
(Source: Nishida, 1985).

Fig. 214. Acartia (Acanthacartia) fossae. Female.
a. dorsal view; b. lateral view.
a b
Fig. 215. Acartia (Acanthacartia) fossae. Male.

a b

Subgenus Odontacartia Steuer, 1915
Reference: Steuer, 1915 (cited by Bradford-Grieve, 1999).
Rostral filaments present. Prosome posterior borders drawn into lateral spines, likewise the
posterior borders of urosome somites, especially in male (with the exception of A. lillieborgi).
Key to Acartia subgenus Odontacartia species

Females
A1 with 2 spines on segment 1; caudal rami short; P5 B elongate, 3 times longer than
broad; exopod in the form of curved spine, with double inflation at its proximal part
…………………………………………………………….… ………………… A. (O.) amboinensis
A1 without spines on segment 1; caudal rami elongated; P5 B short, exopod in the form of a spine
with proximal knob …………………………………………………………… A. (O.) ohtsukai
Males
Ur3 and 4 without posterodorsal spines; P5 right Exp1 with small inner process; left Exp1 with
distal outer spine ………………………………………………………….….. A. (O.) amboinensis
Ur3 with 2 large posterodorsal spines, Ur4 with a pair of smaller spines; P5 right Exp1 without
inner process, left Exp2 with long inner seta ………………………………... A. (O.) ohtsukai
Acartia (Odontacartia) amboinensis Carl, 1907
Female. Length: 1.44-1.47 mm. (Fig. 216 A – G; Fig. 218 a, b)

Prosome posterodorsal corners extend laterally into large spines, with 2 smaller spines dorsally.
Posterodorsal border of Genital somite with 2 small spines. Caudal rami short. Rostrum present
as 2 filaments. A1 with 2 spines on segment 1. P5 B elongate, 3 times longer than broad; exopod
in the form of curved spine, with double inflation at its proximal part.
Male. Length: 1.29-1.33 mm. (Fig. 217 H – J; Fig. 219)

Prosome posterior borders extend laterally into large spines, with 2 pairs of smaller spines
dorsally. Ur1 naked; Ur2 posterolaterally with 2 spines on both sides, these spines evident from
dorsal view; Ur5 with long hairs on lateral margins. P5: right Exp1 with small inner process at
distal part, Exp2 with large inner process; left Exp1 with distal outer spine.

Fig. 216. Acartia (Odontacartia) amboinensis. Female.
A. dorsal view; B. right lateral view; C. anterior part
of head and A1 (dorsal view); D. Ur and last thoracic
segment (right lateral view); E. Ur and P5 (ventral
view); F. Ur and last thoracic segment (dorsal view);
G. P5.
Fig. 217. Acartia (Odontacartia) amboinensis. Male.

H. Ur and last thoracic somite (dorsal view);
I. Ur (right lateral view); J. P5.
a b
Fig. 218. Acartia (Odontacartia) amboinensis. Female. Fig. 219. Acartia (Odontacartia) amboinensis. Male,
a. dorsal view; b. left lateral view. dorsal view.

Acartia (Odontacartia) ohtsukai Ueda and Bucklin, 2006
Acartia (Odontacartia) ohtsukai is a new brackish-water calanoid copepod, registered first in

Ariake Bay, Japan. This species is closely related to A. pacifica Steuer, 1915 (Ueda and Bucklin,
2006).
Female. Length 1.44-1.51 mm. (Fig. 220 A – E, F, J, K; Fig. 221 A – C; Fig. 223 a)
Prosome posterior borders extend laterally into large spines, with 2 smaller spines dorsally.
Posterodorsal border of genital somite with 2 small spines, Ur2 with 2 larger posterodorsal
spines. Caudal rami more than twice longer than broad. Rostrum present as 2 filaments. P5 B
short, exopod in the form of a spine with a knob-like process at its proximal part.
Male. Length 1.26-1.33 mm. (Fig. 220 G, H; Fig. 222 A – C; Fig. 223 b)
Prosome posterior borders extend laterally into small spines, with 2 smaller spines dorsally. Ur1
naked; Ur2 extends posterodorsally into points, with a pair of posterodorsal spines; Ur3 with
2 large posterodorsal spines; Ur4 with a pair of smaller spines more toward midline. P5: right
Exp1 without inner process, Exp2 with large inner process; left Exp2 with long seta on mid inner
margin.
Fig. 220. Acartia (Odontacartia) ohtsukai.

Female: A. dorsal view; B. left lateral view; C. last thoracic somite and Ur, dorsal view; D. Ur, left lateral view; E. R,
ventral; F. P5; J and K. last thoracic somite and Ur, dorsal view (abnormal). Male: G. dorsal view; H. last thoracic
somite and Ur, dorsal view; I. P5.
(Source: Al-Yamani & Prusova, 2003, where this species was previously identified as A. pacifica).

Fig 221. Acartia (Odontacartia) ohtsukai. Female.
a. dorsal view; b. left lateral view; c. Ur dorsal view.
a c b
Fig 222. Acartia (Odontacartia) ohtsukai. Male.

a. dorsal view; b. left lateral view; c. Ur dorsal view.
a c b
Fig 223. Acartia (Odontacartia) ohtsukai.

a. Female Ur lateral view; b. Male Ur lateral view.
a b

Acartiella faoensis Khalaf, 1991
Female. Length:1.0-1.28 mm (0.9-1.15). (Fig. 224 a – c)

The prosome is about 2 times as long as the urosome, proportional length of prosome and
urosome 2.1:1. The cephalosome and the first pedigerous somite are separate, pedigerous
segments 4 and 5 are fused together. The urosome is long and consists of 3 separate somites
and terminate in 2 long caudal rami, which are asymmetrical in both sexes. Urosome somites 1-3
are fused. All urosome somites are devoid of spines. Antennule when folded back reaches to the
middle of the caudal rami. The basipod segment 1 of leg 5 fused medially, segment 2 twice as
long as wide with a single long plumose seta inserted at about 0.8 the distance along the lateral
margin, the outer marginal seta of the same length as exopod. Both the exopod and endopod
are present, the exopod with a sharp point and bearing a single seta-like spine on its external
margin at about the junction of the middle and distal thirds.
Male. Length 0.95-1.2 mm (0.9-1.0 mm). (Fig. 225 a – c)

Prosome is about 2 times as long as urosome. Proportional length of prosome and urosome 1.9-
1.0. The urosome consists of 5 somites. The A1 on the right side is modified to form a grasping
organ. It consists of 17 separate segments.

a b c
Fig. 224. Acartiella faoensis. Female. a. dorsal view; b. lateral view; c. Ur dorsal.
a b c
Fig. 225. Acartiella faoensis. Male. a. dorsal view; b – lateral view; c. P5.

Family Tortanidae Sars, 1902
Description: As in the description of the only genus (see below).
Genus Tortanus Giesbrecht, 1898

References: Brodsky, 1950; Bradford-Grieve, 1999.
Females
Small copepods with one median eye. Rostrum absent. Cephalosome and Pd1 separate, Pd4 and
5 fused or separate; last thoracic segment posterior corners with rounded or pointed extremites.
Urosome of 2 or 3 somites, genital somite without seminal receptacles, often asymmetrical, both
because of asymmetry in individual segments and as a result of curvature along the longitudinal
axis. Caudal rami sometimes asymmetrical, one larger than the other, often fused to the anal
segment. A1 12- to 15-segmented; A2 Exp indistinctly 3-segmented, Enp 2-segmented. Mx1
with much reduced number of lobes (Li1 and one other the only ones present). Mx2 with lobes
1-3 reduced, the remaining lobes bear claw-like setae. Mxp reduced, with 2 long spines on first
segment; the distal part of limb 2-segmented, bearing 3 or 4 setae on inner margin and 1 on
outer margin of distal segment. Swimming legs with 3-segmented Exp; Enp of P1 2-3-segmented,
P2-4 2-segmented. P5 simple, uniramous, 2-3-segmented; coxa and intercoxal plate fused; basis
with outer seta; 1 free exopod segment present, sometimes in form of curved tapering process,
typically with 1 outer margin spine and 3 spinous processes arranged along the margins.
Males
Last thoracic segment posterior corners with rounded or pointed extremites or without
protrusions. Urosome of 5 somites, often asymmetrical. Right A1 prehensile, its middle part
widened and sometimes equipped with a denticulate plate. P5 asymmetrical, uniramous; right
leg 3-segmented, third segment forming a subchela.
Tortanus barbatus (Brady, 1883)
Female. Length: 1.15-1.5 mm (1.32-1.6 mm). (Fig. 226 a – f; Fig. 227 a – c)

Urosome of 3 somites; anal somite and caudal rami fused, asymmetrical; Ur3 in lateral view with
dorsal projection. P5 asymmetrical, terminal segment on one side short and tapering to a point, on
the other side elongated and with an inner median serrated margin, the teeth of which are variable
in appearance.
Male. Length: 0.9-1.2 mm (1.02-1.12 mm). (Fig. 226 g – j; Fig. 228 a – c)

Anal somite and caudal rami fused, slightly asymmetrical. Right P5 second segment with a triangular
projection with a seta halfway along the inner margin; left P5 penultimate segment with no setae,
terminal segment with a long outer seta, with numerous spinules on the inner and outer distal edges.

Fig. 226. Tortanus barbatus.
Female: a. postrior metasome and urosome, dorsal;
b. P5; c. aberrant left P5 with bifid denticle;
d. posterior metasome and urosome with broad-
caudal rami form (other specimen); e. P5 of same;
f. posterior metasome and urosome of broad-caudal
rami form, left lateral view.
Male: g. Ur, dorsal; h. P5, anterior view; i. P5, posterior
view; j. Pd4-5 and Ur, right lateral view.
(Source: Greenwood, 1978).
a b
Fig. 227. Tortanus barbatus. Female. a. dorsal view; b. lateral view; c. P5.
a b c
Fig. 228. Tortanus barbatus. Male. a. dorsal view; b. lateral view; c. P5.

Tortanus forcipatus (Giesbrecht, 1889)
Female. Length: 1.1-1.6 mm (1.09-2.0 mm). (Fig. 229 A – C; Fig. 230 a, b)

Urosome of 3 somites, anal somite and caudal rami fused, asymmetrical. In lateral view Ur3 has
a large irregular notch. P5 asymmetrical, terminal segment on right around half the length of
left. Left P5 terminal segment with 1 small and 2 larger spines on outer margin; right P5 with 2
small spines on mid outer margin.
Male. Length: 0.95-1.2 mm (0.94-1.17 mm). (Fig. 229 D, E; Fig. 231 a, b)

Genital somite and Ur2 free, the remaining part of Ur and caudal rami fused, slightly asymmetrical.
Right P5 second segment with a pointed triangular projection; left P5 terminal segment curved.
P5 left leg segment 3 naked; segment 4 with 1 distal outer seta, 1 inner seta and fine setules
along its inner border, ending with small short spine.
Fig. 229. Tortanus forcipatus.

Female. A. dorsal view; B. left lateral view;
C. P5. Male: D. dorsal view; E. P5.

b a
Fig. 230. Tortanus forcipatus. Female. a. dorsal view; b. Ur left lateral view.
a b
Fig. 231. Tortanus forcipatus. Male. a. dorsal view; b. right lateral view.

Order Cyclopoida Burmeister, 1834
The Cyclopoida are defined by their podoplean tagmosis, by the possession of a combination of
digeniculate A1 bearing a sheath on segment 15 in males with the lack of a defined A2 Exp. The
A2 Exp is represented by up to 3 setae, but no Exp segment remains. They are also characterized
by the fusion of Enp1 and Enp2 of Mx2 and by a 4-segmented Md Exp derived by loss of the
ancestral Exp1 (Huys and Boxshall, 1991; cited by Bradford-Grieve et al., 1999).
Family Oithonidae Dana, 1853

Reference: Nishida, 1985 (cited by Bradford-Grieve et. al., 1999).
Females
Total length: 0.36-1.90 mm. Prosome short oval to long fusiform in dorsal view, of 5 somites.
Anterior end of head quadrate, rounded or produced into pointed rostrum. Urosome slender,
of 5 somites; genital somite longest. Caudal rami frequently with 1 dorsal, 4 apical and 1 outer
marginal setae; 2 of apical setae may be absent in Paroithona. A1 slender, with proximal part
indistinctly segmented making it difficult to determine precise number of segments. A2
uniramous, bent outward at medial geniculation; proximal part 1-segmented, with 1-3 inner and
0-2 outer marginal setae; distal part 1- or 2-segmented. Proximal segment on distal part short,
with 2 medial and 3 distal inner marginal setae; distal segment long, usually with 7 terminal
setae. Md consisting of gnatobase with 1-2 transverse rows of setules on anterior surface near
distal margin. B transversally elongated, with 1 inner seta; fused to Enp1 which has 2-3 elements
varying in shape and size; Enp2 with 2-5 setae; Exp 4-segmented, with 5 setae. Mxp 4-segmented;
praecoxa and coxa incompletely separated; seta formula: 1-3-2 (1-2-2 in Limnoithona); basis with
1 medial and 1 distal inner marginal setae; 1 midmarginal row and/or 1 distal posterior surface
row of minute setae psesent or absent; Enp1 with 1 or 3 setae; Enp2 with 3-4 setae. P1-4 usually
with 3-segmented Endopod and exopod except for Paroithona where P1 Exp2 and 3 and P1-4
Enp2 and 3 are partly or completely fused. In most species P4 Enp2-3 with 1-2 modified setae
which are thick and slightly curved towards tip, with distal flange. P5 represented by 2 lateral
processes on Ur1; its dorsal process usually a free segment with 1-4 terminal setae. P6 reduced
into lateral process with 1-2 setae on anterior part of genital somite.
Males
Total length: 0.37-1.24 mm; usually smaller than female. Prosome longer than urosome, of
5 somites. Forehead rounded in lateral view, truncate in dorsal view. Lateral posterior margin
of cephalosome usually produced into flap with well-developed rows of pores. Urosome of 6
somites excluding caudal rami. A1 with 2 geniculations. A2, Md, Mxp similar to those of female in
basic structure however A2, Md, and P6 sexually dimorphic in some species. P1-4 Exp generally
with 1, 1, 2-3 outer marginal spines. P5 as in female.
Subfamily Oithoninae Kiefer, 1928

Genus Oithona Baird, 1843 (Figs. 232, 233)
References: Nishida, 1985; Shuvalov, 1980.
Females
Small subtle copepods, total length 0.4 – 1.9 mm. Prosome short oval to long fusiform in dorsal
view, of 5 somites. Anterior end of head rounded or produced into 1 pointed rostrum, diverse
in shape, may serve as a character for the species identification. Pd4 posterior margin rounded.
Urosome slender, of 5 somites, excluding caudal rami; genital somite longest. A2 uniramous.
Md B2 with 2 spines on distal end, outer one reduced in some species. P1-4 Enp 3-segmented.
P5 represented by 2 lateral processes on Pd 5. P6 reduced into lateral process with 1-2 setae on
anterior part of genital somite.

a b
c d
Fig. 233. Swimming legs (a, b, d) and exopods of swimming legs (c) of Oithona species: a. Oithona attenuata;
b. Oithona brevicornis; c. Oithona plumifera; d. Oithona nana. E. P1; F. P2; G. P3; H. P4.
(Sources: a, b, c from Al-Yamani and Prusova, 2003; d- from Nishida, 1985).
2 3
1 4 5
Fig. 232. Body shape of Oithona species (schematic figures):

1. O. plumifera; 2. O. attenuata; 3. O. nana; 4. O. brevicornis; 5. O. simplex. 1-5.
(Source: Shuvalov, 1980).

Males
Prosome longer than urosome, short or long roundish rectangular in dorsal view, of 5 somites.
Anterior end of head truncate in dorsal view, rounded in lateral view, newer ending in sharply
pointed rostrum. Pd4 posterior margin rounded. Urosome slender, of 6 somites, excluding caudal
rami. A1 with 2 geniculations. A2, Md, P5 similar to those of female in basic structure.
Keys to Oithona species

Females
1. Forehead produced into pointed rostrum………………....……………….……………………… 2
Forehead not produced into pointed rostrum………………....……………….………………….. 3
2. Forehead rounded in dorsal view. In lateral view, rostrum wide, curved
downwards………………………………………........…………………….... Oithona brevicornis(?)
Forehead narrowing anteriorly in dorsal view. In lateral view bent anteroventrally
into sharply pointed rostrum. Anteroventral surface of genital somite with tuft of fine
setae………………………………………........……………………..................Oithona plumifera
Forehead narrow, truncate anteriorly in dorsal view. Genital somite remarkably swollen laterally
in anterior 1/4, anterior margin of swelling nearly semicircular in
dorsal view………………………………………........…………………….......Oithona nana
2. P4 Exp3 with 2 outer marginal spines; A2 B1 with moderately thick long seta on distal
end of outer margin………………………………………........………….......Oithona attenuata
3. P4 Exp3 with 3 outer marginal spines; A2 B1-2 fused, with small outer midmarginal
seta ………………………………………........……………………..................Oithona sp.
Males
1. P4 Exp3 with 2 outer marginal spines. A2 distal segment long, its length about 4 times
width………………………………………........……………………................Oithona attenuata
2. P4 Exp3 with 3 outer marginal spines. A2 distal segment short, its length about 2 times
width………………………………………........…………………….................Oithona sp.
Oithona attenuata Farran, 1913
Female. Length: 0.6-0.8 mm (0.5-0.9 mm). (Fig. 234 A – I; Fig. 236 a, b)

Prosome rounded rhomboid, greatest width at posterior end of cephalosome; length 1.5 times
width, 1.0 times urosome. Head in lateral view bent ventrally into small process. Flap of caudal rami
lateral seta not extending to posterior margin of caudal rami. A1 short; length 0.95 Pr. A2 B1 with 1
moderately thick seta on distal end of outer margin. Distal segment long rectangular. Md B2 with 1
strong clawlike spine with spinules and 1 thin short seta on distal end, and 1 seta about middle of
inner margin. Enp with 4 setae. Genital somite with a pair of lateral processes located 0.33 times the
genital somite length from anterior end. P1-4 Exp with 1.1.3; 1.1.3; 1.1.3; 1.1.2 outer marginal spines.
Proportional lengths of terminal spine to Exp 1.0, 0.8, 1.0, 0.9. P5 with 1 seta attaching directly to
dorsolateral process of last thoracic segment, and 1 long free segment with 1 long seta.
Male. Length: 0.5-0.6 mm (0.4-0.53 mm). (Fig. 235 J – M; Fig. 237)

Prosome ovoid; length 1.9 times width, 1.2 times urosome. Head in lateral view bent ventrally into
small process. A2, Md B2, P1-5 as in female. P6 with 2 subequal short setae, extending to posterior
end of Ur3, attaching directly to small ventrolateral process on posterior margin of genital somite.

Remarks: Present specimens agree with Nishida’s (1985) description of O. attenuata stocky form.
According to Nishida (1985), O. attenuata typical form is of bigger size (total length 0.78-0.84 mm),
bigger prosome length to width ratio (2.0) and smaller prosome to urosome length ratio (0.8) in
comparison with the stocky form.
Fig. 234. Oithona attenuata. Female. Fig. 235. Oithona attenuata. Male.
A. dorsal view; B. left lateral view; C. Ur1-2 dorsal J. dorsal view; K. Ur1-3 ventral view; L. A2; M. Mdp.
view; D. A2; E. P1; F. P2; G. P3; H. P4; I. Mdp.
a b
Fig. 236. Oithona attenuata. Female. a. dorsal view; Fig. 237. Oithona attenuata. Male, dorsal view.
b. lateral view.

Oithona brevicornis (?) Giesbrecht, 1891
Female. Length: 0.5-0.75 mm (0.58-0.72 mm). (Fig. 238 A – H; Fig. 239 a, b)

Prosome elongate ellipsoid in dorsal view; length 2 times width, 1.3 times urosome. Head rounded
anteriorly in dorsal view; in lateral view produced ventrally into sharply pointed, slightly curved
downwards rostrum. Caudal rami lateral seta extending beyond posterior margin of caudal rami.
Lateral seta and 4 apical setae plumose; outer marginal seta naked. A1 short; length 0.75 times
prosome. A2 B1-2 fused, with 1 moderately long outer seta. Distal segment long rectangular. Md
B2 with 2 thick, blunt curved spines with spinules on distal end. Enp with 4 setae. P1-4 Exp with
1.1.3; 1.1.3; 1.1.3; 1.1.2 outer marginal spines. Proportional lengths of terminal spine 1.0, 0.8, 0.8
and 0.9. P5 with 1 seta attached directly to lateral process of last thoracic segment, and 1 long
free segment with 1 long, plumose seta.
Remarks: Present specimens agree with O. brevicornis Giesbrecht, 1891, but differ from the
latter in proportions of prosome length and width, prosome and urosome length (according
to Nishida’s (1985) description of O. brevicornis, these proportions are 2.4-2.6 and 1.4-1.5
correspondingly) and in caudal rami lateral seta being plumose and extending beyond posterior
margin of caudal rami (according to Nishida (1985), in O. brevicornis, flap of lateral setae not
extending to posterior margin of caudal rami). In structure of lateral seta the present specimens
correspond to O. spinulosa Lindberg, 1950. However, Nishida (1985) considers O spinulosa
Lindberg, 1950 as junior synonym of O. brevicornis Giesbrecht, 1891, and the present specimens
are referred to as O. brevicornis Giesbrecht, 1981. Shuvalov (1980) believed that O. brevicornis s.l.
could be a complex of closely related species. Further study needed to clarify the final status of
the described specimens.
Oithona nana Giesbrecht, 1892 (= O. minuta Krichagin, 1873, non O. minuta T. Scott, 1894)
Female. Length: 0.48-0.6 mm (0.49-0.72 mm). (Fig. 240 A, K; Fig. 241 a, b)

Head narrow, truncate anteriorly in dorsal view. Rostrum blunt and not visible dorsally. Prosome
short and stout, approximately the same length as the urosome. Antennule short, approximately
as long as prosome. Swimming legs (P1-P4) exopod with 1.1.3; 1.13; 1.1.3; 1.1.2 outer marginal
spines. Genital somite remarkably swollen laterally in anterior 1/4, anterior margin of swelling
nearly semicircular in dorsal view; length 1.8-1.9 times width.
Male. Length: 0.45-0.5 mm (0.44-0.53 mm). (Fig. 240 L, M)

Prosome shorter and stouter than female. Genital somite longer than wide. A1 geniculate. Anal
segment as long as wide. Caudal rami twice as long as wide. (Conway et al., 2003).

a b
Fig. 238. Oithona brevicornis (?). Female. Fig. 239. Oithona brevicornis (?). Female.
A. dorsal view; B. R left lateral view; C. A2; D. Mdp; a. dorsal view; b. lateral view.
E. P1; F. P2; G. P3; H. P4.
a b
Fig. 240. Oithona nana. Fig. 241. Oithona nana. Female. a. dorsal view;
Female: A, C. dorsal view; B. lateral view; D, E. forehead, b. lateral view.
dorsal and lateral; F - I. swimming legs P1 - P4; J. Pd4-5
and Gns, dorsal; K. CR. (Source: Shuvalov, 1980). Male:
L. dorsal view; M. basipodal segment 2 of Md
(Source: Grice, 1960).

Oithona plumifera Baird, 1843
Female. Length: 1.0-1.3 mm (1.06-1.51). (Fig. 242 A – H; Fig. 243 a, b)

Prosome fusiform in dorsal view; length 2.4 times width, 1.05 times urosome. Head narrowing
anteriorly in dorsal view; in lateral view bent anteroventrally into sharply pointed rostrum.
Anterior 1/3 of Genital somite swollen laterally, anteroventral surface of genital somite with tuft
of fine setae. Caudal rami lateral seta extending beyond posterior margin of caudal rami. P1-4 Re
with 1.1.2, 1.0.2, 1.0.1, 0.0.1 outer marginal spines.
Male. Length: 0.75-0.8 mm (0.59-0.68 mm).

Anterior of cephalosome very different from female, as it is not pointed and the rostrum is
blunt. Antennule (A1) is twice geniculate, with a sheath just beyond the proximal elbow and a
semicircular process on the first segment beyond the distal elbow. Genital somite large, about
as wide as it long. The anal segment is similar in length to the two preceding segments (Conway
et al., 2003).
a b
Fig. 243. Oithona plumifera. Female. a. dorsal view;

b. lateral view.
Fig. 242. Oithona plumifera. Female.

A. dorsal view; B. Ur dorsal view; C. Gns left lateral
view; D. R left lateral view; E. P1 Exp; F. P2 Exp; G. P3
Exp; H. P4 Exp.
10 0 M A R I N E ZO O P L A N K TO N P R AC T I C A L G U I D E f or t h e N ort h w e s tern A ra b ian G ul f

Salmiya beach in early morning - Photo by Dr. V. Skryabin - Kuwait Institute for Scientific Research
K u wait I n s titute f or Scienti f ic R e s earc h 10 1

Oithona sp.
Female. Length 0.47.-0.54 mm. (Fig. 244 A – I; Fig. 246 a, b)

Prosome oval in dorsal view; length 1.7 times width, 1.6 times urosome. Head truncates anteriorly
in dorsal view; in lateral view bent ventrally into blunt rostrum slightly acute anteroventrally. Flap
of caudal rami lateral seta not extending to posterior margin of caudal rami. A1 short; length 0.7
times Pr. A2 B1-2 fused, with 1 inner seta on distal end and 1 short small outer seta. Md B2 with
1 strong spinulose spine on distal end, and 1 inner and 1 outer marginal setae. Enp small oblong
with 5 setae. P1-4 Exp with 1.1.3, 1.1.3, 1.1.3, 1.1.3 outer marginal spines. Proportional lengths of
terminal spine to Exp 0.5, 0.5, 0.6, 0.6. P5 with bifurcate lateral process with 1 seta on each branch.
Male. Length 0.50-0.53 mm. (Fig. 245 J – M; Fig. 247)

Prosome oval; length 1.8 times width, 1.5 times urosome. Head, broad truncated anteriorly in
dorsal view; in lateral view bent ventrally into blunt ostrum, a little acute anteroventrally. A2, Md,
P1-5 as in female.
Remarks: Present specimens, both females and males, are very similar to O. simplex Farran, 1913
in a body and rostrum shape, Md and P1-4 armature. Present specimens are distinguished from O.
simplex by their bigger size (according to Nishida,1985). O. simplex female length 0.36-0.40 mm,
male – 0.39-0.41 mm), proportional lengths of prosome and urosome (in O. simplex, according to
Nishida, 1985, Pr 1.7-1.9 times Ur in female and 1.7 times in male) and in A2 B1 with 1 short small
seta on outer margin both in female and in male.
Fig. 244. Oithona sp. Female.

A. dorsal view; B. Ur, dorsal view; C. anterior head, left lateral view; D. Mdp; E. A2; F. P1;
G. P2; H. P3; I – P4.

Fig 245. Oithona sp. Male.
J. dorsal view; K. lateral view; L. A2; M. Mdp.
Fig 246. Oithona sp. Female.

a b
Fig 247. Oithona sp. Male, dorsal view.

Family Oncaeidae Giesbrecht, 1892
Reference: Heron and Bradford-Grieve (1995).
Females
Prosome elongate to elongate-oval, or cyclopiform. Cephalosome and Pd1 always separate.
Urosome generally slender, of 5 somites; genital apertures paired, located dorsally or laterally on
anterior part of genital somite; seminal receptacles paired; caudal rami with 6 setae, dorsal seta
may or may not be inserted into dorsal expansion on ramus. Eggs carried in paired sacs.
Rostrum fused to cephalosome; nauplius eye present; labrum well-developed, medially incised.
A1 short 6-7-segmented. A2 uniramous, 3-segmented, subprehensile to prehensile; with
2-segmented Enp, terminal part with 6-7 setae or spines; exopod lacking; coxa and basis fused.
P1-4 biramous with both rami 3-segmented. P5 absent or uniramous, with single segmented
protopod (C+B); inner coxal seta absent; outer seta on basis may be present; exopod 1-segmented
bearing 1-4 setae; endopod absent. P6 represented by opercular plates bearing 1-2 setae or
spines closing off genital openings.
Males
Urosome of 6 somites, genital apertures paired, located dorsolaterally at posterior border of large
genital somite. A1 never geniculate, last 3 segments often fused; 3-6-segmented. P6 represented
by paired flaps at posterior of genital somite.
Genus Oncaea Philippi, 1843

Reference: Heron and Bradford-Grieve (1995).
Females
Prosome cyclopiform. Cephalosome and Pd1 always separate. Prosome and urosome divisions
well-defined, the latter generally slender. Prosome and urosome of 5 somites; genital apertures
paired, located dorsally or laterally on anterior part of genital somite; seminal receptacles paired;
caudal rami with 6 setae, dorsal seta may or may not be inserted into dorsal expansion on ramus.
Eggs carried in paired sacs. Rostrum fused to cephalon; nauplius eye present; labrum well-
developed, medially incised. A1 short 6-7-segmented. A2 uniramous, 3-segmented, with terminal
segment’s length shorter than that of first segment; exopod lacking. P1-4 biramous with both
rami 3-segmented; P2-P4 Enp of some species terminated with a conical process between two
apical spines. P5 small, rod- or knob-shaped, free or unsegmented. P6 represented by opercular
plates bearing 1-2 setae or spines closing off genital openings.
Males
Urosome of 6 somites, genital apertures paired, located dorsolaterally at posterior border of
large genital somite. A1 never geniculate, last 3 segments often fused; 3-6-segmented.

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

Oncaea clevei Fruchtl, 1923
Female. Length: 0.65-0.75 mm (0.68-76 mm). (Fig. 248 A – G; Fig. 250 a, b)

In lateral view pedigerous somite 2 with conspicuous dorsoposterior projection; pedigerous
somite 4 with rounded lateroposterior corners. Prosome 1.8 urosome. Genital somite 1.2-1.4
times width, at the widest part and longer than total length of somites which follow. Caudal rami
length about 3 times width and approximately equal to sum of 2 preceding somites. Endopod
segment of leg 4 without conical process between terminal and subterminal spines.
Leg 5 with small free segment, 2 terminal setae of approximately equal length.
Male. Length 0.55-0.7 mm (0.46 mm). (Fig. 249 H – M; Fig. 251 a, b)

Prosome 1.7 times of urosome. Caudal rami 1.1-1.2 times anal somite.
Family Sapphirinidae Thorell, 1859

Females
Body more or less flattened, often leaf-like. Two ocular lenses present on anterior cephalosome.
Cephalosome and Pd1 fused (Copilia) or separate; Pd3 and 4 may be expanded laterally into
rounded lateral plates. Urosome may be slender or tapering gradually from width of posterior
prosome, of 3-5 somites (in Sapphirina fused Ur2 and 3 have obvious demarcation between
somites); Ur1 (Pd5) small or may be fused to Ur2; genital apertures paired, located laterally or
dorsolaterally on anterior part of genital somite; caudal rami leaf- shaped or in form of narrow,
elongated rod, with 4-5 setae. Eggs carried in paired sacs. A1 short with at most 6 segments.
A2 uniramous, 5-segmented, prehensile, terminated by claw. Md with 1 large terminal toothed
element usually with 1-2 small setae at its base. Mxp claw-like 3-segmented, basis with 2 inner
edge spines; Enp1 fused with Enp2 and in form of claw which bears 2 proximal setae. P1-3
biramous with both rami 3-segmented; P4 with 3-segmented Exp and Enp 1-segmented (Copilia),
2-segmented (Vettoria) and 3-segmented (Sapphirina). P5 1-segmented with 2 small terminal
setae and small basal seta on each side, or 1 spine and seta, or represented by 3 alone (Vettoria).
P6 represented by 3 small rounded appendages (Sapphirina), or singe seta (Vettoria).
Males
Body shape usually different from that of female, usually more leaf-shaped with no distinct break
in width between the prosome and urosome. Urosome of 6 somites, Ur1 and anal somite often
covered by adjacent urosome somites; genital apertures paired, located ventrolaterally near
posterior border of the genital somite; caudal rami leaf- or rod-shaped. A1 never geniculate.
A2 more slender than in female. Md may be reduced. Mxp final hook is longer than in female,
4-segmented. P2 Enp3 distal part with modified spines in Sapphirina. P6 represented by 3 setules
in Sapphirina.

Fig. 248. Oncaea clevei. Female. Fig. 249. Oncaea clevei. Male.
A. dorsal view; B. left lateral view; C. Ur, dorsal; H. dorsal view; I. left lateral view; J. Ur, dorsal;
D. Ur, left lateral view; E. P4 Enp; F. A2; G – Mxp. K. Ur, left lateral view; L. A2; M. Mxp.
a b a b
Fig. 250. Oncaea clevei. Female. a. dorsal view; Fig. 251. Oncaea clevei. Male. a. dorsal view; b.
b. lateral view. lateral view.

Genus Copilia Dana, 1849
References: Wilson, 1932; Mori, 1937; Bradford-Grieve et al. 1999.
Females
Body transparent and strongly depressed; cephalosome fused with Pd1, squarely truncated
anteriorly, with a pair of knob-like corneal lenses near the corners. Urosome of 4 somites;
genital somite being divided by a median groove; caudal rami much longer than urosome. A1
6-segmented. A2 4-segmented. P1-P3 Exp and Enp 3-segmented; P4 Exp 3-segmented, Enp
1-segmented. P5 uniramous, 1-segmented.
Males
Body shape different from that of female, more leaf-shaped with no distinct break in width
between the prosome and urosome., body segments widened and flattened; posterior margin
of Pd4 with a median knob. Cephalosome separated from Pd1, without corneal lenses. Urosome
of 5 somites; caudal rami rod-like and shorter than in female. A1 never geniculate.
Copilia mirabilis Dana, 1849
Female. Length 3.27-3.29 mm. (Fig. 252 A – E; Fig. 253)

Cephalosome quadrangular, widened posteriorly and as long as the rest of body excluding caudal
rami. Ocular lenses separate by distance of approximately 1.5 their length. Pd4 with median
dorsal spine pointed backward. Anal somite anterior and posterior ends, being the widest part
of the segment, 1.4-1.5 times wider than the narrowest, central part. Caudal rami 1.5 times longer
than urosome. A2 with stout apical claw, with numerous small and 1 large spine on coxa; large
spine on basis with 4-6 lateral spines. P4 endopod and exopod 1 of equal length.
Fig. 252. Copilia mirabilis. Female. Fig. 253. Copilia mirabilis. Female, dorsal view.
A. dorsal view; B. urosome (dorsal); C. urosome
(ventral); D. A2; E. P4.

Salmiya city - Photo by Melad M. Helani

Genus Sapphirina Thompson, 1829
References: Wilson, 1932; Mori, 1937; Bradford-Grieve et al. 1999.
Females
Entire body strongly depressed, prosome much widened, with a pair of cuticular lenses on
forehead. Prosome of 5 somites; cephalosome and Pd1 separated. Urosome of 5 somites, much
narrower than prosome; caudal rami leaf-like, with 4 short marginal setae. A1 3-6-segmented. A2
4-segmented, prehensile, terminal segment with a hook-like spine. P1-P4 endopod and exopod
3-segmented; P4 endopod often smaller than exopod. P5 rudimentary, 1-segmented, each with
2 minute apical setae.
Males
Urosome of 6 somites; Ur1 very small; Ur2 (genital somite) nearly as wide as last prosome somite;
anal somite small. Caudal rami leaf-shaped. A1 never geniculate. A2 more slender than in female.
P2 Enp3 distal part with modified spines.
Sapphirina nigromaculata Claus, 1863
Female. Length 1.71-2.10 mm. (Fig. 254 A – D; Fig. 255 a, b)

Cuticular lenses visible in dorsal view. Prosome 1.8 times urosome. Pd1-3 tapered regularly
backward. Caudal rami with 1 inner projection on distal border; length 2.3-2.5 times width. P4
endopod shorter than exopod.
Male. Length 1.93 mm. (Fig. 254 E – H; Fig. 256 a, b)

Caudal rami with 1 projection on inner distal border, length 1.8 times width. P2 Enp3 with 2
foliaceous spines and 1 spine with denticulate margin on the outer side only.
Fig. 254. Sapphirina nigromaculata. Female.

A. dorsal view; B. urosome (dorsal view);
C. A2; D. P4. Male: E. dorsal view;
F. caudal rami; G. A2; H. P2 Enp.

a b
Fig. 255. Sapphirina nigromaculata. Female. a. dorsal view; b. Ur, dorsal view.
a b
Fig. 256. Sapphirina nigromaculata. Male. a. dorsal view; b. Ur, dorsal view.

Family Corycaeidae Dana, 1852
Females
Body cylindrical to conical, tapering posteriorly. Ocular lenses close together, sometimes
contiguous. Cephalosome and Pd1 fused or separate; Pd3 and 4 prolonged into lateral points;
Pd2, 3 and 4 may be fused or separate. Urosome generally slender, of 2-3 somites, Ur1 (Pd5)
small; genital apertures paired, located dorsolaterally on anterior part of genital somite;
seminal receptacles paired; caudal rami elongate with 4 setae. Eggs carried in paired sacs.A1
short, 6-segmented with naked setae; without sensory organs. A2 uniramous, 5-segmented,
prehensile; coxa and basis large, each with thick, long seta; endopod with strong, hooked spines.
P1-3 biramous with both rami 3-segmented, exopod longer than endopod; P4 with 3-segmented
exopod; Enp1-segmented with 1 or 2 seta, or completely absent; P1 C with or without inner seta.
P5 in form of 2 small setae on each side. P6 represented by opercular plates closing off genital
openings.
Males
Urosome of 2-3 somites, genital apertures paired, located ventrolaterally near posterior border
of genital somite. A1 never geniculate. A2 terminal hook longer than in female. Mxp final hook
longer than in female, 4-segmented; basis with or without inner edge spine. P6 represented by 1
plumose seta on flaps closing off genital apertures.
Genus Corycaeus Dana, 1845

References: Tanaka, 1957; Bradford-Grieve et al. 1999.
Females
Body slender, subclavate; cephalosome much longer than the rest of prosome; ventral projection
on cephalon rounded. Ocular lenses large, placed close together. Pd3 and 4 prolonged into
lateral points. Urosome generally slender, of 2 somites (except females of s.g. Agetus) genital
apertures paired, located dorsolaterally on anterior part of genital somite; seminal receptacles
paired; caudal rami elongate with 4 setae. A1 short, 6-segmented. A2 uniramous, 5-segmented,
prehensile; coxa and basis large, each with thick, long nude seta; Endopod with strong, hooked
spines. P1-3 biramous with both rami 3-segmented, Exopod longer than endopod; Exp P1-3 with
1, 1, 3 spines; P4 (besides basal seta) with small, peg-like endopod with 1-2 setae. P5 in form of 2
small setae on each side. P6 represented by opercular plates closing off genital openings.
Males
Urosome generally of 2 somites. A2 with 2 proximal setae;1 seta finely plumose along one edge;
terminal spine elongate. Ventral projection on cephalosome rounded.

Subgenus Dithrichocorycaeus M. Dahl, 1912
Females and males
Reference: Tanaka, 1957.
Urosome of 2 somites; caudal rami not longer than remainder of urosome; peg-like endopod of
P4 with 2 setae and small posterior projection.
Key to Corycaeus subgenus Dithrichocorycaeus species

Females
1. Prosome at least twice longer than urosome; caudal rami shorter than anal somite .………2
Prosome less than twice as long as urosome; caudal rami longer than anal somite .…...…..3
2. Genital somite massive, widest at its anterior end; A2 B2 marginal spine nearly 0.6 times as
long as that of B1 …………………………………………………………….…C. (D.) andrewsi
3. Prosome 1.7 times urosome; genital somite with small ventral hook …….…...C. (D.) dahli
Prosome 1.7 times urosome; genital somite without ventral hook …………...C. (D.) lubbocki
Corycaeus (Dithrichocorycaeus) andrewsi Farran, 1911
Female. Length 0.73-0.76 mm. (Fig. 257 A – E; Fig. 258 a, b)

Prosome 2.3 – 2.4 times urosome. Pd3 distal corners produced into small wing-like expansions.
Pd4 distal corners not extending to the middle of genital somite. Genital somite 1.1 times the
reminder of urosome, without ventral hook in lateral view. Urosome somites and caudal rami in
the proportional lengths 24:13:12. A2 B2 marginal spine 0.6 times as long as that of B1.
Remarks: The described specimens slightly differ from Tanaka’s (1957) and Dahl (1912) C. (D.)
andrewsi by their smaller size. The C. (D.) andrewsi female length according to Tanaka, 1957 is
1.00-1.07 mm, and is 0.88-0.95 mm according to Dahl (1912); and Pr/Ur proportions according
to both Dahl (1912) and Tanaka (1957) who indicated that prosome to be 2 times the urosome.
a b
Fig. 258. Corycaeus (Dithrichocorycaeus) andrewsi. Female.

a. dorsal view; b. left lateral view.
Fig. 257. Corycaeus (Dithrichocorycaeus) andrewsi. Female.

A. dorsal view; B. right lateral view; C. A2; D. P4; E. P2 Exp3.

Corycaeus (Dithrichocorycaeus) dahli Tanaka, 1957
Female. Length 0.97-1.10 mm. (Fig. 259 A – F; Fig. 261 a – c)

Prosome about 1.7 times as long as urosome. Urosome somites and caudal rami in the
proportional lengths 30:18:38. Posterior corners of Pd3 extend beyond the middle of genital
somite. Pd4 produced into short pointed processes. Genital somite ventral surface flat, with very
small hook on anterior end. A2 B2 marginal spine about 3 times shorter than that of B1.
Male. Length 0.88-0.93 mm. (Fig. 260 G – J; Fig. 262 a – c)

proportional lengths 28:11:17. Genital somite oval in dorsal view; in lateral view ventral surface
with a small median hook. A2 B1 marginal spine a little longer than that of B2; terminal spine
about 1.4 times as long as B2.
Fig. 259. Corycaeus (Dithrichocorycaeus) dahli. Female.

A. dorsal view; B. left lateral view; C. Ur, dorsal; D. Ur, right lateral view; E. A2; F. P4.

Fig. 260. Corycaeus (Dithrichocorycaeus) dahli. Male.
G. dorsal view; H. right lateral view; I. Ur, right lateral
view; J. A2.
Fig. 261. Corycaeus (Dithrichocorycaeus) dahli. Female.

a. dorsal view; b. left lateral view; c. Ur, left lateral view.
a b c
Fig. 262. Corycaeus (Dithrichocorycaeus) dahli. Male.

a. dorsal view; b. right lateral view; c. Ur, right lateral
a b c view.

Corycaeus (Dithrichocorycaeus) lubbocki Giesbrecht, 1891

proportional lengths 38:17:47. Posterior corners of Pd3 extend to the middle of genital somite.
Pd4 produced into short pointed processes. Genital somite ventral surface flat, without a hook.
A2 B2 marginal spine about 3 times shorter than that of B1.
rter than that of B1.
Subgenus Onychocorycaeus M. Dahl, 1912

Reference: Tanaka, 1957
Females and males

Small forms at most 1.18 mm. Females with very wide cephalosome; peg-like endopod of P4 with
1 seta, without posterior projection.
Key to Corycaeus subgenus Onychocorycaeus species

Females
Urosome slender; genital somite relatively small, shorter than caudal rami…… C. (O.) agilis
Urosome robust; caudal rami and anal somite of approximately equal length; Pd3 in its widest part
wider than cephalosome in dorsal view …………………………………....…… C. (O.) pacificus
a b
Fig. 264. Corycaeus (Dithrichocorycaeus) lubbocki. Female.

Fig. 263. Corycaeus (Dithrichocorycaeus) lubbocki. Female.

A. dorsal view; B. right lateral view; C. Ur, dorsal view; D. Ur, left
lateral view; E. A2; F. P4.

Corycaeus (Onychocorycaeus) agilis Dana, 1849
Female. Length 0.97 mm. (Fig. 265 A – F; Fig. 266 a, b)

Prosome robust, 1.5 times as long as urosome. Urosome slender. Urosome somites and caudal
rami in the proportional lengths 34:29:39. A2 B1 marginal spine 2.4 times longer than that of B2.
a b
Fig. 266. Corycaeus (Onychocorycaeus) agilis. Female.

Fig. 265. Corycaeus (Onychocorycaeus) agilis. Female.

A. dorsal view; B. left lateral view; C. Ur, dorsal;
D. Ur, left lateral view; E. A2; F. P4.

Corycaeus (Onychocorycaeus) pacificus F. Dahl, 1894

Prosome robust, about 2 times as long as urosome. Pd3 very wide, posterior corners extend
beyond ¾ of genital somite. Pd4 posterior corners short and bluntly pointed. Urosome somites
and caudal rami in the proportional lengths 48:19:19. Genital somite length 1.3 times width. A2
B2 marginal spine about 3 times shorter that of B1.
a b
Fig. 268. Corycaeus (Onychocorycaeus) pacificus. Female.

Fig. 267. Corycaeus (Onychocorycaeus) pacificus. Female.

A. dorsal view; B. left lateral view; C. Ur, dorsal; D. Ur, left lateral
view; E. A2; F. P4.

Salmiya Marina - Photo by Melad M. Helani

Order Harpacticoida Sars, 1903
The Harpacticoida are defined by their podoplean tagmosis, the presence of short A1 with at most
9 segments in females and 14 in males, the fusion of the endopod and basis into baseoendopod
on P5 of both sexes of most species, those with a separate Enp having a maximum of 2 setae on
it (Huys, 1996), cited by Bradford-Grieve et al., 1999.
Family Ectinosomatidae Sars, 1903

Reference: Rose, 1933 (cited by Bradford-Grieve et al., 1999).
Body smooth, spindle-shaped; without distinct demarcation between prosome and urosome.
Rostrum prominent, fused to cephalosome. Prosome prolonged laterally by well-developed
epimeral plates, except for last somite. Female urosome of 4 somites; male urosome of 5 somites;
last somite small and more or less cleft posteriorly. Caudal rami divergent; its 2 median setae
elongate. A1 small. A2 with terminal spines denticulate; exopod 2-3-segmented. Rami of P1-4
3-segmented. P5 lamellar with strong spines. Ovisac simple.
Genus Microsetella Brady and Robertson, 1873

References: Wilson, 1932; Mori, 1937.
Females
Body slender and compressed laterally. Urosome as wide as prosome; Cephalosome and Pd1
fused. Urosome of 4 somites, the Genital somite being completely divided; caudal rami divergent;
its 2 median setae elongate. A1 5-segmented, with two elongate aesthetasks. A2 biramous, Exp
3-segmented. P1-P4 with 3-segmented exopod and endopod, endopod longer than exopod. P5
symmetrical, 2-segmented; distal segment imperfectly trilobate, with 3 setae, inner expansion of
basal segment with 2 setae.
Males
Body as in female, except urosome of 5 somites. A1 geniculate. P1-P5 as in female; P5 inner
expansion of basal segment rudimentary.
Microsetella sp.
Female. Length 0.65-0.72 mm. (Fig. 269 A – H; Fig. 270 a, b)

The last 3 urosomal somites with transverse rows of spinules, caudal rami outer apical seta short,
the inner seta 1.2-1.4 times longer than the body length. P1 Enp1 inner distal seta reaching the
distal end of Enp2. P5 Enp (i.e., inner expansion of basal segment) well developed, with inner
apical seta nearly twice shorter than outer seta; Exp (i.e., distal segment) with 2 apical setae of
which the inner one longer, 1 relatively short inner lateral seta, and an accessory seta on posterior
surface at central part.
Male. Length 0.54-0.58 mm. (Fig. 269 I – L; Fig. 271 a, b)

Caudal rami inner seta1.3-1.5 times longer than the body length. P5 Exp with 2 apical setae of
which the inner one longer, and 1 accessory seta on posterior surface; basal outer seta 0.5 times
as long as Exp inner seta.
Remarks: The described species is very close to M. rosea (Dana, 1847), but differs from the latter
one in shorter caudal rami inner apical seta (in M. rosea caudal rami apical setae nearly twice the
body length) and in details of P1 and P5 armament (according to Giesbrecht, 1892, in M. rosea
female P1 Enp1 inner distal seta nearly twice longer than Enp2; P5 Enp apical setae about equal
in length; in male basal outer seta 0.3 times as long as Exp inner seta).

b a
Fig. 270. Microsetella sp. Female. a. dorsal view;

b. lateral view.
Fig. 269. Microsetella sp.

Female: A and B. left lateral view; C. Ur last somites, left
lateral view; D. P1, E. anterior head, left lateral view;
F. P5; G. A1; H. A2. Male: I. dorsal view; J. lateral view;
K. P5; L. A1.
a b
Fig. 271. Microsetella sp. Male. a. dorsal view; b. lateral view.

Family Miraciidae Dana, 1846
Reference: Wells, 1970 (cited by Bradford-Grieve et al., 1999).
Cephalosome and Pd1 fused. Caudal rami about as long as last 2 urosomal somites. P1 Exp
3-segmented, Enp 2-segmented. P2-4 with both rami 3-segmented. P2 Enp of male modified,
2-segmented. P5 basal segment expanded medially. Cephalosome usually with pair of large
cuticular lenses.
Genus Macrosetella A. Scott, 1909

Reference: Wilson, 1932.
Females
Body slender and cylindrical, slightly compressed laterally. Cephalosome and Pd1 fused. Urosome
of 4 somites, caudal rami slender, cylindrical, longer than last 2 urosome somites combined,
and closely appressed, apical setae much longer than the entire body. A1 9-segmented. A2 Exp
lacking. P1-P4 EXp 3-segmented, P1 Enp 2-segmented, P2-P4 Enp 3-segmented. P5 2-segmented.
Males
Body as in female except urosome of 5 somites. A1 geniculate. P1 and P2-P4 Exp as in female; P2
Enp 2-segmented, P3-P4 Enp 3-segmented. P5 2-segmented.
Macrosetella gracilis (Dana, 1847)
Female. Length 1.22 mm. (Fig. 272 A, B; Fig. 273 a, b)

A1 relatively long, reaching Pd4. P5 basal expansion not reaching central part of distal segment,
with 4 apical setae, the second inner seta plumose and the longest. Distal segment elongate,
with 3 outer edge setae and 3 apical setae, of which 2 inner setae equal in length.
Male. Length 1.01 mm. (Fig. 272 c, d)

A1 geniculate. P5 basal expansion very short and tipped with 2 setae; distal segment elongate,
with 3 apical and 1 outer setae.

Fig. 272. Macrosetella gracilis.
Female: A. right lateral view; B. P5. Male: C. lateral view; D. P5.
a b
Fig. 273. Macrosetella gracilis. Female. a. left lateral view; b. dorsal view.

Family Euterpinidae Brian, 1921
Body variable in shape. A1 short, prehensile in male. A2 Exp well-developed. Md with biramous
palp. P1 not prehensile, similar to P2; all with 3-segmented Exp and Enp. P5 lamelliform,
2-segmented. Ovisac simple.
Genus Euterpina Normann, 1903

Reference: Mori, 1937.
Females
Cephalosome and Pd1 fused. Prosome of 4 somites; rostrum uniramous and stout. Urosome of 6
somites. A1 7-segmented. A2 Exp well-developed, 3-segmented. Mdp Exp and Enp 1-segmented.
P1 Exp and Enp 2-segmented, P2-P4 Exp and Enp 3-segmented. P5 symmetrical, lamelliform,
1-segmented, with 4 apical setae.
Males
A1 geniculate on both sides, with segments 4 and 5 fused and much thickened; segments 6 and
7 fused. P5 symmetrical, rudimentary; each ramus furnished with 2 apical and 2 outer setae.
Euterpina acutifrons (Dana, 1848)
Female. Length 0.68-0.72 mm. (Fig. 274 A – E; Fig. 275 a, b)

Body fusiform. Caudal rami and anal somite of approximately equal length. P1 basis with teeth
along distal edge.
Family Clytemnestridae A. Scott, 1909

Prosome of 4 somites. Its posterolateral corners flattened and prolonged into points. Urosome
of 5 somites in female and of 6 somites in male. Caudal rami short, setae very short. A1
7-8-segmented, with short setae and long sensory organs; male A1 prehensile on both sides. A2
3-segmented; exopod rudimentary, represented by 1-2 setae. Mxp 2-segmented, elongate; with
terminal claw; male differs from female. P1 Enp 1-segmented, exopod 3-segmented. P2-4 with
both rami 3-segmented. P5 rudimentary, with 2 long segments; symmetrical or almost in male.
a b
Fig. 274. Euterpina acutifrons. Female. Fig. 275. Euterpina acutifrons. Female.
A. dorsal view; B. left lateral view; C. P1; D – Mxp; E.P5. a. dorsal view; b. lateral view.

Genus Clytemnestra Dana, 1848
Females
Prosome of 4 somites with posterolateral corners flattened and prolonged into points. Urosome
of 5 somites. Caudal rami short, setae very short. A1 7-8-segmented, with short setae and
long sensory organs; A2 3-segmented; exopod rudimentary, represented by 2 setae. Mxp
2-segmented, elongate; with terminal claw. P1 Enp 1-segmented, Exp 3-segmented. P2-4 with
both rami 3-segmented. P5 rudimentary, symmetrical, with 2 long segments.
Males
Urosome of 6 somites. A1 prehensile on both sides. P5 rudimentary, 2-segmented; almost
symmetrical.
Clytemnestra scutellata Dana, 1848
Female. Length 0.86-0.92 mm. (Fig. 276 A – G; Fig. 277 a – c)

Genital somite longer than the following 2 somites combined; caudal rami 0.7 times as long
as anal somite. A2 Exp in the form of 2 long setae, of nearly the same length as Enp2 and 3
combined. Mxp terminal claw reaches the middle of terminal segment. P5 proximal segment
with 1 outer distal seta; distal segment with 3 apical and 2 lateral setae.
a b
Fig. 277. Clytemnestra scutellata. Female.

a. dorsal view; b. lateral view; c. cephalon (dorsal
view).
Fig. 276. Clytemnestra scutellata. Female.
A. dorsal view; B. right lateral view; C. Ans and CR,
dorsal view; D. P5; E. P1; F. A2; G. Mxp.

2.14. Mysida
Subphylum Crustacea, Brunnich, 1772

Class Malacostraca, Latreille, 1802
Subclass Eumalacostraca, Grobben, 1892
Superorder Peracarida, Calman, 1904
Order Mysida, Haworth, 1825
Family Mysidae, Haworth, 1825
Subfamily Rhopalophthalminae, Hansen, 1910
Genus Rhopalophthalmus, Illig, 1906
Rhopalophthalmus sp. (Fig. 278 a – g)

References: Grabe, 1989; Grabe et al., 2004; Panampunnayil and Biju, 2006; Price, 1982.
Female (Fig. 278). TL=11.7 mm. General form slender (Fig. 278 a). Carapace with dorso-median
nodules (Fig. 278 g); anterior margin lacking rostrum, postorbital spines prominent, keels
prominent (Fig. 278 f ). Eyes stout, extending to distal end of first segment of antennule, cornea
occupying little more than half of eye and wider than stalk (Fig. 278, b). Apical spines of telson
are sub-equal (Fig. 278 c). Number of spines on lateral margin of telson – 14 (Fig. 278 d).
Distribution: Indo-Pacific Region; Arabian Gulf.

a
1 mm
f g
d e
Fig. 278. Mysid shrimp Rhopalophthalmus sp. Female. a. general view (scale bar 1mm);
b. anterior part of body; c. dorsal view of telson; d. lateral view of telson; e. dorsal view of telson (T) with uropods;
f. anterior margin of carapace; g. lateral fragment of carapace with nodules.
M. marsupium; OAS. apical spines of telson (outer spines); IAS. inner spines; LS. lateral spines of telson; ExUr. exopod
of uropods; EndUr. endopod of uropod.

2.15. Decapoda
Phylum Arthropoda
Subphylum Crustacea Brunnich, 1772
Class Malacostraca Latreille, 1802
Subclass Eumalacostraca Grobben, 1892
Superorder Eucarida Calman, 1904
Order Decapoda Latreille, 1802
Suborder Dendrobranchiata Bate, 1888
Superfamily Penaeoidea Rafinesque, 1815
Family Penaeidae Rafinesque, 1815
Genus Metapenaeus Wood-Mason, 1891
Metapenaeus ensis (De Haan, 1844) (Fig. 279 a, b)

Reference: Jackson et al.,1989.
Description: Protozoea II, TL = 2.5 mm; CL = 0.88 mm. Carapace does not completely cover
thorax (Fig. 279 a). Abdomen bifurcate posteriorly with each furca bearing at least seven spines.
No spines arise from posterior half of carapace. Eyes mobile (E in Fig. 279 a). Uropods absent. Two
pairs of supraorbital spines (Fig. 279 b). Inner pair of supraorbital spines filamentous and less
than half as long as outer pair.
Distribution: Indo-West Pacific Region; Arabian Gulf.
b
a
0.5 mm
Fig. 279. Metapenaeus ensis protozoea II. a. general dorsal view, b. anterior part of carapace.
E. eye; i1-i7 – 1st-7th telson spines.

Phylum Arthropoda
Genus Parapenaeopsis Alcock, 1901
Parapenaeopsis stylifera (H. Milne Edwards, 1837). (Fig. 280 a - d)

References: Habib-ul-Hassan, 1984; Jackson,C.J. et al.,1989.
Description: Protozoea III, TL = 1.6 mm; CL = 0.4 mm (Fig.280). Carapace with only rostral spine
round (Ca, Ro in Fig. 280 a, b). Stalked compound eyes. Biramous uropods present (Ex Ur, En Ur
in Fig. 280 c, d). Dorsal abdominal spines present (DSp in Fig. 280 d). Caudal furca wide (CF in Fig.
280 c, d). There are eight pairs of caudal spines (i1 - i8 in Fig. 280 c, d).
a b c
0.2 mm
Fig.280. Parapenaeopsis stylifera, protozoea III. a. dorsal view; b. dorsal view of carapace; c. dorsal view of telson;
d. lateral view of abdomen and telson.
Ro. rostrum; Ca. carapace; T. telson; CF. caudal furca; Ex Ur. exopod of uropod; En Ur. endopod of uropod; i1-i8.
1st-8th telson spines; DSp. dorsal spines; I-VI. abdominal somites.

Phylum Arthropoda
Genus Penaeus Fabricius, 1798
Penaeus semisulcatus de Haan, 1850 (Fig. 281 a - d)

References: Habib-ul-Hassan, 1982; Jackson et al., 1989.
Description: Protozoea I, TL = 1.1 mm, CL = 0.4 mm (Fig. 281).

Carapace is round (Fig.281 a, b) with a median notch at the anterior end (MN in Fig. 281 b).
The last six thoracic segements are free from the carapace (Fig. 281 c). Abdomen is unsegmented.
An ocellus is present (Fig. 281 a, b). The antennule is about the same size as the antenna (Fig. 281 a).
a b
0.2 mm
d
c
Fig. 281. Penaeus semisulcatus, protozoea I. a. general dorsal view; b. dorsal view of carapace; c. dorsal view of
thorax (Th, I-VI - thoracic segments) and abdomen; d. dorsal view of telson.
An1. antennules; An2. antenna; E. eye, O. ocellus; MN. median notch; Ab. abdomen; T. telson; CF. caudal furca;
i1-i7. 1st-7th telson spines.

Phylum Arthropoda
Genus Penaeus Fabricius, 1798
Penaeus semisulcatus de Haan, 1850 (Fig. 282 a - d)

References: Habib-ul-Hassan, 1982; Jackson et al., 1989.
Description: Protozoea III, TL = 2.3 mm, CL = 0.6 mm. There is a pair of prominent supraorbital
spines (SSp in Fig. 282 b) and projecting rostrum (R in Fig. 282 b). Tere is presance of biramous
uropods (Ur in Fig. 282 d) and dorsomedian spines (DSp in Fig. 282 c). There is one pair of lateral
spines on the fifth and sixth abdominal segments (LSp in Fig. 282 c, d). Caudal furca bears 8+8
spines (i1-i8 in Fig. 282 d).
a b
c d
0.5 mm
Fig. 282. Penaeus semisulcatus, protozoea III. a. dorsal view; b. dorsal view of frontal segments of carapace; c. dorsal
view of abdomen; d. dorsal view of distal part of abdomen (III-VI abdominal segments) and telson.
R. rostrum; SSp. supraorbital spines; DSp. dorsomedian spines; I-VI. abdominal segments; T. telson; LSp. lateral
spines; Ur. biramous uropods; i1-i8. telson spines.

Phylum Arthropoda
Family Solenoceridae Wood-Mason, 1891
Genus Solenocera Lucas 1850
Solenocera crassicornis (H. Milne Edwards, 1837) (Fig. 283)

References: Dos Santos, Lindley, 2001; Cheung, 1963; Farfante, Kensley, 1997.
Description: Protozoea III (Elaphocaris). TL = 2.8 mm; CL = 1.1 mm. Carapace with spines and
processes. Carapace margin serrate, spines robust (Fig. 283). Margin of carapace fringed with
5 pairs of lappets, serrated between 1st and 2nd and between 3rd and 4th (Lap in Fig. 283). Each
abdominal tergum with one dorsal and pair of lateral spines (LSp). Telson (T) bifurcate posteriorly,
each half with 7 setae (i1-i7).
Distribution: Arabian Gulf, along the northeast and east coasts of the Arabian Sea from Pakistan
to south India and off Sri Lanka. Further east it extends as far as China, Japan and New Guinea.
0.5 mm
Fig. 283. Solenocera crassicornis, protozoea III, Dorsal view.

Lap. lappets; LSp. lateral abdominal spines; T. telson; Ur. uropods; i1 -i7. 1st -7th caudal setae.

Phylum Arthropoda
Solenocera crassicornis (H. Milne Edwards, 1837) (Fig. 284 a - e)

References: Holthuis, 1980; Cheung, 1963.
Description: Mysis (Acanthosoma), TL = 5.9 mm; CL - 1.6 mm. No spinules on surface of carapace
(Fig. 284 a, b), no lappets; small spines near serrated lateral margins; dorsal organ still present
(DO in Fig. 284 b); carapace fused with all thoracic tergites. Spines on abdominal
terga as in protozoea, but lateral spines much smaller and 2nd dorsal spine largest (Fig. 284 a).
Telson deeply forked (Fig. 284 d, e), 4 setae on each inner margin of each prong (i1 - i4 in Fig.
284 e), pair of small spines on outer margins proximal to fork (i5 in Fig. 284 e). Uropods well-
developed, numerous long setae on each margin of endopod and on inner margin of exopod
(Ex Ur, End Ur in Fig. 284 d). First 3 pereiopods with rudimentary chelae (Ch in Fig.284 c); all
pereiopods with natatory exopods.
Distribution: Arabian Gulf, along the northeast and east coasts of the Arabian Sea from Pakistan
to south India and off Sri Lanka. Further east, it extends as far as China, Japan and New Guinea.
e
b d
Fig. 284. Solenocera crassicornis, mysis. a. lateral view; b. dorsal surface of carapace; c. distal part of 3rd pereopod;
d. caudal fan; e. telson.
DO. dorsal organ; DSp. dorsal spines; I-VI. abdominal tergites; DO. dorsal organ; Ch. rudiment chelae; P3. 3rd
pereopod; T.telson; Ex Ur. exopod of uropod; En Ur. endopod of uropod; i1-i4. inner setae; i5. outer setae.

Phylum Arthropoda
Solenocera hextii Wood-Mason and Alcock, 1891. (Fig. 285 a, b)

References: Dos Santos, Lindley, 2001; Cheung, 1963.
Description: Protozoea III (Elaphocaris), TL = 2.6 mm; CL = 1.0 mm Carapace with spines and
processes (Fig. 285). Carapace margin serrate, spines robust. Dorsal organ (on anterior median
part of carapace) large (DO in Fig. 285 a). Each abdominal tergum with one dorsal and pair of
lateral spines (Fig. 285 a). Telson bifurcate posteriorly, each half with 8 setae (, i1 - i8 in Fig. 285 b).
Distribution: It occurs in the Arabian Sea from the Gulf of Aden to the south coast of India. It is
found at depths between 115 and 500 m.
0.5 mm
Fig.285. Solenocera hextii, protozoea III. a. lateral view; b. dorsal view of telson (T).
Ro. rostrum; DO. dorsal organ; Lap. lappets; DSp. dorsal abdominal spines; P1-P5. rudiments of thoracic legs
(pereopods); Ur. uropod; i1-i8. 1st-8th caudal setae.

Phylum Arthropoda
Superfamily Sergestoidea Dana, 1852
Family Luciferidae de Haan, 1849
Genus Lucifer Thompson, 1829
Lucifer hanseni Nobili, 1905 ( Fig. 286 a - d)

References: Hashizume, 1999; Dos Santos and Lindley, 2001.
Description: Protozoea III, TL = 0.98 mm; CL = 0.34 mm (Northwestern Arabian Gulf ). Carapace
posterior margin with 3 small spines (Sp in Fig.286 a, b). Both rudimentary eyestalks situated
closely together (h in Fig.286 c). Third spine (i3) of telson about 3 times as long as 4th one (i4).
a b
0.1 mm
d
Fig. 286. Lucifer hanseni, protozoea III. a. dorsal view (scale bar 0.1 mm); b. lateral view;
c. frontal part of carapace (dorsal view); d. telson (dorsal view).
Sp. spines; Ur. uropod; E. eye, h. distance between rudimentary eyestalks; i1-i5. 1st - 5th spines of telson.

Phylum Arthropoda
Lucifer hanseni Nobili, 1905 (Fig. 287 a - d)

References: Hashizume K., 1999; Dos Santos, A. and Lindley, J.A. 2001.
Description: Mysis I, TL= 2.15 mm; CL=0.50 mm (Northwestern Arabian Gulf ). Carapace smooth,
dorsomedian spines absent on abdominal somites 1-5, abdominal somites with ventral spines
(Sp in Fig. 287 b). Uropod with tooth (J) on lateral margin of exopod not reaching to posterior
edge (K) of exopod (Fig. 287 c). Second spine (i2) of telson about 2 times as long as the first one
(i1) (Fig. 287 d).
a d
0.2 mm
b
c
Fig. 287. Lucifer hanseni, mysis I. a. general lateral view; b. abdomen; c. distal part of exopodal uropod;
d. telson.
I-VI. abdominal somites; Sp. ventral spines; J. tooth on lateral margin of exopodal uropod; K. posterior edge of
exopodal uropod; i1-i5. 1st-5th spines of telson.

Phylum Arthropoda
Lucifer hanseni Nobili, 1905 (Fig. 288 a-d)

References: Omori, 1992; Hayashi and Tsumura, 1981
Description: Male, TL=10.4 mm; CL= 2.9 mm. It is easily identified by the rather stumpy shape
of its body (Fig. 288 a) as well by the shape of the petasma and the sixth abdominal segment
in males and by the comparative length of the outer spine of uropodal exopods in both sexes.
Terminal portion of petasma (P) acute and curved (Fig. 288 c). Eyestalk (E) less than half distance
between bases of eye and labrum (N) (Fig. 288 a). Outer marginal spine of uropodal exopod (J)
not reaching lamellar part (K) (Fig. 288 b).
Distribution: Indo-West Pacific Region from Madagascar to Australia; Arabian Gulf.
1 mm
b
c
Fig. 288. Lucifer hanseni, male. a. general lateral view; b. distal portion of uropodal exopod;
c. two-sided view of petasma; d. lateral view of telson.
E. eyestalk; N. distance between base of eyestalk and labrum; J. outer marginal spine of uropodal exopod, K.
lamellar part; P. petasma; VP. ventral process.

Phylum Arthropoda
Lucifer typus H. Milne Edwards, 1837 (Fig. 289 a - d)

References: Brooks, 1882; Omori, 1992; Hayashi, Tsumura S., 1981; Khan, 1976. Kensley, 1971.
Description: Male: TL=10.8 mm; CL=2.8 mm. L. typus is distinguished by long eyestalk which is
more than half distance between bases of eye and labrum (Fig. 289 a). The petasma has a robust
terminal portion the sheath is apically blunt and encloses the broad processus ventralis and a
strong hooked process (Fig. 289 b). In males the ventral process of telson situated some distance
from the apex (more anteriorly) (Fig. 289 d) and the posterior ventral process on sixth abdominal
segment well curved (Fig. 289 c).
Distribution: Circumglobal distribution in tropical and subtropical waters of the Atlantic and
Indo-West Pacific; Arabian Gulf.
1 mm
c
d
Fig. 289. Lucifer typus, male. a. general lateral view; b. fragment of 1st pleopod; c. sixth abdominal segment
d. lateral view of telson (uropods removed).
E. eyestalk; N. distance between base of eyestalk and labrum; P. petasma; PP. posterior process; VP. ventral
process.

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

Phylum Arthropoda
Subphylum Crustacea Brunnich,1772
Class Malacostraca Latreille,1802
Subclass Eumalacostraca Grobben,1892
Superorder Eucarida Calman,1904
Order Decapoda Latreille,1802
Suborder Dendrobranchiata Bate,1888
Superfamily Sergestoidea Dana,1852
Family Sergestidae Dana, 1852
Genus Acetes H.Milne-Edwards, 1830
Acetes japonicus (Kishinouye, 1905) (Fig. 290)

References: Farfante and Kensley, 1997; Costa et al., 2003.
Description: Small sized shrimps. The adult total length ranging between 1 and 4 cm. The body
is translucent or semitranslucent with black eyes and several pairs red pigment spots on the
basis and endopod of uropods. Carapace with poorly developed crests and grooves, rostrum
shorter than the eyestalk, generally small (ES, R in Fig. 290). Rostrum with first dorsal tooth (R,
DT in Fig. 290). The pleurae of the second abdominal segment not overlapping those of first
segment. Fourth and fifth pair of pereopods entirely lacking (xx in Fig. 298). First maxillae and
first maxillipeds without palp (IMXLP in Fig. 290). Second maxillae with a single undivided lobe
(IIMXL in Fig. 290).
Female: No protuberance in genital area. Petasma absent. Lower antennular flagella without
spines (AF in Fig. 290). First segment of main branch of lower antennular flagella without
triangular projection. Apex of telson rounded or truncated (T, AT in Fig. 290). Third thoracic
sternite produced posteriorly. Procurved tooth absent between bases of 1st pair of pleopods
(1Pl in Fig. 290).
Male: A pair of protruberances (genital coxae) is present between third pereopods and first
pleopods. Petasma present on first pleopods. Lower antennular flagella with two clasping
spines. First segment of main branch of lower antennular flagella without traingular projection.
Apex of telson rounded or truncated. Third thoracic sternite produced posteriorly. Procurved
tooth absent between bases of first pair of pleopods.

A
Fig. 290. Acetes japonicus, female. Scale - 1 mm.

A. general view; ES. eyestalk; R. rostrum; DT. dorsal tooth; ANTL. antenulla; AF. lower antennular flagella; 3P. third
pereopod; xx. lack of fourth and fifth pereopod; 1Pl. first pleopod; IIMXL. second maxillae; IMXLP. first maxilliped;
T. telson; AT. apex of telson.

Phylum Arthropoda
Genus Sergestes H. Milne Edwards, 1830
Sergestes sp. (Fig. 291)

References: Dos Santos, Lindley, 2001; Illig, 1927; Karuppasamy et al., 2006.
Description: Protozoea II (Elaphocaris). TL = 1.2 mm; CL = 0.75 mm. Eyes stalked (E in Fig. 291).
Carapace with spines and processes. Carapace margin smooth , spines long and slender. Lateral
and posterior processes with long lateral spines. Anterior, lateral and posterior carapace processes
with numerous long lateral spines. Lateral lappets of carapace with 10 spines (Lap in Fig. 291).
0.2 mm
Fig. 291. Sergestes sp. protozoea II.

E. eyes; Ca. carapace; Lap. lateral lappets; T. telson

Phylum Arthropoda
Genus Sergestes H. Milne Edwards, 1830
Sergestes sp. (Fig. 292 a-b)

References: Dos Santos, Lindley, 2001. 2. Illig, 1927. 3. Karuppasamy et al, 2006.
Description: Mysis (Acanthosoma), TL = 2.9 mm; CL = 0.8 mm. Carapace with many spines and
processes (Lap, LSp in Fig. 292 a). Carapace margin smooth, spines long and slender (Lap, LSp in
Fig. 292 a). Carapace without posterior dorsal spine (Ca in Fig. 292 a), eye asymmetrical (E in Fig.
292 a). Eyestalk equal width of eye (E in Fig. 292 a).
0.5 mm
Fig. 292. Sergestes sp. mysis (acanthosoma). a. lateral view, b. telson.

An1. antennule; An2. antenna; Ro. rostrum; SSp. supraorbital spine; E. eye; Lap. lappets; Ca. carapace; LSp. lateral
spines; P1-P3. 1st - 3rd pereopods; Plp1. Plp5 - 1st - 5th pleopods; T. telson; Ex Ur. exopod of uropod; En Ur.
endopod of uropod.

Phylum Arthropoda
Suborder Pleocyemata Burkenroad, 1963
Infraorder Caridea Dana, 1852
Family Hippolytidae Spence Bate, 1888
Genus Tozeuma Stimpson, 1860
Tozeuma sp. (Fig. 293 a - d)

References: Gurney, 1936; Bourdillon-Casanova, 1960; Williamson, 1976.
Description: Zoea I, TL = 3.8 mm; CL = 0.6 mm. Body not covered with spicules; abdomen with one
spines; postero-ventral carapace margins rounded (Fig. 293 a, b); a median dorsal spine of somite 3
(DSp in Fig. 293 c); no median ventral spine on somite 5 (V in Fig.293 c), no pleural spines on
somites 1- 6 (I-VI in Fig.293 c); no supraorbital spines; rostrum (Ro) longer than antennular
peduncle (Fig. 293 b); telson triangular with concave posterior margin (T in Fig. 293 d), telson
formula 3 + 3 (i1 – i3 in Fig.293 d).
0.5 mm
c d
Fig. 293. Tozeuma sp. zoea I. a. lateral view; b. anterior part; c. posterior part; d. telson (T).
Ro. rostrum; DSp. dorsal spine; I-V. 1st-5th abdominal somites; i1-i3. 1st-3rd spines.

Phylum Arthropoda
Family Thalassocarididae Bate, 1888
Genus Thalassocaris Stimpson, 1860
Thalassocaris obscura Menon and Williamson, 1971 (Fig. 294 a, b)

Reference: Menon and Williamson. 1971
Description: Zoea XI, TL = 6.4 mm; CL = 1.9 mm. Larvae of Thalassocaris obscura are most easily
distinguished from those of the other two species by the comparatively narrow antennal scale
(An2 in Fig. 294 b).
Distribution: Indo-west Pacific; Arabian Gulf.
1 mm
Fig. 294. Thalassocaris obscura, zoea XI. a. later l view; b. dorsal view.
An2. antennal scale.

Phylum Arthropoda
Family Oplophoridae Dana, 1852
Genus Acanthephyra A. Milne Edwards, 1881
Acanthephyra sp. (Fig. 295 a, b)

References: Williamson,1957a; Williamson, 1962.
Description: Zoea IV, TL =3.4 mm; CL = 0.6 mm. Abdominal somite. 3 of zoeal stages with
conspicuous fat-filled hump (H in Fig. 295 a) rounded in lateral view but forming a dorsal carina.
Cornea of eye wider than stalk in zoeal stages (E in Fig. 295 a). Rostrum with a few dorsal teeth in
late zoeal stages (Ro in Fig. 295 a). Pair of lateral spines on posterior margin of abdominal somite
5 from stage II or III. Number of telson processes 6 + 6 (T, i1- i6 in Fig. 295 b).
a b
0.5 mm
Fig. 295. Acanthephyra sp. zoea IV. a - lateral view, b. telson (T).
T. telson; i1-i6. 1st-6th telson spines; E. eyes; Ro. rostrum; H. hump; I-VI. abdominall somites.

Phylum Arthropoda
Family Palaemonidae Rafinesque, 1815
Genus Palaemon Weber, 1795
Palaemon sp. (Fig. 296 a - d)

References: Williamson, 1957a; Fincham and Williamson, 1978; Shy et al., 2005.
Description: Zoea IV, TL =4.0 mm; CL = 0.6 mm. Carapace with 1 -3 dorsomedian teeth on the
anterior part (DT in Fig. 296 b). Dactylus of fifth pereiopod armed by 4 curved spines (d, 1-4 in
Fig. 296 c).
0.5 mm
D
c
Fig. 296. Palaemon sp. zoea IV. a. lateral view; b. anterior part of carapace;
c. distalmost part of fifth pereopod; d. teson.
Ro. rostrum; DT. dorsomedian teeth; P5. fifth pereopod; D. dactylus; 1-4. spines; T. teson; i1-i4.
1st-4th telson spines.

Phylum Arthropoda
Family Palaemonidae Rafinesque, 1815
Subfamily Pontoniinae Kingsley, 1878
Genus Periclimenes Costa, 1844
Periclimenes sp. (Fig. 297 a - d)

References: Williamson,1957a; Fincham and Williamson, 1978; Wear, 1976.
Description: Zoea IV, TL =2.7 mm; CL = 0.4 mm. Body with double bend (mesocaris form), acutely
angled or “humped” (Fig.297 a, b). Rostrum short (Ro in Fig. 297 c). Carapace with supraorbital
spines only (SSp in Fig. 297 c).
a b
0.5 mm 0.4 mm
c d
Fig. 297. Periclimenes sp. zoea IV. a, b. zoea IV lateral view; c. anterior part of carapace; d. telson.
Ro. rostrum; SSp. supra-orbital spine; i1-i6. 1st 6th telson spines.

Phylum Arthropoda
Family Alpheidae Rafinesque, 1815
Genus Alpheus Fabricius, 1798
Alpheus spp. (Fig. 298 a - d)

References: Dos Santos and Gonzales-Cordillo, 2004; Williamson, 1967
Description: Zoea IV, TL = 3.7 mm; CL = 0.9 mm (Fig. 298 a). Zoea IV, TL = 3.5 mm ; CL = 0.8 mm
(Fig. 298 b). Eyes stalked, rounded. Leg 5 uniramous and much longer than leg 4. Leg 5 reaching
anterior part of carapace. Telson widest in anterior half. Telson with 4 + 4 setae (i1-i4 in Fig. 298 c).
1 mm
b c
0.5 mm
Fig. 298. Alpheus sp. a, b. lateral view of Alpheus sp. zoea IV; c. telson of Alpheus sp. zoea IV;
d. adult specimen of Alpheus sp.
i1-i4. 1st-4th telson spines.

Phylum Arthropoda
Family Hippolytidae, Dana, 1852
Genus Latreutes Stimpson, 1860
Latreutes sp. (Fig. 299 a - e)

References: Williamson,1957a; Williamson, 1957b; Yang, 2005; 2007.
Description: Zoea IX, TL = 4.0 mm, CL = 0.8 mm. Eye-stalks cylindrical (Fig. 299 a). Telson
flat without a median spine (Fig. 299 e). Rostrum prominent, slender, and tapering in early
stages, deep and most oblong in later stages (Fig. 299 c); carapace with anterior and posterior
dorsomedian papillae (DP in Fig. 299 b); ventral margin of carapace with anterior, middle, and
posterior denticles in first zoea and series of anterior and strong posterior denticles in late zoea
(Fig. 299 d).
b
0.5 mm
e
0.5 mm
Fig. 299. Latreutes sp. zoea IX. a, b. lateral view; c. anterior part; d. carapace; e. telson.
DP. dorsomedian papillae; Ro. rostrum; AD. anterior denticles, PD. posterior denticles; i1-i7. 1st-7th
telson spines.

Phylum Arthropoda
Family Hippolytidae, Dana, 1852
Genus Lysmata Stimpson, 1866
Lysmata sp. (Fig. 300 a - h)

References: Williamson,1957a; Williamson, 1957b.
Description: Zoea IV, TL = 4.0 mm; CL = 0.6 mm (Fig. 300 a). Zoea VI, TL = 7.5 mm; CL = 1.3 mm
(Fig. 300 f ). Eye-stalk very long (Fig. 300 a, f, e).
a b
0.5 mm
e d
f 1 mm
Fig. 300. Lysmata sp. a. zoea IV, lateral view, b. anterior part,
c. ventral part of carapace and maxillipeds; d. distal part of abdomen and telson,
e. telson; f. zoea VI, lateral view, g. anterior part, h. rostrum.
E. eyes; An1. antennules; An2. antenna; Ro. rostrum; Dsp. dorsal spine; i1-i8. 1st-8th telson spines; P5. fifth pereopod;
Ro. rostrum, i1-i7. 1st-7th telson spines.

Phylum Arthropoda
Infraorder Thalassinidea Latreille, 1831
Family Callianassidae Dana, 1852
Genus Callianassa Leach, 1814
Callianassa spp. (Fig. 301 a - e)

References: Gurney, 1938; Sandifer, 1973.
Description: Zoea I, TL = 3.2 mm; CL = 0.6 mm (Fig. 301 a, b). Zoea I, TL = 3.25 mm; CL = 0.6 mm
(Fig. 301 c, d, e). Anterolateral carapace margin toothed. Rostrum narrow, pointed (Fig. 301 a - d).
Eyes sessile (Fig. 301 a, b). Abdomen (Fig. 301 a, b) of 5 somites; 6th somite fused with telson (Fig.
301 a, b, e). Somite 2 with very large dorsal spine overlapping all of somite 3 (DSp in Fig. 301 a);
somites 3 to 5 with small medial spines on posterodorsal border (MSp in Fig. 301 a). Telson (Fig.
301 e) triangular with 7 + 1 + 7 terminal processes (i1 - i7 in Fig. 301 e); unpaired medial spine
large and unarticulated (USp in Fig.301 e); 2nd processes hair-like (i2 in Fig. 301 e).
0.2 mm
b
d
0.2 mm
Fig. 301. Callianassa spp. zoea I. a, c. lateral view; b, d. anterior part; e. telson.
DSp. dorsal spine; MSp. medial spines; I - V. abdominal somites; Ro. rostrum; i1-i7. 1st-7th terminal processes;
USp. unpaired medial spine.

Phylum Arthropoda
Family Laomediidae Dana, 1852
Genus Laomedia De Haan, 1841
Laomedia sp. (Fig. 302 a-e)

Reference: Yaldwin and Wear, 1972.
Description: Zoea IV, TL= 5.0 mm; CL = 1.4 mm; (A). Zoea IV, TL= 4.60 mm; CL = 1.4 mm; (B).
Rostrum slender and unarmed, tapering to a point and extending well beyond the level of the
unstalked eyes (Fig. 302 a, b, c); region between mouth and antennae not lengthened, and there
is no evidence of a “neck”. Abdomen (Fig. 302 a, b, d) consists of five segments and a telson;
abdominal segments unarmed and lacking pro-curved pleural hooks. Telson (Fig. 302 e) almost
straight, the posterior margin with temperate posterior cleft. Posterior telson margin armed with
eigth pairs of setae: first (outer) seta very short and stout, articulating with telson plate, and finely
serrated along its inner margin; second seta reduced to a fine hair as is usual among thalassinid
larvae; third setae prominent, fourth to eight setae about equal in length, finely serrated along
their inner and outer margins, and with scattered fine lateral hairs; eight (inner) seta smaller than
fourth to seventh pairs, but otherwise similar to these. Unpaired medial spine (USp) small.
b
a
0.5 mm
c
0.5 mm
e d
1 mm
Fig. 302. Laomedia sp. zoea IV. a. lateral view of anterior part; b. lateral view of posterior part;
c. anterior part of carapace; d. telson; e. lateral view of zoea IV.
I-V. 1st-5th abdominal somites, T. telson; Ro. rostrum; i1-i8. 1st-8th telson spines, USp. unpaired medial spine.

Phylum Arthropoda
Family Upogebiidae Borradaile, 1903
Genus Upogebia Leach, 1814
Upogebia sp. (Fig. 303 a - f )

References: Ngoc-Ho, 1981; Andryszak, 1986.
Description: Zoea I, TL= 3.0 mm; CL = 1.0 mm (Fig. 303 a-d); Zoea III, TL=3.6 mm; CL = 1.2 mm
(Fig. 303 e, f ). Pair of large lateral spines (LSp in Fig. 303 b, c,) on the 5th abdominal segments.
These spines constitute the main features of interest. The presence of abdominal spines can be
considered as an important distinguishing character. Telson formula for zoea I 7+7. No unpaired
medial spine. Telson formula for zoea III 8+1+8. Small unpaired medial spine.
a b
0.5 mm
e f
0.5 mm
Fig. 303. Upogebia sp. zoea I (A-D), zoea III (E, F). a. lateral view of anterior part of zoea I; b. lateral view of posterior
part; c. dorsal view of 5th and 6th abdominal somites; d. telson; e. lateral view of zoea III; f. telson of zoea III.
Ro. rostrum; III-VI. 3rd-6th abdominal somites; LSp. lateral spines; i1-i8. 1st-8th terminal spines; T. telson.

Phylum Arthropoda
Infraorder Palinura Latreille, 1802
Superfamily Palinuroidea Latreille, 1802
Family Scyllaridae Latreille, 1825
Genus Thenus Leach, 1816
Thenus orientalis Lund, 1793 (Fig. 304).

Reference: Barnett et al., 1984.
Description: Phyllosoma stage I, total length 1.12 mm. Cephalic shield broader than long (A in Fig.
304). Eyestalk elongate, unsegmented (E in Fig. 304). Antennule longer than eyestalk (A1 in Fig.
304), unsegmented with 3 terminal setae (S in Fig. 304); Antenna 1/3 length of antennule (A2 in
Fig. 304), slender, unsegmented with terminal seta. 2nd maxilliped of 5 segments without exopod
(M2 in Fig. 304). 3rd maxilliped bears ventral coxal spine and comb-like setae on terminal segment
(M3 in Fig. 304). Pereiopods 1, 2 biramous with setose exopods (P1, P2 in Fig. 304). Pereiopod
3 bears short exopod, usually non-setose (P3, P3ex in Fig. 304). Pereiopods 4 and 5 are absent.
Abdomen narrow, approximately 1/2 length and 1/4 width of thorax (A, B Fig. 304). Uropod buds
absent. Posterior margin of telson slightly indented at midpoint, 2 short posterolateral spines
flanked by 3 short setae.
1 mm
Fig. 304. Phyllosoma stage I of scyllarid lobster Thenus orientalis. Scale - 1 mm.
A1. antennule; A2. antenna; E. eyestalk; S. terminal seta; M2. 2nd maxilliped; M3. 3rd maxilliped; P1, P2, P3.
pereiopods; P3ex. pereiopod 3 exopod; AB. abdomen.

Thenus orientalis (Lund, 1793) (Fig. 305).
Description: Phyllosoma stage II, total length 6.0 mm. Cephalic shield slightly pear-shaped,
twice the width of thorax. Eyestalk segmented (E in Fig. 305). Biramous antennule of 4 segments,
shorter than eyestalk; inner ramus well developed (A1 in Fig. 305). Terminal segment bears 4
major terminal setae and 8 rows of shorter, wide-based setae. Antenna bifurcate, 1/2 length of
antennule; lateral process short (A2 in Fig. 305). 2nd maxilla a single paddle-shaped lobe curved
slightly towards posterior, setae now absent. 1st maxilliped a small bud at base of 2nd maxilla.
Pereiopod 5 bud 1/2 length of abdomen. Thorax more than 1/2 width of cephalic shield. 4 pairs
of pleopod buds evident (PL in Fig. 305). Small biramous uropods. Single spine flanked by 2 setae
at each postero-lateral corner of telson (SP in Fig. 305).
Fig. 305. Phyllosoma stage II of scyllarid lobster Thenus orientalis. Scale - 1 mm.
A1. antennule; A2. antenna; E. eyestalk; P5. pereiopod 5; PL. pleopod buds; SP. spine; UR. uropods.

Thenus orientalis (Lund, 1793) (Fig. 306).
Description: Phyllosoma stage III, total length 9.3 mm. Cephalic shield almost round. Antennule
of 4 segments, medial spur a pointed extension of segment 3; distal segment with 4 long terminal
setae and 8 rows of shorter wide-based setae on inner margin (A1 in Fig. 306). Antenna over 1/2
length of antennule, bifurcate and flattened (A2 in Fig. 306). 1st maxilliped now small lobe at base
of 2nd maxilla (M1 in Fig. 306). 2nd maxilliped of 4 segments (M2 in Fig. 306). Pereiopod 5 of 3
segments, slightly longer than abdomen (P5 in Fig. 306). Abdomen broader than in stages I and
II, slightly less 1/2 width of thorax. Paired uropods developed, flattened and rounded (UR in Fig.
306). Short spine on each lateral margin of telson (SP in Fig. 306). Posteriorly directed median
carina on dorsal surface of each of 5 abdominal segments (CA in Fig. 306).
Fig. 306. Phyllosoma stage III of scyllarid lobster Thenus orientalis. Scale - 1 mm.
A1. antennule; A2. antenna; E. eyestalk; M1. 1st maxilliped; M2. 2nd maxilliped; P5. pereiopod 5; SP. spines;
UR. uropods; CA. median carina; Ad. adult specimen of T. orientalis.

Phylum Arthropoda
Infraorder Anomura MacLeay, 1838
Family Galatheidae Samouelle, 1819
Genus Galathea Fabricius, 1793
Galathea sp. (Fig. 307 a - e)

References: Pike and Williamson, 1972; Dos Santos and Gonzales-Cordillo, 2004.
Description: Zoea III, TL = 2.0 mm; CL = 0.4 mm (Fig. 307). Carapace laterally flattened (Fig. 307 b).
Each posterolateral margin of carapace extended into a spinous process (Fig. 307 a, d). Rostrum
shorter than body (Fig. 307 a). Posterolateral margin process of carapace never extending
beyond the end of telson (Fig. 307 a). Posterolateral margins of carapace denticulate (Fig. 307
d). Antennal exopod flattened (Fig. 307 b). Rostrum narrower than diameter of eye (Fig. 307 b).
a c
0.4 mm
b e
Fig. 307. Galathea sp. zoea III. a. lateral view; b. dorsal view; c. dorsal portion of carapace; d. posterior portion of
carapace; e. telson.
Ro. rostrum; E. eye; Ex An2. antennal exopod; dent. denticles; SpPr. spinous process; T. telson; Ex Ur. exopod of
uropod; En Ur. endopod of uropod.

Phylum Arthropoda
Family Porcellanidae Haworth, 1825
Genus Pachycheles Stimpson, 1858
Pachycheles spp. (Fig. 308 a - d)

References: Dos Santos and Gonzales-Cordillo, 2004; Tirmizi and Yaqoob, 1979; Wear, 1965;
Konishi, 1987.
Description: Zoea I, CL = 1.4 mm; rostral spine length = 5.5 mm; posterior spine length = 2.0
mm. Carapace laterally flattened, typically porcellanid (Fig. 308 a). Each posterolateral margin of
carapace extended into a spinous process (PSp in Fig. 308 a, c). Rostrum much longer than body
(4 times carapace length) armed overall with spinules to its tip (Sp in Fig. 308 a). Posterio-lateral
margin process of carapace extended beyond the end of telson (1.4 times carapace length).
Both posterior spines bearing ventral pointed spinelets near their basal portion (Sp in Fig. 308
c). Lower margin of carapace smooth (Ca in Fig. 308 c). Abdomen with lateral spines increasing
in size posteriorly (LSp in Fig. 308 d). Telson bearing 7 pairs of posterior processes, long and
plumose except for first and second (i1-i7 in Fig. 308 d).
0.5 mm
b d
Fig. 308. Pachycheles sp. zoea I. a. lateral view; b. fragment of rostrum; c. posterior portion of carapace; d. telson.
Ro. rostrum; Ca. carapace; PSp. posterio-lateral margin process; Sp. spinulets; T. telson; LSp. lateral spines; i1-i7.
1st-7th posterior processes.

Phylum Arthropoda
Subphylum Crustacea Brunnich,1772
Order Decapoda Latreille,1802
Family Diogenidae Ortmann, 1892
Genus Diogenes Dana, 1851
Diogenes sp. (Fig. 309 a - e)

Reference: Dos Santos and Gonzales-Cordillo, 2004.
Description: Zoea III, TL = 2.5 mm; CL = 0.8 mm. Carapace laterally flattened, smooth with small
swellings behind eyes. Each posterolateral margin of carapace rounded (Fig. 309 c). Rostrum
large with pointed end, longer than antenna length (Fig. 309 b). Eye stalk relatively short. 5th
somite of abdomen bears dorsal spine (Fig. 309 d). Telson flattened (Fig. 309 e). Posterior margin
without median spine and slightly concave. Each half of telson bears six spines. Small setules
between 2nd - 6th spines. Uropods with exopod and rudimentary endopod.
b a c
d
e
0.5 mm
Fig. 309. Diogenes sp. zoea III. a. lateral view; b. anterior fragment; c. posterior part of carapace; d. fragment of
abdomen.
Ro. rostrum; Ca. carapace; An2. antenna; DSp. dorsal spine on 5th somite; III-VI. 3rd-6th somite; E. distal fragment
of abdomen; T. telson; Ex Ur. exopod of uropod; En Ur. endopod of uropod; i1-i6. 1st-6th caudal spines.

Phylum Arthropoda
Genus Dardanus Paulson, 1875
Dardanus sp. (Fig. 310 a - c)

Reference: Pike and Williamson, 1960.
Description: Zoea I, TL = 1.9 mm; CL = 0.6 mm. Carapace without posterior spine (Fig. 310 a).
Antennal scale without terminal spine. Rostrum broad narrowing more abruptly near tip (Fig. 310
b). No dorsal spine on 5th abdominal somite (Fig. 310 a). Lateral spines on 5th abdominal somite.
Rostrum reaching well beyond antennal scale (Fig. 310 b). Carapace with small dorsal carina (DC in
Fig. 310 a).
0.2 mm
b c
Fig. 310. Dardanus sp. zoea I. a. lateral view, b. dorsal view of anterior part; c. telson.
Ro. rostrum; DC. dorsal carina; (T); i1-i6. 1st-6th telson spines.

Phylum Arthropoda
Genus Paguristes Dana, 1852
Paguristes sp. (Fig. 311 a - d)

References: Pike and Williamson, 1958; Roberts, 1971; Dos Santos and Gonzales-Cordillo, 2004.
Description: Zoea I, TL = 1.8 mm; CL = 0.6 mm (Fig. 311 a, b). Carapace laterally flattened. Eyes
sessile. Posterior margins of carapace rounded. Rostrum not reaching apex of antennule (Fig. 311
a). Eight pairs of caudal spines (i1-i8 in Fig. 311 b).
Zoea I, TL = 2.6 mm; CL = 1.0 mm (Fig. 311 c, d). Carapace laterally flattened. Eyes sessile. Posterior
margins of carapace rounded. Rostrum reaching apex of antennule (Fig. 311 c). Eight pairs of
caudal spines (i1-i8 in Fig. 311 d).
a b
0.5 mm
c d
0.5 mm
Fig. 311. Paguristes sp. zoea I. a, c. lateral view; b, d. telson.

T. telson; i1-i8. 1st-8th telson spines.

Phylum Arthropoda
Family Paguridae Latreille, 1802
Genus Pagurus Fabricius, 1775
Pagurus sp. (Fig. 312 a - c)

References: Pike and Williamson, 1958; Roberts, 1971; Dos Santos and Gonzales-Cordillo, 2004.
Description: Zoea III, TL = 3.4 mm; CL = 1.1 mm. Carapace laterally flattened (Fig. 312 a). Eyes
sessile. Biramous uropods present. Each posterolateral margin of carapace extended into a
spinous process (SpPr in Fig. 312 a, b). Rostrum shorter than body (Fig. 312 a, b). Posterolateral
margins of carapace smooth (Fig. 312 a, b). Eight pairs of telson spines (i1-i8 in Fig. 312 c).
a c
0.5 mm
Fig. 312. Pagurus sp. zoea III. a. lateral view, b. lateral view of carapace, c. telson.
SpPr. spinous process; T. telson; i1-i8. 1st-8th telson spines.

Phylum Arthropoda
Infraorder Brachyura Latreille, 1802
Family Leucosiidae Samouelle, 1819
Genus Ebalia Leach, 1817
Ebalia sp. (Fig. 313 a, b)

References: Dos Santos and Gonzales-Cordillo, 2004; Ingle, 1992; Pohle et al., 1999.
Description: Zoea I, CL = 0.6 mm. Telson with triangular shape and short furcal spines lacking
medial arch. Five pairs of furcal spines (Fig. 313 b). Carapace without dorsal spine (Fig. 313 a).
0.2 mm
Fig. 313. Ebalia sp. zoea I. a. lateral view; b. telson.

Ro. rostrum; i1-i5. 1st-5th telson spines; T. telson.

Phylum Arthropoda
Family Leucosiidae Samouelle, 1819
Genus Leucosia Weber, 1795
Leucosia sp. (Fig. 314 a, b)

References: Dos Santos and Gonzales-Cordillo, 2004; Ingle, 1992; Pohle et al., 1999.
Description: Zoea I, TL = 3.8 mm; CL = 0.8 mm. Telson with triangular shape and short furcal
spines lacking medial arch. Four pairs of furcal spines (Fig. 314 b). Carapace with long rostral,
dorsal and lateral spines (Fig. 314 a).
1 mm
Fig. 314. Leucosia sp. zoea I. a. lateral view; b. telson.

Ro. rostrum; DSp. dorsal spine; LSp. lateral spine; T. telson; i1-i4. 1st-4th spines.

Phylum Arthropoda
Family Parthenopidae MacLeay, 1838
Genus Parthenope Weber, 1795
Parthenope sp. (Fig. 315 a - e)

References: Yang 1971; Guerao and Abello, 1999.
Description: Zoea IV, TL = 3.5 mm, CL = 1.1 mm. Carapace globose with the full complement of
dorsal, rostral and lateral spines but without conspicuous projection (Fig. 315 a, b). Abdominal
somites 3-5 with prominent posterolateral spines, about as long as length of somite (PLSp in Fig.
315 c, d); telson furca arched (T in Fig. 315 e).
a b
0.5 mm
0.5 mm
c d e
Fig. 315. Parthenope sp. zoea IV. a, b. lateral view; c. lateral view of abdomen, d. dorsal view of abdomen, e. dorsal
view of telson.
Pl. pleopod buds; I-V. abdominal somites; T. telson; PLSp. posterolateral spines; LSp. lateral spines; i1-i5. 1st-5th
furcal spines.

Phylum Arthropoda
Family Ocypodidae Rafinesque, 1815
Genus Macrophthalmus Desmarest, 1823
Macrophthalmus sp. (Fig. 316 a, b)

References: Fielder and Greenwood, 1986; Hashmi, 1969; Pasupathi and Kannupandi, 1988.
Description: Zoea III, CL = 0.7 mm. Carapace smooth and globose (Fig. 316 a). Dorsal and lateral
spines absent. Rostrum very short and evenly tapered (Fig. 316 a). Eyes immobile. Telson slightly
wider than last abdominal somite (Fig. 316 b). Telson posterior margin transverse with 3 + 3
subequal biplumose setae (i2-i4 in Fig. 316 b). Telson rami short (i1 in Fig. 316 b).
0.2 mm
Fig. 316. Macrophthalmus sp. zoea III. a. lateral view, b. telson.

Ro. rostrum; T. telson; i1-i4. 1st-4th telson spines.

Phylum Arthropoda
Family Ocypodidae Rafinesque, 1815
Genus Ilyoplax Stimpson, 1858
Ilyoplax frater (Kemp, 1919) (Fig. 317 a - c)

References: Al-Khayat and Jones 1996.
Description: Zoea VI , TL = 4.5 mm, CL = 0.75 mm. Carapace wider than long. Dorsal spine longer
than length of carapace. Rostral spine approximately two thirds length of dorsal spine (Fig.
317 a). Pair of lateral spines on both sides of carapace. Eyes rounded (Fig. 317 b). Five subequal
abdominal somites. Telson elongate with posterior margin with deep median cleft and 3 + 3
plumose setae (i1 - i3); long furcae without rows of spinules (T in Fig. 317 c).
Distribution: Indian Ocean; Arabian Gulf.
0.5 mm
Fig. 317. Ilyoplax frater, zoea VI. a. lateral view; b. lateral view of carapace; c. distal part of abdomen and telson.
E. eye; T. telson; i1-i3. 1st -3rd setae.

2.16. Chaetognatha
Phylum Chaetognatha (Leukart, 1894)

Class Sagittoidea Claus and Grobben, 1905
Order Aphragmophora Tokioka, 1965
Family Sagittidae Claus and Grobben, 1905
Genus Sagitta Quoy and Gaimard, 1827
Sagitta enflata Grassi, 1881 (Fig. 318 a - e)
Description: Hooks: 8-10 (Fig. 318 b, c); anterior teeth: 4-10; posterior teeth: 4-15 (AT, PT in Fig.
318 b). Maximum adult body length: 25 mm; tail: 14-17% (Fig. 318 a). Body flaccid, transparent;
transversal musculature absent; head of medium width (Fig. 318 a, c); hooks not serrated (H in
Fig. 318 b). Fin bridge absent. Anterior fins very short, partially rayed, round; posterior fins short,
partially rayed, round (Fig. 318 a). Collarette absent (Fig. 318 c); gut diverticula absent. Small eyes,
with star-shaped pigment spot (Fig. 318 c). Seminal vesicles round; position of seminal vesicles:
touching tail fin, well-separated from posterior fins (Fig. 318 a, d). Ovaries short, reaching to
middle of posterior fins; ova large (Fig. 318 e).
Distribution: Circumglobal, between 40° N and 40° S; Arabian Gulf.
b c
a
d e
Fig. 318. Sagitta enflata. a. general view; b. part of head; c. head; d. seminal vesicle; e. ovary. Scale - 0.1 mm.
SV. seminal vesicle; O. ovary; AT. anterior teeth; PT. posterior teeth; H. hooks.

Sagitta neglecta Aida, 1897 (Fig. 319 a - f )
Description: Hooks: 3-7 (H in Fig. 319 d); anterior teeth: 7 (AT in Fig. 319 d); posterior teeth: 18 (PT
in Fig. 319 d). Maximum adult body length: 8 mm. Body firm, opaque; transversal musculature
absent; head narrow; hooks not serrated (H in Fig. 319 d). Fin bridge absent (Fig. 319 a). Anterior
fins of medium length, fully rayed, rounded. Posterior fins of medium length, fully rayed (PF in
Fig. 319 f ), rounded. Long and narrow collarette (Fig. 319 c). Gut diverticula absent (Fig. 319 a).
Distribution: Indo-Pacific, between 30° N and 30° S; Arabian Gulf.
a
c
e f
Fig. 319. Sagitta neglecta. a. general view; b. head; c. collarette; d. hooks; e. aboral end;
f. posterior fin. Scale - 1 mm.
AT. anterior teeth; PT. posterior teeth; H. hooks; PF. posterior fin; TF. tail fin.

Sagitta pulchra Doncaster, 1902 (Fig. 320)

Description: Hooks: 6-7; anterior teeth: 8-10; posterior teeth: 8-16. Maximum adult body length:
24 mm; tail: 15-20 % (Fig. 320). Body firm, opaque; transversal musculature absent; head wide;
hooks not serrated. Fin bridge absent. Anterior fins long, partially rayed, angular; posterior fins of
medium length, partially rayed, angular. Collarette short; gut diverticula absent. Small eyes, with
star-shaped pigment spot. Seminal vesicles with knob and trunk; position of seminal vesicles:
touching both posterior fins and tail fin. Ovaries of medium length, reaching to region of ventral
ganglion; ova small. Adhesive papillae and adhesive organs absent. Apical glands absent.
Fig. 320. Sagitta pulchra, general view. Scale - 1 mm.

Sagitta regularis Aida, 1897 (Fig. 321)
Description: Hooks: 10-11 (H in Fig. 321); anterior teeth: 4; posterior teeth: 6. Maximum adult
body length: 6 mm; tail: 33%. Body firm, opaque; transversal musculature absent; head narrow;
hooks not serrated. Fin bridge absent. Anterior fins of medium length, fully rayed, rounded;
posterior fins medium length, fully rayed, rounded (AF, PF in Fig. 321). Collarette long and very
conspicuous (C in Fig. 321); gut diverticula present (G in Fig. 321). Small eyes with T-shaped
pigment spot. Seminal vesicles with knob and trunk (SV in Fig. 321); position of seminal vesicles:
touching posterior fins and separated from tail fin. Ovaries of medium length, reaching to region
of ventral ganglion; ova large. Adhesive papillae and adhesive organs absent. Apical glands
absent.
Fig. 321. Sagitta regularis. Scale - 1 mm.

C. collarette; H. hooks; G. gut diverticula. AT. anterior teeth; PT. posterior teeth; AF. anterior fin; PF. posterior fin;
TF. tail fin; SV. seminal vesicles; G. gut diverticula.

2.17. Urochordata
Phylum Chordata Bateson, 1885

Subphylum Urochordata (=Tunicata) Giribet et al., 2000
Class Appendicularia (=Copelata or Larvacea) (Fol 1872)
Family Oikopleuridae (Lohman, 1915)
Subfamily Oikopleurinae Lohman, 1896
Supergenus Labiata Fenaux, 1993
Genus Oikopleura Mertens, 1830
Oikopleura dioica Fol, 1872 (Fig. 322)
Description: Trunk compact, 1.5 mm long (T, R in Fig. 322). Buccal glands spherical and small (BG
in Fig. 322). Dioecious. Testis or ovary adjoining the coil of the gut hemispherically (O in Fig. 322).
Tail with 2 spindle-shaped subchordal cells arranged in a line, musculature narrow. Tail six times
as long as the trunk. Mouth terminal or lying antero-dorsally (M in Fig. 322).
Distribution: Semi-cosmopolitan, neritic species from warm and temperate waters. Atlantic,
Indian and Pacific Oceans. Also present in Mediterranean Sea and Red Sea. From all Appendicularia,
O. dioica is the most eurytherm and euryhaline species
Fig. 322. Oikopleura dioica. Scale - 0.1 mm.

b. trunk (left side) covered by “house”; CH. chorda; TM. tail musculature; TF. tail fin; T. trunk, top view; BG. buccal
glands; O. ovary; R. trunk right side; M. mouth.

Family Fritillariidae Lohmann, 1915
Subfamily Appendiculariinae Seeliger, 1895
Genus Appendicularia Fol, 1874
Appendicularia sicula Fol, 1874 (Fig. 323 a - c)

Reference: Marine Species Identification Portal (2000-2010).
Description: Trunk is short, slightly pear-shaped, swollen posteriorly and depressed anteriorly
(Fig. 323 a, b). Mouth round, with tacticle hairs; there are no extended lips. Rectum vast, pear-
shaped, occupying a big part of the digestive tract. The short endostyle is located close to the
mouth, with broadened proximal part. No anus visible. Gonads located above the digestive tract,
in the middle of the dorsal part of the trunk. The ovary is spherical, enveloped by the testis,
except for the upper part. Tail narrow near the body, gradually widening to moderately wide,
bifurcated at the free end (Fig. 323 c). Tail muscles very thin. The shape of the tail is characteristic
for the species. Size: Trunk 0.45-0.5 mm. Tail length 1.5 mm. The oikopleurid house is 1.5-2.5 mm
in diameter.
Distribution: Warm and temperate waters of the Atlantic, Pacific and Indian Oceans, the
Mediterranean and Red seas.
a c
Fig. 323. Appendicularia sicula.

a. general view; b. trunk; c. tail.

Subfamily Fritillarinae Seeliger, 1895
Genus Fritillaria Fol, 1872
Fritillaria sp. (Fig. 324 a - b)
Description: Trunk very elongate or flat (Fig. 324 a, b). Oral part of the trunk covered by
membranous hood (H in Fig. 324). Mouth with lobes. Stomach spherical (S in Fig. 324). Intestine
and rectum below and behind. Gonads behind digestive tract. Tail with broad fin, strongly
intended near its root.
Fig. 324. Fritillaria sp. a. general view (left side); b. trunk (left side).
H. hood; S. stomach.

Class Thaliacea Nielsen, 1995
Order Salpida Uljanin, 1884
Family Salpidae Lahille, 1888
Genus Weelia Yount, 1954
Weelia cylindrica (Cuvier, 1804) (Fig. 325 a - d)
Description: In both generations, the transverse ribs of the gill bar meet dorsally, narrowing
sharply toward the ventral surface, so that they are far apart from one another ventrally. Solitary
zooids: up to 45 mm long. Test soft and smooth. Body elongate and cylindrical (Fig. 325 a).
Eight or 9 body muscles that extend rather far ventrally and are usually separated. Ml to MIV
converging, contiguous, or fused in the mid-dorsal line. A great variation is also found in the
number of muscular fibers. Aggregate zooids: up to 17 mm long, excluding projections. Body
fusiform because of the presence of short anterior and posterior projections (MV, MIV in Fig. 325
d).
a b
Fig. 325. Salpa Weelia cylindrica. a. solitary zooid; b. aggregate zooid (dorsal view); c. detail of gill bar;
d. embrio. Scale - 1 mm.
E. embryo. MIV, MV. muscles.

Genus Thalia Blumenbach 1798
Thalia sp. (Fig. 326)
Description: Aggregate zooids: body oval, rounded or pointed posteriorly (Fig. 326). Five body
muscles fused dorsally into two groups: MI to MIII and MIV to MV (MI-MIII, MIV in Fig. 326). All the
muscles interrupted ventrally (I in Fig. 325). Nucleus compact (N in Fig. 326). Testis arranged in 4
lobes. Ovary and embryo near MV (O in Fig. 326).
Fig. 326. Salpa Thalia sp. Aggregate zooid (dorsal view). Scale - 1 mm.
MI-MIV. muscles; O. ovary; I. muscles ventral interruption; N. nucleus.

Class Ascidiacea Nielsen, 1995
Tadpole larva (Fig. 327 a - b)

Reference: Marine Species Identification Portal (2000-2010).
Description: Tunicates of class Ascidiaceae, commonly called sea squirts, have free-swimming,
nonfeeding larvae (Fig. 327 a) and benthic, sessile adults (Fig. 327 b). Tadpole larva has adhesive
suckers (papilli) on its head and a muscular tail. The larval nervous system includes a dorsal
pigment-containing light receptor (ocellus) and a dorsal, hollow nerve cord (neural tube) that
runs the length of its tail. The ascidian larva has a nonfunctional digestive tube. After one day
of swimming, ascidian larvae adhere with their anterior ends to the solid surface and undergo
extensive metamorphosis into the sessile adult. Tail, nerve cord, and notochord are resorbed as
the body rotates about 180 degrees; the oral siphon migrates to open opposite the attachment
to the substrate; other internal organs also rotate to their adult locations.
Fig. 327. a. tadpole larva of Ascidiacea; b. adult ascidia Phallusia nigra.

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

2.18. Cephalochordata

Subphylum Cephalohordata Owen, 1846
Family Branchiostomidae Bonaparte, 1841
Genus Branchistoma Bonaparte, 1841
Branhiostoma sp. (Fig. 328 a - c)
Description: The animal with fishlike (fusiform or lanceolate) shape. The body is divided into
head, trunk and tail (H, TR, TL in Fig. 328 a). The head, at the anterior end, is small and poorly
defined. The rostrum extends anteriorly and overhangs the mouth and buccal cavity (R in Fig.
328 a, b). The large mouth lies under the rostrum and opens into a spacious buccal cavity (M, BC
in Fig. 328 b). The mouth is surrounded by a ring of tentacle-like buccal cirri (=oral cirri). Most of
the body is the trunk (TR in Fig. 328 a), which extends posteriorly from the head to the anus. The
trunk contains most of the gut, including the large conspicuous pharynx and the musculature.
The segmental arrangement of the axial musculature (body musculature) is readily apparent
through the translucent integument. The muscles are arranged in 50-75 V-shaped segmental
bundles called myomeres (MS in Fig. 328 a). There are no paired appendages but on either side
of the trunk is a ventro-lateral longitudinal ridge, the metapleural fold. These ridges run from the
oral hood to a position just posterior to the gonadal region. The atrium opens to the exterior via
the atriopore, located on the midventral margin at the point where the two metapleural folds
join the ventral margin (AT in Fig. 328 a). Farther posteriorly, beside a slight dip between the
ventral fin and the caudal fin, is the anus, located slightly to the left side of the ventral midline
(AN in Fig. 328 a). The anus is the posterior external opening of the gut and marks the posterior
limit of the trunk. The region of the body posterior to the anus is the tail (TL in Fig. 328 a). One
of the characteristics of chordates is the presence of this postanal tail. A posterior caudal fin
(=tail fin) extends around the dorsal and ventral margins of the tail. There is a long dorsal fin
along most of the dorsal margin of the body (DF in Fig. 328 a). A short ventral fin is located on
the ventral margin of the trunk just anterior to the caudal fin (VF in Fig. 328 a). It extends from
atriopore to anus. Segmentally arranged, paired, rectangular swellings along the ventral margins
of the myomeres are about 26 pairs of segmental gonads.

a
Fig. 328. Branchiostoma sp. a, c. general view; b. anterior part. Scale - 0.1 mm.
H. head; TR. trunk; TL. tail; R. rostrum; MS. miomers; AT. atriopore; VF. ventral fin; AN. anus; CF. caudal fin; DF.
dorsal fin; M. mouth; BC. buccal cavity; NC. nerve cord; FS. fin supports; N. notochord; ES. eyespot.

Addendum
While this guide book was in press, we registered in Kuwait waters two new and very interesting
species of Medusa and Larvacea. Hence, we decided to add these species at the end of this guide
book.
Medusa Catostylus mosaicus belongs to Class Scyphozoa. Scyphozoan medusae are included in
volume 1, pages 150-151. The larvacea Fritillaria pellucida belongs to Family Fritillaridae (Phylum
Chordata). The rest of fritillarids are described in volume 2, pages 174-175.
Phylum Cnidaria Verrill, 1865

Subphylum Medusozoa Petersen, 1979
Superclass Hydrozoa Owen, 1843
Class Scyphozoa Goette, 1887
Subclass Discomedusae Lankester, 1877
Order Rhizostomeae Cuvier, 1799
Suborder Daktyliophorae Stiasny, 1921
Family Catostylidae Gegenbaur, 1857
Genus Catostylus Agassiz, 1862
Catostylus mosaicus (Quoy and Gaimard, 1824) (Fig. 229)
Description: C. mosaicus is characterized by lack of tentacles. Instead, it has eight three-winged

mouth arms. There is a network of branching canals linked with the primary ring canal, but
these are not joined to the gastrovascular cavity except through the 16 radial canals. Some of
these radial canals do not extend to the edge of the bell. There are eight sense organs, known
as rhopalia, which have canals extending to the margin of the bell. The bell of C. mosaicus is
creamy white, brown or blue. Color variations depend on a symbiotic relationship of the jellyfish
with microalgae that are found inside its body. The algae photosynthesize and can be used by
the jellyfish. There is no obvious mouth on the underside, but there are small openings on each
arm, through which food is passed into the stomach. The tentacles have stinging cells that can
capture plankton. C. mosaicus can live for up to 13 months; within 1 to 3 months can grow to
sexual maturity.
Measurements: Reported diameter of bell up to 35 cm. Maximum bell diameter of specimens

from Kuwait waters up to 19-20 cm, length of mouth arms up to 15-16 cm. Most of specimens
from Kuwait waters are of creamy white color.
Distribution: Intertidal estuaries and coastal waters of Indo-West Pacific Region; Arabian Gulf.
Bloom of this species was registered in Kuwait waters (Kuwait Bay) in June 2011.
C. mosaicus is part of commercial fisheries in Asian countries. Salted and dried medusas are
popular in China and Japan, where they are considered as a culinary delicacy. The Chinese
believe that eating this jellyfish reduces high blood pressure. They also use this species for
treating arthritis and back pain.

Fig. 229. Catostylus mosaicus, general view.

Subphylum Urochordata (=Tunicata) Giribet et al., 2000
Class Appendicularia (=Copelata or Larvacea) (Fol 1872)
Family Fritillariidae Lohmann, 1915
Subfamily Fritillarinae Seeliger, 1895
Genus Fritillaria Fol, 1872
Fritillaria pellucida (Busch, 1851) (Fig. 230 a-b)
Description: Trunk rectangular, flattened dorso-ventrally, with two conical, well separated
cuticular appendices on its genital end. Mouth with protruding upper lip. Digestive tract axis is
transverse. Gonads asymmetrically arranged. Testis on the right side of the body; it is cylindrical
and transverse in young specimens, later becoming Y-shaped. Ovary is spherical, on the left
side of the body. Tail has broad musculature and two pairs of amphichordal glandular cells with
visible exit canals. Distal extremity of the tail is notched or truncated.
Reported Measurements: Length of trunk ranges from 1.0 to 2.2 mm.
Distribution: Warm and temperate waters of Atlantic, Pacific and Indian oceans; Arabian Gulf.

a
Fig. 230. Fritillaria pellucida. a. trunk (left side); b. general view (right side).

References
Alvarino A. 1967. The Chaetognatha of the Naga expedition (1959-1961) in the China Sea and the Gulf of Thailand. Part 1 - Systematics.
Naga Rep. 4(2): 1-197.
Ali, M., F. Al-Yamani, and I. Prusova. 2007. Bestiolina arabica sp. nov. (Copepoda, Calanoida, Paracalanidae), a new species from the
northwestern Arabian Gulf. Crustaceana. 80: 195-205.
Al-Yamani, F.Y. 1989. Plankton studies in the ROPME Sea Area: Present status and future prospects. Regional Organization for the
Protection of the Marine Environment, Report No ROPME/GC-6/004, Kuwait.
Al-Yamani, F.Y. and A.M. Fahmi. 1995. New copepod host records for ellobiopsid parasites from the Northwestern Arabian Gulf. Arab
Gulf J. Scientific Res. 13(3): 571-581.
Al-Yamani, F.Y.; J. Bishop; K. Al-Rifaie; W. Ismail; A. Al-Yaqout; L. Al-Omran; A. Kwarteng; A. Al-Ghadban; and C. Sheppard. 1997a.
Assessment of the effects of the Shatt Al-Arab’s altered discharge regimes on the ecology of the Northern Arabian Gulf.
Kuwait Institute for Scientific Research, Report No. KISR 5174, Kuwait.
Al-Yamani, F.; R. Durvasula; W. Ismail; K. Al-Rifaie; T. Al-Saeed; A. Al-Yaqout; and L. Al-Omran. 1997b. Dynamic oceanography of the
northwestern waters of the Arabian Gulf: Ecological significance of the marine food web. Kuwait Institute for Scientific
Research, Report No. KISR 5173. Kuwait.
Al-Yamani, F.; J. Bishop; M. Al-Hossaini; D.V. Subba Rao; A.N. Al-Ghadban; N.Y. Khan; M. Al-Salman; M. Abdel-Jawad; M. Tabtabaei; and
M. Behbehani. 2000. Development study for the optimal utilization of the marine environment: characterization of Kuwait’s
marine environment. Kuwait Institute for Scientific Research, Report No. KISR 5991, Kuwait.
Al-Yamani F.Y. and I.Y. Prusova. 2003. Common Copepods of the Northwestern Arabian Gulf: Identification Guide. Kuwait: Kuwait
Institute for Scientific Research, 162 pp.
Al-Yamani, F.Y.; J. Bishop; E. Ramadhan; M. Al-Husaini; and F.N. Al-Ghadban. 2004. Oceanographic Atlas of Kuwait’s Waters. Kuwait:
Kuwait Institute for Scientific Research, 203 pp.
Al-Yamani F.Y. and V.A. Skryabin. 2006. Identification Guide for Protozoans from Kuwait’s Waters. Coastal Planktonic Ciliates: 1.Tintinnids.
Kuwait: Kuwait Institute for Scientific Research, 109 pp.
Al-Yamani F.Y., S.A. Khvorov, and A.S. Khvorov. 2010. Interactive Guide of Planktonic Decapod Larvae of Kuwait’s Waters. Kuwait
Institute for Scientific Researches. Electronic resource. Registry number: 2010/009 – 4, Kuwait.
Al-Ghadban, A.N. 2003. Geological oceanography of the Arabian Gulf, In: Khan, N.Y, Munawar, M., and Price, A.R.G (Eds.), The Gulf
Ecosystem, Health and sustainability, Kuwait Institute for Scientific Research, Backhuys Publishes, pp 23-40.
Al-Ghadban, A.N., D. Alajmi, and M.S. Massoud. 1998. Sedimentation rate and bioturbulation in the Arabian Gulf. Environment
International. 24(1-2), pp. 23-31.
Al-Khayat, J.A. and D.A. Jones. 1996. Two new genera, Manningis and Leptochryseus (Decapoda, Camptandriinae) and descriptions of
first zoea of six brachyurans from the Arabian Gulf. J. Crust. Biol. 16(4): 797-813.
Al-Majed, N., H. Mohammadi, and A. Al-Ghadban. 2000. Regional report of the state of the environment. ROPME/GC-10/001/1. Revised
by A. Al-Awadi, Regional Organisation for the Protection of the Marine Environment, Kuwait, 178 pp.
Al-Muzaini, S., O. Samhan, and M.F. Hamoda. 1991. Sewage-related impact on Kuwait’s marine environment: A case study. Water
Science and Technology. 23: 181-189.
Andronov, V.N. 1972. Some new species of the genus Paracalanus (Copepoda, Paracalanidae). Zoologicheskii Zhurnal. 51(1): 139-141.
Andronov, V.N. 1977. Paracalanus tropicus sp.n. (Copepoda, Paracalanidae) from the South East Atlantic. Zoologicheskiy Zhurnal. 56:
154-156.
Andryszak B.L. 1986. Upogebia affinis (Say): its postlarval stage described from Louisiana plankton, with a comparison to postlarvae of
other species within the genus and notes on its distribution. J. Crust.Biol. 6(2): 214-226.
Angel M.V. 1999. Ostracoda. In: South Atlantic Zooplankton. Edited by D. Boltovskoy, Backhuys Publishers, Leiden, The Netherlands,
pp. 815-868.
Angel, M.V., Blachowiak-Samolyk K. and Chavtur V.G. 2008. Atlas of Atlantic Planktonic Ostracods. Available from http://www.nhm.
ac.uk/research-curation/research/projects/atlantic-ostracods/index.html.
Appeltans, W.; P. Bouchet; G.A. Boxshall; K. Fauchald; D.P. Gordon; B.W. Hoeksema; G.C.B. Poore; R.W.M. van Soest; S. Stöhr; T.C. Walter;
and M.J. Costello. (eds) 2010. World Register of Marine Species. Availabie from: http://www.marinespecies.org/index.php.
Arnold, C. 2004. The Distribution and Ecology of Tintinnids, In Coastal Waters Around South-East Anglessey, North Wales. University of
Wales, Bangor. Available from: http://216.239.39.104/search?q=cache:toDn3d7Y4jkJ:www.sos.bangor.ac.uk/marine/mb/
O3B32_Field_Course_II/Arnold_03.htm+food+of+tintinnidsandhl=enandie=UTF-8.
Basaham, A.S. and S.S. Al-Lihaibi. 1993. Trace elements in sediments of the western Gulf. Mar. Pollution Bull. 27: 103-107.
Barnett B.M., R.F. Hartwick, and N.E. Milward. 1984. Phyllosoma and Nisto stage of the Morton Bay Bug, Thenus orientalis (Lund)
(Crustacea: Decapoda: Scyllaridae), from shelf waters of the Great Barrier Reef. Aust. J. Mar. Freshw. Res. 35: 143-152.
Bayly, I.A.E. and G.H. Arnott. 1969. A new centropagid genus (Copepoda: Calanoida) from Australian estuarine waters. Australian J.
Mar. Freshw. Res. 20:189-198.
Boltovskoy, D. (editor). 1999. South Atlantic Zooplankton. Volume I & II, Backhuys Publishers, Leiden, The Netherlands, 1705 pp.
Boltovskoy, D., N.R. Swanberg, and S.M. Vivequin. 1995. Tintinnids and other microplankton from the Greenland Sea: abundance and
distribution in the Marginal Ice Zone (May-June 1989). Marine Ecology. 16(2): 1-15.

Boschma, H. 1956. Ellobiopsidae. Zooplankton. Fiches d’identification du zooplancton, ICES, Conseil Perm. Intern. Explor. Mer., Sheet
65, pp. 1-4.
Bouillon, J. 1999. Hydromedusae. In: South Atlantic Zooplankton. Edited by D. Boltovskoy, Backhuys Publishers, Leiden, The
Netherlands, pp. 385-465.
Bourdillon-Casanova, L. 1960. Le meroplankton du Golfe de Marseille: Les larves de Crustaces Decapodes. Recueil des Travaux de la
Staion Marine d’Endoume. 30(18): l-286.
Boxshall, G. A. and S. H. Halsey, 2004. An Introduction to Copepod Diversity. The Ray Society Series. Ray Society, London, 966 pp.
Bradford, J.M. 1976. Partial revision of the Acartia subgenus Acartiura (Copepoda: Calanoida: Acatriidae). New Zealand J. Mar. Freshw.
Res. 10: 159-202.
Bradford J.M. and J.B. Jillet. 1974. A revision of generic definitions in the Calanidae (Copepoda, Calanoida). Crustaceana, 27: 5-16.
Bradford, J.M., L. Haakonsen, and J.B. Jillett. 1983. The marine fauna of New Zealand: pelagic calanoid copepods: familiesEuchaetidae,
Phaennidae, Scolesithricidae, Diaxididae and Tharybidae. Mem. New Zealand Oceanogr. Inst. 90: 1-150.
Bradford-Grieve, J.M. 1994. The marine fauna of New Zealand: Pelagic Calanoid Copepoda: Families Megacalanidae, Calanidae,
Paracalanidae, Mecynoceridae, Eucalanidae, Spinocalanidae, Clausocalanidae. Mem. New Zealand Oceanogr. Inst. 102: 1-160.
Bradford-Grieve, J.M. 1999. The marine fauna of New Zealand: Pelagic Calanoid Copepoda: Bathypontiidae, Arietellidae, Augaptilidae,
Heterorhabdidae, Lucicutiidae, Metridinidae, Phyllopodidae, Centropagidae, Pseudodiaptomidae, Temoridae, Candaciidae,
Pontellidae, Sulcanidae, Acartiidae, Tortanidae. Mem. New Zealand National Inst. of Water and Atmospheric Res. (NIWA)
111: 1-268.
Bradford-Grieve, J.M.; E.L. Markhaseva; C.E.F. Rocha; and B. Abiahy. 1999. Copepoda. In: South Atlantic Zooplankton. Edited by D.
Boltovskoy, Backhuys Publishers, Leiden, The Netherlands, pp. 869-1098.
Brodsky, K.A. 1950. Calanoida of the far eastern seas and polar basin of the U.S.S.R. In: Opredeliteli po Faune SSSR, Akad. Nauk SSSR,
Moskva-Leningrad 35: 1-442.
Brodsky, K.A.; N.V. Vyshkvartseva; M.S. Kos; and E.L. Markhaseva. 1983. Copepods (Copepoda: Calanoida) of the USSR Seas and
contiguous waters. In: Opredeliteli po faune SSSR, Acad. Nauk SSSR, Moskva-Leningrad. 135: 1-358.
Brooks, W.K. 1882. Lucifer: A Study in Morphology. Phil. Trans. R. Soc., London, CLXXIII, pp. 57-138.
Brusca, R.C. and G.J. Brusca. 1990. Invertebrates. Sinauer Associates, Inc., Sunderland, MA, 922 pp.
Campbell, A.S. 1926. On Tintinnopsis neriticus sp. nov. from San Francisco Bay. Univ. Calif. Publ. Zool. 29: 237-239.
Casanova, J.P. 1999. Chaetognatha. In: South Atlantic Zooplankton. Edited by D. Boltovskoy, Backhuys Publishers, Leiden, The
Netherlands, pp.1353-1374.
Chen, Q.C. and S.Z. Zhang. 1965. The planktonic copepods of the Yellow Sea and the East China Sea. I. Calanoida, Studia Marina Sinica
7: 20–131.
Chen, Q.C., S.Z. Zhang, and C.S. Zhu. 1974. The planktonic copepods of the Yellow Sea and the East China Sea. II. Cyclopoida and
Harpacticoida. Studia Marina Sinica 9: 27–76.
Cheung, T.S. 1963. The natural history of the commercial species of Hong Kong Penaeidae (Crustacea, Decapoda). Ann.Mag.nat.Hist.
13(6): 401–33
Conway, D.V.P.; R.G. White; J. Hugues-Dit-Ciles; C.P. Gallienne; and D.B. Robins. 2003. Guide to the coastal and surface plankton of the
southwestern Indian Ocean, Occasional Publication of the Marine Biological Association of the United Kingdom, Plymouth,
UK (15), 345 pp.
Costa, R.C.; A. Fransozo; G.A.S. Melo; and F.A.M. Freire. 2003. An illustrated key for dendrobranchiata shrimps from the northern
coast of São Paulo State, Brazil. Biota Neotropica 3(1). Available from: http://www.biotaneotropica.org.br/v3n1/pt/
abstract?identification-key+BN01503012003.
Dahl, M. 1912. Die Copepoden der Plankton-Expedition I. Die Corycaeinen. Mit Berucksichtigung aller bekannten Arten. Ergebnisse
der Plankton-Expedition, Kiel and Leipzig, 2(G)f(1): 1-136.
Dales, R.P. 1957. Pelagic polychaets of the Pacific Ocean. Bulletin of the Scripps Institute of Oceanography. 7(2): 99-168.
Deevey, G.B. 1968. Pelagic ostracods of the Sargasso Sea of Bermuda. Peabody Museum Bull. 26: 116-118.
Dolan, J.R. and. C.L. Gallegos. 2001. Estuarine diversity of tintinnids (planktonic ciliates). Journal of Plankton Research. 23: 1009-1027.
Doncaster, L. 1902. Chaetognatha, with a note on the variation and distribution of the group. Fauna Maldive and Laccadive Archipel.
1: 209-218.
Dos Santos, A. and J.A. Lindley. 2001. Crustacea Decapoda: larvae. II. Dendrobranchiata (Aristeidae, Benthesicymidae, Penaeidae,
Solenoceridae, Sicyonidae, Sergestidae and Luciferidae). Fishes d’Identification du Zooplancton. 186: 1-9.
Dos Santos, A. and J.I. Gonzales-Cordillo. 2004. Illustrated keys for identification of the Pleocyemata (Crustacea: Decapoda) zoeal
stages, from the coastal region of south-western Europe. J.Mar.Biol. Ass. UK 84: 205-227.
Emig, C.C. 1982. The biology of Phoronida. Adv. Mar. Biol. 19: 1-89.
Esnal, G.B. 1999. Appendicularia. In: South Atlantic Zooplankton. Edited by D. Boltovskoy, Backhuys Publishers, Leiden, The Netherlands,
pp.1375-1399.
Esnal, G.B. and M.C. Daponte. 1999. Salpida. In: South Atlantic Zooplankton. Edited by D. Boltovskoy, Backhuys Publishers, Leiden, The
Netherlands, pp.1423-1444.
Farfante, P.I. and B. Kensley. 1997. Penaeoid and Sergestoid shrimps and prawns of the world. Keys and diagnoses for the families and
genera. Memoires du Museum National, d’histoire Naturelle, 175: 1-233.

Fazal-Ur-Rehman, 1974. Nouvelle description of the female and redescription of the male Pontella investigatoris (Sewell, 1912) from
inshore waters of Karachi. Pakist. J. Zool. 6(1-2): 105-110.
Fenaux, R. 1967. Les appendiculaires des mers d’Europe et du Bassin Méditerranéen. Faune de l’Europe et du Bassin Méditerranéen,
2. Masson et Cie, Paris, 116 pp.
Fernandes, L.F. 2004. Tintininos (Ciliophora, Tintinnina) de agues subtropicas na regiao Sueste-Sul do Brasil. I. Familias Codonellidae,
Codonellopsidae, Coxliellidae, Cyttarocilidae, Epyplocilidae, Petalotrichidae, Ptychocylidae, Tintinnididae e Undellidae.
Revista Brasiliera de Zoologia. 21(3): 521-576.
Fernandez-Alamo, M.A. and E.V. Thuesen. 1999. Polychaeta. In: South Atlantic Zooplankton. Edited by D. Boltovskoy, Backhuys
Publishers, Leiden, The Netherlands, pp. 595-619.
Fielder D.R. and J.G. Greenwood. 1986. Zoeal larvae of Macrophthalmus setosus H. Milne-Edwards, 1852 and M. punctulatus Miers,
1884 (Decapoda, Ocypodidae). Mem. Qd. Mus. 22(2): 155-164.
Fincham, A.A. and D.I. Williamson. 1978. Decapoda, Larvae, VI. Caridea, Families: Palaemonidae and Processidae. Fich. Ident.
Zooplancton. 159/160: 1-8.
Fleminger, A. 1973. Pattern, number, variability and taxonomic significance of the integumental organs (sensilla and glandular pores)
in the genus Eucalanus (Copepoda: Calanoida). Fish. Bull. 71: 965-1010.
Frost, B. and A. Fleminger. 1968. A revision of the genus Clausocalanus (Copepoda: Calanoida) with remarks on distribution patterns
in diagnostic characters. Bull. Scripps Inst. Oceanogr. 12: 1-235.
Geletin, Yu.V. 1976. The ontogenetic abdomen formation in copepods of genera Eucalanus and Rhincalanus (Calanoida, Eucalanidae)
and new system of these copepods. Issledovaniya Fauny Morei. 18(26): 75-93.
Gershwin, L. and A.G. Collins. 2002. A preliminary phylogeny of Pelagiidae (Cnidaria, Scyphozoa), with new observations of Chrysaora
colorata comb. nov. Journal of Natural History. 36: 127-148.
Giesbrecht, W. 1892. Systematik und Faunistik der pelagishen Copepoden des Golfes von Neapel und der angrenzenden Meers-
abschnitte. In: Fauna e Flora del Golfo di Napoli, pp. 1-831.
Gotthard, R. and R.R. Seapy. 1999. Heteropoda. In: South Atlantic Zooplankton. Edited by D. Boltovskoy, Backhuys Publishers, Leiden,
The Netherlands, pp. 621-647.
Grabe, S.A. 1989. Some aspects of the biology of Rhopalophthalmus tattersallae Pillai, 1961 (Crustacea, Mysidacea) and extension of
range into the Khor Al Sabiya, Kuwait (Arabian Gulf). Proceedings of the Biological Society of Washington. 102(3): 726-731.
Grabe, S.A.; W.W. Price; E.A.A. Abdulqader; and R.W.J. Heard. 2004. Shallow water Mysida (Crustacea: Mysidacea) of Bahrain (Arabian
Gulf): species composition, abundance and life history characteristics of selected species. J. Nat. Hist. 38 (18): 2315-2329.
Greenwood, J.G. 1978. Calanoid copepods of Moreton Bay (Queensland). III. Families Temoridae to Tortanidae, excluding Pontellidae.
Proc. R. Soc. Qd. 89: 1-21.
Greenwood, J.G. 1979. Calanoid copepods of Moreton Bay (Queensland). IV. Family Pontellidae. Proc. R. Soc. Qd. 90: 93-111.
Grice, G.D. 1960. Copepods of the genus Oithona from the Gulf of Mexico. Bull. Mar. Sci. Gulf Caribb. 10(4): 485-490.
Grice, G.D. 1963. A revision of the genus Candacia (Copepoda: Calanoida) with an annotated list of the species and a key for their
identification. Zool. Mededelingen. 38: 171-194.
Guerao G. and P. Abello. 1999. The first zoeal stage of Parthenope macrochelos (Herbst, 1790) hatched in the laboratory (Crustacea,
Brachyura, Parthenopidae). Sci. Mar. 63(1): 9-14.
Gurney, R. 1931. British Freshwater Copepoda. London: Ray Soc. Publ., 239 pp.
Gurney, R. 1936. Larvae of decapod Crustacea. Part I: Stenopodidea. Discovery Reports. 12: 379-392.
Gurney, R. 1938. Larvae of decapod Crustacea. Part V. Nephropsidea and Thalassinidea. Discovery Reports. 17: 291-344.
Habib-ul-Hassan, 1982. The larval development of Penaeus semisulcatus de Haan, 1850 (Decapoda, Penaeidae) reared in the laboratory.
J. Plank. Res. 4(1): 1-17.
Habib-ul-Hassan, 1984. Larval development of Parapenaeopsis stylifera (H. Milne-Edwards) (Decapoda, Penaeidae). ICES J. Mar. Sci. 41:
293-303
Hada, Y. 1932. Report of the Biological Survey of Mutsu Bay. 24. The Pelagic Ciliata, Suborder Tintinnoinea. Sci.Rep.Tohoku Imp.Univ.
Ser.IV (Biol.) VII(4): 553-573.
Hada, Y. 1937. The fauna of Akkeshi Bay IV. The Pelagic Ciliata. J. Fac. Sci. Hokkaido Imp. Univ., ser. 4, Zool. 3: 143-216.
Hashizume, K. 1999. Larval development of seven species of Lucifer (Dendrobranchiata, Sergestoidea), with a key for identification
of their larval forms. In: Crustaceans and Biodiversity Crisis: Proceedings of the Fourth International Crustacean Congress,
Amsterdam, The Netherlands, July 20-24, 1: 753-779.
Hashmi S.S. 1969. Studies on larval Ocypodidae (Macrophthalmus) hatched in the laboratory (Decapoda: Crustacea). Pak. J. Sci. Res.
21: 50-56.
Hayashi, K.I. and S. Tsumura. 1981. Revision of the Japanese Luciferinae (Decapoda, Penaeidae, Sergestidae). Bull. Jap. Soc. Sci. Fish.
47: 1437-1441.
Heron, G.A. and J.M. Bradford-Grieve. 1995. Pelagic Copepoda: Poecilostomatoida: Oncaeidae. Mem. New Zealand Oceanogr. Inst.
104: 1-57.
Holthuis, L.B. 1980. Shrimps and Prawns of the World: An Annotated Catalogue of Species of Interest to Fisheries. FAO Fisheries
Synopsis 1(125). FAO. Rome, 271 pp.
Hooge, M.D. and S. Tyler. 2003. Acoels (Platyhelminthes, Acoela) from the Atlantic coast of North America. Meiofauna Marina. 12: 1-30.

Hunter, J.R. 1982. The physical oceanography of the Arabian Gulf: A review and theoretical interpretation of previous observations.
Proceedings of the First Arabian Gulf Conference on Environment and Pollution, Kuwait University, February 7-9, pp. 1-23.
Huys, R. and G.A. Boxshall. 1991. Copepod Evolution. Ray Society, London 159:468.
Huys, R. 1996. Superornatiremidae fam. nov. (Copepoda: Harpacticoida): An enigmatic family from North Atlantic anchihaline caves.
Scientia Marina 60(4): 497-542.
Iles, E.J. 1961. The appendages of the Halocyprididae. Discovery Reports 31:299-326.
Illig, G. 1927. Die Sergestiden der Deutschen Tiefsee-Expedition. Wiss. Ergebn. Detschen Tiefsee-Expedition. Valdivia 1898-1899. 23(7):
277-354.
Ingle R. 1992. Larval Stages of Northeastern Atlantic Crabs. An Illustrated Key. Chapman and Hall Identifcation Guide. London-New
York-Tokyo-Melbourne-Madras, 363 pp.
Inamori, Y. 1996-2010. The World of Protozoa, Rotifera, Nematoda and Oligochaeta [on-line]. National Institute for Environmental
Studies, Japan Environmental Agency 16-2 Onogawa, Tsukuba, Ibaraki 305-0053 Japan. Available from: http://web2.nies.
go.jp/chiiki1/protoz/.
Jackson, C.J.; P.C. Rothlisberg; R.C. Pendrey and M.T. Beamish. 1989. A key to genera of the penaeid larvae and early postlarvae of the
Indo-west Pacific region, with descriptions of the larval development of Atypopenaeus formosus Dall and Metapenaeopsis
palmensis Haswell (Decapoda: Penaeoidea: Penaeidae) reared in the laboratory. Fishery Bull. 87: 703-733.
Jereb, P. and C.F.E. Roper. 2005. Cephalopods of the World - An annotated and illustrated catalogue of cephalopod species known
to date. Vol. 1 Chambered Nautiluses and Sepioids (Nautilidae, Sepiidae, Sepiolidae, Sepiadariidae, Isiosepiidae and
Spirulidae). FAO Species Catalogue for Fishery Purposes. 4(1): 1-262.
Karuppasamy, P.K.; N.G. Menon; K.K.C. Nair; and C.T. Achuthankutty. 2006. Distribution and abundance of pelagic shrimps from the
deep scattering layer of the eastern Arabian Sea. J. Shellfish. Res. Available from: http://findarticles.com/p/articles/mi_
m0QPU/is_3_25/ai_n17134279
Khalaf, T.A. 1991. A new calanoid of the genus Acartia from Khor Abdulla and Khor Al-Zubair waters, Iraq. Marina Mesopotamica 6(1):
80-91.
Khan, M.A. 1976. Luciferinae (Crustacea, Decapoda, Sergestidae) collected from Manora Channel (northern Arabian Sea). Agriculture
Pakistan 27: 113-117
Kensley, B.F. 1971 The family Sergestidae in the waters around southern Africa (Crustacea, Decapoda, Natantia). Annals of the South
African Museum. 57: 215-265.
Kofoid, C.A. and A.S. Campbell. 1929. A conspectus of the marine and freshwater Ciliata belonging to the suborder Tintinnoinea, with
descriptions of new species principally from the Agassiz expedition to the eastern tropical Pacific, 1904-1905. Univ. Calif.
Publ. Zool. 403 pp.
Kofoid, C.A. and A.S. Campbell. 1939. Reports on the scientific results of the expedition to the eastern tropical Pacific in charge of
Alexander Agassiz, by the U.S. Fish Commission steamer “Albatross” from October 1904 to March 1905, Lieut. –commander
L.M. Garrett, U.S.N. commanding XXXVII. The Ciliata: the Tintinnoinea. Bull. Mus. comp. Harv. 84: 1-473.
Konishi, K. 1987. Larval Development of Pachycheles stevensii Stimpson, 1858 (Crustacea:Anomura: Porcellanidae) under Laboratory
Conditions. Journal of Crustacean Biology, 7(3): 481-492.
Kunz, T.J. and A.J. Richardson. 2006. Impacts of climate change on phytoplankton. In: Impacts of climate change on Australian marine
life. Report to the Australian Greenhouse Office, Canberra. Part C, pp. 8–18.
Lalli, C.M. and R.W. Gilmer. 1989. Pelagic snails: the biology of holoplanktonic gastropod mollusks, Stanford University Press, Stanford,
California, 259 pp.
Lalli, C.M. and T.R. Parsons. 2001. Biological Ocenaography An Introduction 2nd Edition. Oxford, Pergamon Press, 406 pp.
Lardner, R.; A.H. Al Rabeh; N. Gunay; R. Khan; H. Hossani; R.M. Reynolds; and W.J. Lehr. 1993. Computation of the residual flow in the
Gulf using the Mt Mitchell data and KFUPM/RI Hydro dynamical models. Marine Pollution Bulletin. 27: 61-70.
Laval-Peuto, M. 1977. Reconstruction d’une lorica de forme Coxliella par le trophonte nu de Favella ehrenbergii (Ciliata, Tintinnina).
C.R.Acad.Sc.Paris. 284: 547-550.
Lebour, M.V. 1916. Stages in the life history of Calanus finnzarchicus (Gunnerus), experimentally reared by Mr. L.R. Crawshay in the
Plymouth Laboratory. J. mar. biol. Ass. U.K. 11(1): 1-17.
Lee, J.J., G.F. Leedale, and P. Bradbury. 2000. An Illustrated Guide to the Protozoa. 2nd Edition, Vol.1. Society of Protozoologists,
Lawrence, KS, USA, 689 pp.
Literathy, P.; L.N. Ali; M.A. Zarba; and K. Puskas. 1986. Effects of sewage discharge on intertidal ecosystems in Kuwait. Water Science
and Technology. 18(11): 35-45.
Lynn, D.H. and E.B. Small. 2000. Phylum Ciliophora. In An Illustrated Guide to the Protozoa, Second Edition, vol. 1. Edited by J.J. Lee, G.F.
Leedale and P. Bradbury. Society of Protozoologists, Lawrence, Kansas, U.S.A., pp. 371-656.
Madhupratab, M. and P. Haridas. 1978. Archidiaptomus aroorus, a new genus and species of Copepoda (Calanoida, Pseudodiaptomus)
from Cochin backwaters, India. Crustaceana. 35: 253-258.
Marine Species Identification Portal (2000-2010). Available from: http://species-identification.org/index.php.
Marshall, S.M. 1969. Protozoa. Order Tintinnida. Conseil International pour l’Exploration de la Mer, Fiches d’Identification de
Zooplancton, pp. 117-127.
Mauchline, J. 1998. The Biology of Calanoid Copepods. Academic Press, San-Diego, 138 pp.

Menon P.G. and D.I. Williamson. 1971. Decapod Crustacea from the international Indian Ocean Expedition. The species of Thalassocaris
(Caridea) and their larvae. J. Zool. London. 165: 27-51.
Mianzan, H.W. 1999. Ctenophora. In: South Atlantic Zooplankton. Edited by D. Boltovskoy, Backhuys Publishers, Leiden, The
Netherlands, pp. 561-573.
Mianzan, H.W. and P.V.S. Cornelius. 1999. Cubomedusae and Scyphomedusae. In: South Atlantic Zooplankton. Edited by D. Boltovskoy,
Backhuys Publishers, Leiden, The Netherlands, pp. 513-559.
Micro*scope. 2010. The Internet Site of Marine Biological Laboratory. The Astrobiology Institute, Woods Hole, Massachusetts, USA.
Available from: http://starcentral.mbl.edu/microscope/portal.php?pagetitle=index.
Michel, H.B.; M. Behbehani; D. Herring; and M. Arar. 1986a. Zooplankton of the western Arabian Gulf south of Kuwait waters. Kuwait
Bulletin of Marine Science. 8: 1-36.
Michel, H.B.; M. Behbehani; D. Herring; M. Arar; M. Shoushani; and T. Brakoniecki. 1986b. Zooplankton diversity, distribution and
abundance in Kuwait waters. Kuwait Bulletin of Marine Science. 8: 37-105.
Mori, T. 1937. The Pelagic Copepoda from the Neighbouring Waters of Japan. Tokyo: The Soyo Company, 150 pp.
Motoda, S. 1959. Devices of simple plankton apparatus. Mem. Fac. Fish. Hokkaido Univ. 7: 73-94.
Muller, C.W. 1906. Ostracoda. Wissenshaftliche Ergebniss der Deatschen Tiefsee-Expedition “Valdivia” 1898-1899. Bd.8. L.2, pp. 1-129.
Muus, B.J. 1953. Polychaeta. Families: Aphroditidae, Phyllodocidae and Alciopidae. Zooplankton, Sheet 52. Conseil International Pour
L’Exploration de la Mer, pp. 1-5.
Naidu, W.D. 1983. Tintinnida (Protozoa: ciliate) – a vital link in the estuarine food web. Mahasagar – Bulletin of the National Institute of
Oceanography. 16: 403-407.
Ngoc-Ho N. 1981. A taxonomic study of the larvae of four thalassinid species (Decapoda, Thalassinidea) from the Gulf of Mexico. Bull.
Br. Mus. Nat. Hist. (Zool.) 40(5): 237-27.
Nicholson, H.A. 1880. A Manual of Zoology for the Use of Students with a General Introduction on the Principles of Zoology. William
Blackwood and Sons, 698 pp.
Nishida, S. 1985. Taxonomy and distribution of the Family Oithonidae (Copepoda, Cyclopoida) in the Pacific and Indian Oceans. Bull.
Ocean Res. Inst., Univ. Tokyo. 20: 167.
Omori, M. 1992. Occurrence of two species of Lucifer (Dendrobranchiata: Sergestoidea: Luciferidae) off the Pacific coast of America. J.
Crust. Biol. 12(1): 104-110.
Onbe, T. 1999. Ctenopoda and Onychopoda (=Cladocera). In: South Atlantic Zooplankton. Edited by D. Boltovskoy, Backhuys
Publishers, Leiden, The Netherlands, pp. 797-813.
Othman, B.H.R. and T. Toda. 2006. Pontellid copepods from Singapore. Coastal Marine Science 30(1): 305-319.
Owre, H.B. and M. Foyo. 1967. Copepods of the Florida Current. Fauna Caribaea, number 1, Crustacea, part 1: Copepoda, Institute of
Marine Science, University of Miami, 137 pp.
Park, T.S. 1995. Taxonomy and distribution of the marine Calanoid Copepod Family Euchaetidae. Bull. Scripps Inst. Oceanogr. 29: 203.
Panampunnayil, S.U. and A. Biju. 2006. Four new species of the genus Rhopalophthalmus (Mysidacea: Crustacea) from the North West
coast of India. J. Nat. Hist. 40: 1389.
Pasupathi, K. and T. Kannupandi. 1988. The complete larval development of the mangrove ocypodid crab Macrophthalmus depressus
Ruppel, 1830 (Brachyura, Macrophthalminae) reared in the laboratory. J. Nat. Hist. 22: 1533-1544.
Paulmier, G. 1997. Tintinnides (Ciliophora, Oligotrichida, Tintinnina) de’l Atlantique boreal, de’l Ocean Indien et de Queloques mers
adjacents: Mediterranee, mer Caribe, mer Rouge. Inventarie et distribution. Observations basees sur les loricas. Station
IFREMER. DRV/RH/97-17, 126 pp.
Pennington, J.T. and S.A. Stricker. 2002. Phylum Brachiopoda. In: Atlas of Marine Invertebrate Larvae. Edited by C.M. Young. Academic
Press, pp. 441-461.
Perez-Farfante, I. and B. Kensley. 1997. Penaeoid and sergestoid shrimps and prawns of the worlds. Keys and diagnoses for the families
and genera. Paris, Memoires du Muzeim National d’Histoire Naturelle, 233 pp.
Pike, R.B. and D.I. Williamson. 1958. Crustacea, Decapoda: larvae, XI. Paguridea, Coenobitidea, Dromiidea, and Homolidea. Fich. Ident.
Zooplancton. 81: 1-9.
Pike, R.B. and D.I. Williamson. 1960. Larvae of decapod crustacea of the families Diogenidae and Paguridae from the Bay of Naples.
Pubbl. Staz. Zool. Napoli. 31(3): 493-543.
Pike, R.B. and D.I. Williamson. 1972. Crustacea, Decapoda: larvae. X. Galatheidea. Fich. Ident. Zooplancton. 139: 1-5.
Pohle G., Fernando L.M. Montellato, M.L. Fransozo N. and Fransozo, A. 1999. Larval Decapoda (Brachyura). In: South Atlantic
Zooplankton. Edited by D. Boltovskoy, Backhuys Publishers, Leiden, The Netherlands, pp.1281-1351.
Poulsen, E.M. 1962. Ostracoda - Myodocopa. Part 1. Cypridiniformes - Cypridinidae, Dana - Report 57, Copenhagen, pp. 255-258.
Price, W.W. 1982. Key to the shallow water Mysidacea of the Texas coast with notes on their ecology. Hydrobiologia 93: 9-21.
Prusova, I., F. Al-Yamani, and H. Al Mutairi. 2001. Subeucalanus flemingeri sp. n. from the Arabian Gulf (Copepoda: Eucalanidae).
Zoosystematica Rossica. 10(1): 47-54.
Raymont, J.E.G. 1983. Plankton and Productivity in the Оceans. Vol. 2. Zooplankton. Pergamon Press, Oxford, pp. 166-228.
Razouls, C. and F. de Bovée. 1998. Diversity and geographic distribution of pelagic Copepoda. 3. An overview and initial interpretation.
Ann. Inst. Océanogr. 74(2): 139-200.

Razouls C., F. de Bovee, and N. Desreumaux. 2005-2010. Diversity and Geographical Distribution of Pelagic Copepoda. Available from:
http://copepodes.obs-banyuls.fr.
Reynolds, R.M. 1993. Physical oceanography of the Gulf Strait of Hormuz and the Gulf of Oman. Results from the Mt. Mitchell
expedition. Marine Pollut. Bull. 27: 35-59.
Reynolds, R.M. 2002. Oceanography. In: Khan, N.Y., Munawar, M. and Price, A.R.G. (eds), The Gulf Ecosystem: Health and Sustainability.
Backhuys Publishers. Leiden, pp. 53–64.
Richardson, A.J. and T.J. Kunz. 2006. Impacts of climate change on zooplankton. In: Impacts of climate change on Australian marine
life. Report to the Australian Greenhouse Office, Canberra. Part C, pp. 19–26.
Roberts P.E. 1971. Zoea larvae of Pagurus campbelli Filhol, 1885, from Perseverance Harbour, Campbell Island (Decapoda, Paguridae).
Journ. Roy. Soc. New Zealand. 1(3/4): 187-196.
Rose, M. 1933. Copepodes Pelagiques. Faune de France, Paris, 374 pp.
Ruppert, E.E. 1997. Cephalochordata (Acrania), In Harrison FW, Ruppert E.E (Eds): Microscopic anatomy of Invertebrates, vol. 15. Wiley-
Liss, New York, pp. 349-504.
Russell, F.S. 1955. Hydromedusae. Families: Pandeidae and Tiarannidae. Zooplankton. Sheet 54, Conseil International Pour L’Exploration
de la Mer, pp. 1-6.
Russell, F.S. 1963. Hydromedusae. Families: Phialuciidae, Eirenidae, Eutimidae. Zooplankton Sheet 102, Conseil International Pour
L’Exploration de la Mer, pp. 1-4.
Russell, F.S. 1970. Hydromedusae. Family: Aequoreidae. Zooplankton, Sheet 128. Conseil International Pour L’Exploration de la Mer,
pp. 1-4.
Russell, F.S. 1977. Hydromedusae: Families Clavidae and Hydractiniidae, Fich. Ident. Zooplancton. 154: 1-4.
Sazhina, L.I. 1987. Reproduction, development, productivity of marine Copepoda. Naukova dumka, Kiev, pp. 1-156.
Sandifer P.A. 1973. Mud shrimp (Callianassa) larvae from Virginia plankton (Crustacea, Decapoda, Callianassidae). Chesapeake Science.
14(3):149-159.
Scott T. 1909. On new and rare Crustacea from Scottish waters. Ann. Mag. nat. Hist. 8(4): 31-36.
Sewell R.B.S. 1914. Notes on the surface Copepoda of the Gulf of Mannar. Spolia zeylan. 9: 191-262.
Sewell R.B.S. 1932. The Copepoda of Indian Seas. Calanoida. Mem. Indian Mus. 10: 223-407.
Sheppard, C. 1993. Physical environment of the Gulf relevant to marine pollution: An overview. Mar. Pollut. Bull. 27: 2-8.
Shunbo, F.; P. Literathy; M.A. Ali; O. Samhan; and P.G. Jacob. 1986. Baseline studies of Oil and Non-oil Pollutants in the Marine
Environment of Kuwait. Kuwait Institute for Scientific Research. Report No. KISR 1324, Kuwait.
Shuvalov, V.S. 1980. Copepoda crustaceans. Cyclopoida of the family Oithonidae of the world oceans. (Veslonogie rachki-tsiklopoidy
semeistva Oithonidae mirovogo okeana). In: Opredeliteli po Faune SSSR, Nauka, Leningrad, pp. 1-197.
Shy J.Y., J.J. Chang, and H.T. Lai. 2005. Complete larval development of Palaemon serrifer (Stimpson, 1860) (Crustacea, Decapoda,
Palaemonidae) reared in laboratory. J. Fish. Soc. Taiwan. 32(1): 79-86.
Silas, E.G. and P.P. Pillai. 1973. The calanoid copepod family Pontellidae from the Indian Ocean. J. Mar. Biol. Ass. India. 15: 771-858.
Skryabin, V.A. and F.Y. Al-Yamani. 2006. New species of genera Tintinnopsis, Metacylis and Eutintinnus (Ciliophora, Spirotrichea,
Tintinnida) from Kuwait’s Waters of the Arabian Gulf. Russian Journal of Marine Biology. 32(5): 302-308.
Skryabin, V.A. and F.Y. Al-Yamani. 2007. New species of genera Leprotintinnus and Luminella (Ciliophora, Spirotrichea, Tintinnida) from
Kuwait’s Waters of the Arabian Gulf. Kuwait Journal of Science and Engineering (IJOO) 34(1A): 79-90.
Skryabin, V.A. and F.Y. Al-Yamani. 2007a. Composition, distribution and seasonal changes of tintinnids in the northwestern Arabian
Gulf around the Bubiyan Island. International Journal of Oceans and Oceanography (IJOO) 2(1): 103-115.
Sleigh, M.A. 1973. The Biology of Protozoa. London: Edward Arnold, 315 pp.
Smith, J.P.S. and S. Tyler. 1985. The acoel turbellarians: kingpins of metazoan evolution or a specialized offshoot? In: The Origins and
Relationships of Lower Invertebrates. Edited by C.C. Morri, J.D. George, R. Gibson, and H.M. Platt. Oxford: Oxford University
Press, pp 123-142.
Smithsonian National Museum of Natural History, Internet Site of the Department of Botany. 2009. Available from: http://botany.
si.edu/references/dinoflag/taxa.htm .
Steuer, A. 1915. Revision der Gattung Acartia Dana. Zool. Anz. 45(9): 392-397.
Steuer, A. 1923. Bausteine zu einer Monographie der Copepodengattung Acartia. Arb. zool. Inst. Univ. Innsbrück. 1(5): 91-144.
Tanaka, O. 1957. On Copepoda of the family Corycaeidae in Japanese waters. J. Faculty of Agriculture, Kyushu University. 11(1): 77-97.
Tanaka, O. 1964. The pelagic copepods of the Izu region, Middle Japan. Systematik account XII. Families Arietellidae, Pseudocyclopidae,
Candaciidae and Pontellidae. Publs Seto mar. biol. Lab. 12(3): 231-271.
Tirmizi, N. M. and M. Yaqoob. 1979. Larval development of Pachycheles tomentosus Henderson (Anomura, Porcellanidae) with
descriptive remarks on the adults from Karachi waters (northern Arabian Sea). Pakistan Journal of Zoology. 11: 29–42.
Todd, C.D., M.S. Laverack, and G.A. Boxshall. 1996. Coastal marine zooplankton: a practical manual for students. Cambridge University
Press, 106 pp.
Totton, A.K. 1954. Siphonophora of the Indian Ocean together with systematic and biological notes on related specimens from other
oceans. Discovery Rep. 27: 162.

Totton, A.K. 1965. A Synopsis of the Siphonophora. British Museum of Natural History, London, 230 pp.
Uchida, T. and Y. Sugiura. 1978. On the polyp of the scyphomedusa, Sanderia malayensis and its reproduction. Journal of the Faculty
of Science. Hokkaido University, series VI, Zoology 21: 279-286.
Ueda, H. and A.C. Bucklin. 2006. Acartia (Odontacartia) ohtsukai, a new brackish-water calanoid copepod from Ariake Bay, Japan, with
redescription of the closely related A. pacifica from the Seto Inland Sea. Hydrobiologia. 560: 71-91.
Van der Spoel, S. and J.R. Dadon. 1999. Pteropoda. In: South Atlantic Zooplankton. Edited by D. Boltovskoy, Backhuys Publishers,
Leiden, The Netherlands, pp. 649-706.
Vervoort, W. 1963. Pelagic copepoda Part 1. Copepoda Calanoida of the families Calanidae up to and including Euchaetidae. Atlantide
Rep. 7: 77-194.
Vervoort, W. 1964. Notes on two Pacific species of Centropages (Copepoda, Calanoida) C. australiensis Fairbridge, 1944, and C.
aucklandicus Kramer, 1985. Crustaceana. 7: 293-311.
Walter, T.C. 1986. New and poorly known Indo-Pacific species of Pseudodiptomus (Copepoda: Calanoida), with a key to the species
groups. Journal of Plankton Research. 8(1): 129-168.
Walter, T.C. 1989. Review of the new world species of Pseudodiptomus (Copepoda: Calanoida), with a key to the species. Bull. of Marine
Science. 45(3): 590-628.
Walter, T.C. 1998. A redescription of Pseudodiaptomus salinus (Giesbrecht, 1896) and a new species from Arabian Sea (Copepoda,
Calanoida, Pseudodiaptomidae). Journal of Marine Systems, Spec. 15(1-4): 451-456.
Wear, R.G. 1965. Larvae of Petrocheles spinosus Miers, 1876 (Crustacea, Decapoda, Anomura) with keys to New Zealand porcellanid
larvae. Transactions of the Royal Society of New Zealand. 5: 147–168.
Wear R.G. 1976. Larva of the commensal shrimp Periclimenes (Periclimenes) soror Nobili, 1904 (Crustacea: Decapoda: Pontoninae) from
Fiji. N.Z.J. Mar. Fresh. Res. 10(3): 527-32.
Wells, J.B.J. 1970. Copepoda. I. Sub-order Harpacticoida. Fiches d’Identif. Zoopl., Cons. Perm. Int. Explor. Mer. 133: 1-7.
Wickstead, J. 1967. Chordata, Sub-phylum Acrania, Family Branchiostomatidae. In: ICES Identification Leaflets for Plankton, Leaflet N
111, ICES, Copenhagen, Denmark, pp. 1-4.
Williamson, D.I. 1957a. Crustacea, Decapoda: larvae, I. General. Fich. Ident. Zooplancton. 67: 1-7.
Williamson, D.I. 1957b. Crustacea, Decapoda: larvae, V. Caridea, Family Hippolytidae. Fich. Ident. Zooplancton. 68: 1-5.
Williamson, D.I. 1962. Crustacea, Decapoda: larvae, III. Caridea, Families Oplophoridae, Nematocarcinidae and Pasiphaeidae. Fich.
Ident. Zooplancton. 92: 1-5.
Williamson, D.I. 1967. Crustacea, Decapoda: larvae, IV. Caridea, Families: Pandalidae and Alpheidae. Fich. Ident. Zooplancton. 109: 1-5.
Williamson, D.I. 1976. Larvae of Stenopodidea (Crustacea Decapoda) from the Indian Ocean. J. Nat. Hist. 10: 497-509.
Wilson, C.B. 1932. The copepods of the Woods Hole region, Massachusetts. Bull. Smithsonian Inst., U.S. National Museum. 158: 634.
Yaldwin, J.C. and R.G. Wear. 1972. The eastern Australian, burrowing mud-shrimp Laomedia healyi (Crustacea, Macrura, Reptantia,
Laomediidae) with notes on larvae of the genus Laomedia. Australian Zoologist. 17(2): 126-143.
Yang, W.T. 1971. The larval and postlarval development of Parthenope serrata reared in the laboratory and the systematic position of
the Parthenopinae (Crustacea, Brachyura). Biol. Bull. 140: 166-189.
Yang, H.J. 2005 Larval development of Latreutes anoplonyx (Decapoda: Hippolytidae) reared in the laboratory. J. Crust. Biol. 25(3):
462-479.
Yang, H.J. 2007. Larval development of Latreutes acicularis Ortmann (Crustacea: Decapoda: Hippolytidae) reared in the laboratory.
Integrative Biosciences. 11: 79-92.
Yashnov, V.A. 1939. Instruction on methods of plankton collection and its analysis in a field environment. M.: Publ. VNIRO, 1939, 22 pp.
Zingel, P. 2002. Introduction to protozoan plankton ecology [on-line]. Aquatic resources. University of Jyväskylä, Finland. Available
from: http://www.jyu.fi/bio/hyb/temp/Protozoa.htm.

Index of Scientific Names
Acanthephyra sp. Vol.2, 146 Clytemnestra scutellata Vol.2, 125
Acartiidae Vol.2, 80 Clytia discoida Vol.1, 140-141
Acartia Vol.2, 80 Codonellidae Vol.1, 20
Acartia (Acanthacartia) fossae Vol.2, 82-83 Codonellopsidae Vol.1, 56
Acartia (Odontacartia) amboinensis Vol.2, 84-85 Codonellopsis Vol.1, 56
Acartia (Odontacartia) ohtsukai Vol.2, 86-87 Codonellopsis lusitanica Vol.1, 56-57
Acartiella faoensis Vol.2, 88-89 Codonellopsis morchella Vol.1, 58
Actinotrocha Larvae Vol.1, 178-179 Codonellopsis ostenfeldi Vol.1, 59
Aequorea pensilis Vol.1, 128-129 Copepoda Vol.2, 1-7
Aglaura hemistoma Vol.1, 146-147 Copilia Vol.2, 108
Alpheus spp. Vol.2, 149 Copilia mirabilis Vol.2, 108-109
Amphinema rugosum Vol.1, 126-127 Corycaeidae Vol.2, 112
Amphorellopsis Vol.1, 96 Corycaeus Vol.2, 112
Amphorellopsis acuta Vol.1, 96-97 Corycaeus (Dithrichocorycaeus) andrewsi Vol.2, 113
Amphorides Vol.1, 98 Corycaeus (Dithrichocorycaeus) dahli Vol.2, 114-115
Amphorides amphora Vol.1, 98-99 Corycaeus (Dithrichocorycaeus) lubbocki Vol.2, 116
Amphorides quadrilineata Vol.1, 100 Corycaeus (Onychocorycaeus) agilis Vol.2, 117
Appendicularia sicula Vol.2, 174 Corycaeus (Onychocorycaeus) pacificus Vol.2, 118
Ascampbelliedidae Vol.1, 76 Coxliella Vol.1, 64
Ascampbelliella Vol.1, 76 Coxliella annulata Vol.1, 64-65
Ascampbelliella obscura Vol.1, 76-77 Creseis chierchiae Vol.1, 175
Atlanta sp. Vol.1, 172-173 Cunina octonaria Vol.1, 144-145
Cyclopoida Vol.2, 94-95
Balanus amphitrite Vol.1, 182-183 Cyphonautes Larvae Vol.1, 180-181
Beroe sp. Vol.1, 156-157 Cypridina sp. Vol.1, 192-193
Bestiolina Vol.2, 20
Bestiolina arabica Vol.2, 20-21 Dadayiella Vol.1, 101
Brachiopoda Larvae Vol.1, 170-171 Dadayiella cuspis Vol.1, 101
Branchiostoma sp. Vol.2, 180-181 Dadayiella ganymedes Vol.1, 102
Dardanus sp. Vol.2, 161
Calanidae Vol.2, 10 Dicranophoridae Vol.1, 164-165
Calanoida Vol.2, 10 Dictyocystidae Vol.1, 60
Calanopia Vol.2, 58 Dictyocysta Vol.1, 60
Calanopia elliptica Vol.2, 58-59 Dictyocysta duplex Vol.1, 60-61
Calanopia minor Vol.2, 60-61 Diogenes sp. Vol.2, 160
Callianassa sp. Vol.2, 152 Diphyes chamissonis Vol.1, 152-153
Candaciidae Vol.2, 56 Dithrichocorycaeus Vol.2, 113
Candacia Vol.2, 56
Candacia bradyi Vol.2, 57 Ebalia sp. Vol.2, 164
Canthocalanus Vol.2, 12 Ectinosomatidae Vol.2, 120
Canthocalanus pauper Vol.2, 12-13 Eirene viridula Vol.1, 130-131
Catostylus mosaicus Vol.2, 182-183 Ellobiopsis chattoni Vol.1, 122-123
Centropagidae Vol.2, 42 Eucalanidae Vol.2, 29
Centropages Vol.2, 42 Euchaetidae Vol.2, 36
Centropages furcatus Vol.2, 44-45 Euchaeta Vol.2, 36
Centropages orsinii Vol.2, 46-47 Euchaeta concinna Vol.2, 38-39
Centropages tenuiremis Vol.2, 47 Euchaeta rimana Vol.2, 40-41
Cirripedia Vol.1, 182-183 Euterpinidae Vol.2, 124
Clausocalanidae Vol.2, 33 Euterpina Vol.2, 124
Clausocalanus Vol.2, 33 Euterpina acutifrons Vol.2, 124
Clausocalanus minor Vol.2, 34-35 Eutima gegenbauri Vol.1, 132-133
Clytemnestridae Vol.2, 124 Euconchoecia aculeata Vol.1, 190-191
Clytemnestra Vol.2, 125 Eutintinnus Vol.1, 103

Eutintinnus apertus Vol.1, 103 Metacylis jorgenseni Vol.1, 68-69
Eutintinnus conicus Vol.1, 104 Metacylis lucasensis Vol.1, 70-71
Eutintinnus contractus Vol.1, 106-107 Metacylis pithos Vol.1, 72-73
Eutintinnus fraknoi Vol.1, 108 Metacylis sp. Vol.1, 75
Eutintinnus lusus-undae Vol.1, 109 Metacylis tropica Vol.1, 74
Eutintinnus macilentus Vol.1, 110 Metapenaeus ensis Vol.2, 128
Eutintinnus sp. Vol.1, 113 Microsetella Vol.2, 120
Eutintinnus tenue Vol.1, 111 Microsetella sp. Vol.2, 120-121
Eutintinnus turgescens Vol.1, 112 Miraciidae Vol.2, 122
Favella Vol.1, 89 Noctiluca scintillans Vol.1, 124-125

Favella adriatica Vol.1, 89
Favella campanula Vol.1, 90 Obelia spp. Vol.1, 136-137
Favella ehrenbergi Vol.1, 91 Octophialucium funerarium Vol.1, 134-135
Favella panamensis Vol.1, 92-93 Odontacartia Vol.2, 84
Fritillaria pellucida Vol.2, 184-185 Oikopleura dioica Vol.2, 173
Fritillaria sp. Vol.2, 175 Oithonidae Vol.2, 94
Oithona Vol.2, 94-96
Galathea sp. Vol.2, 158 Oithona attenuata Vol.2, 96-97
Oithona brevicornis Vol.2, 98-99
Harpacticoida Vol.2, 120 Oithona nana Vol.2, 98-99
Helicostomella Vol.1, 66 Oithona plumifera Vol.2, 100
Helicostomella longa Vol.1, 66-67 Oithona sp. Vol.2, 102-103
Oncaeidae Vol.2, 104
Ilyoplax frater Vol.2, 168 Oncaea Vol.2, 104
Oncaea clevei Vol.2, 106-107
Labidocera Vol.2, 62 Onychocorycaeus Vol.2, 116
Labidocera acuta Vol.2, 62-65
Labidocera bengalensis Vol.2, 66-67 Pachycheles sp. Vol.2, 159
Labidocera kroyeri Vol.2, 58-69 Paguristes sp. Vol.2, 162
Labidocera minuta Vol.2, 70-71 Pagurus sp. Vol.2, 163
Labidocera sp. Vol.2, 72-73 Palaemon sp. Vol.2, 147
Laomedia sp. Vol.2, 153 Paracalanidae Vol.2, 14-15
Latreutes sp. Vol.2, 150 Paracalanus Vol.2, 22
Leprotintinnus Vol.1, 14 Paracalanus indicus Vol.2, 22-23
Leprotintinnus bubiyanicus Vol.1, 14-15 Paracalanus sp. Vol.2, 24-25
Leprotintinnus elongatus Vol.1, 16-17 Parapenaeopsis stylifera Vol.2, 129
Leprotintinnus nordqvisti Vol.1, 18-19 Parthenope sp. Vol.2, 166
Leucosia sp. Vol.2, 165 Parundella Vol.1, 86
Limacina bulimoides Vol.1, 174 Parundella lohmanni Vol.1, 86
Liriope tetraphyla Vol.1, 148-149 Parvocalanus Vol.2, 26
Lopadorhynchus henseni Vol.1, 166-167 Parvocalanus crassirostris Vol.2, 26-27
Lucifer hanseni Vol.2, 135-137 Parvocalanus elegans Vol.2, 28
Lucifer typus Vol.2, 138 Pelagobia longicirrata Vol.1, 168-169
Luminella Vol.1, 62 Penaeus semisulcatus Vol.2, 130-131
Luminella kuwaitensis Vol.1, 62-63 Penilia avirostris Vol.1, 184-185
Lysmata sp. Vol.2, 151 Periclimenes sp. Vol.2, 148
Petalotrichidae Vol.1, 78
Macrophthalmus sp. Vol.2, 167 Petalotricha Vol.1, 78
Macrosetella Vol.2, 122 Petalotricha ampulla Vol.1, 78-79
Macrosetella gracilis Vol.2, 122-123 Pilidium Larvae Vol.1, 162-163
Metacylididae Vol.1, 64 Pleopis polyphemoides Vol.1, 186-187
Metacylis Vol.1, 68 Pleurobrachia pileus Vol.1, 154-155

Pneumodermatidae gen.sp. Vol.1, 176 Temora turbinata Vol.2, 54-55
Podocoryne sp. Vol.1, 138-139 Thalassocaris obscura Vol.2, 145
Pontellidae Vol.2, 58 Thalia sp. Vol.2, 177
Pontella Vol.2, 74 Thenus orientalis Vol.2, 155-157
Pontella danae Vol.2, 74-75 Tintinnidae Vol.1, 96
Pontella investigatoris Vol.2, 76-77 Tintinnidiidae Vol.1, 14
Pontellopsis Vol.2, 78 Tintinnopsis Vol.1, 20
Pontellopsis herdmani Vol.2, 78-79 Tintinnopsis acuminata Vol.1, 20-21
Protorhabdonella Vol.1, 80 Tintinnopsis ampla Vol.1, 22-23
Protorhabdonella curta Vol.1, 80 Tintinnopsis angusta Vol.1, 24
Protorhabdonella simplex Vol.1, 81 Tintinnopsis baltica Vol.1, 25
Pseudevadne tergestina Vol.1, 188-189 Tintinnopsis beroidea Vol.1, 26
Pseudodiaptomidae Vol.2, 48 Tintinnopsis compressa Vol.1, 27
Pseudodiaptomus Vol.2, 48 Tintinnopsis dadayi Vol.1, 28-29
Pseudodiaptomus arabicus Vol.2, 49 Tintinnopsis directa Vol.1, 30-31
Pseudodiaptomus ardjuna Vol.2, 50 Tintinnopsis failakkaensis Vol.1, 32-33
Tintinnopsis gracilis Vol.1, 34-35
Rhabdonellidae Vol.1, 80 Tintinnopsis karajacensis Vol.1, 36
Rhabdonella Vol.1, 82 Tintinnopsis lobiancoi Vol.1, 37
Rhabdonella conica Vol.1, 82-83 Tintinnopsis lohmannii Vol.1, 38
Rhabdonella sp. Vol.1, 85 Tintinnopsis mortensenii Vol.1, 39
Rhabdonella striata Vol.1, 84 Tintinnopsis nana Vol.1, 40
Rhopalophthalmus sp. Vol.2, 126-127 Tintinnopsis orientalis Vol.1, 41
Rotifera Vol.1, 164-165 Tintinnopsis parva Vol.1, 42
Tintinnopsis parvula Vol.1, 43
Sagitta enflata Vol.2, 169 Tintinnopsis radix Vol.1, 44-45
Sagitta neglecta Vol.2, 170 Tintinnopsis rotundata Vol.1, 46
Sagitta pulchra Vol.2, 171 Tintinnopsis sacculus Vol.1, 47
Sagitta regularis Vol.2, 172 Tintinnopsis schotti Vol.1, 48-49
Salpingacantha Vol.1, 117 Tintinnopsis tocantinensis Vol.1, 50-51
Salpingacantha unguiculata Vol.1, 117 Tintinnopsis turbo Vol.1, 52
Salpingella Vol.1, 114 Tintinnopsis undella Vol.1, 53
Salpingella attenuata Vol.1, 114-115 Tintinnopsis urnula Vol.1, 54-55
Salpingella rotundata Vol.1, 116 Tortanidae Vol.2, 90
Sanderia malayensis Vol.1, 150-151 Tortanus Vol.2, 90
Sapphirinidae Vol.2, 106 Tortanus barbatus Vol.2, 90-91
Sapphirina Vol.2, 110 Tortanus forcipatus Vol.2, 92-93
Sapphirina nigromaculata Vol.2, 110-111 Turbellaria Larvae Vol.1, 160-161
Sepiolidae gen.sp. Vol.1, 177
Sergestes sp. Vol.2, 142-143 Undellidae Vol.1, 94
Solenocera crassicornis Vol.2, 132-133 Undella Vol.1, 94
Solenocera hextii Vol.2, 134 Undella dilatata Vol.1, 94-95
Solmundella bitentaculata Vol.1, 142-143 Upogebia sp. Vol.2, 154
Stenopus sp. Vol.2, 144
Subeucalanus Vol.2, 29 Weelia cylindrica Vol.2, 176
Subeucalanus flemingeri Vol.2, 30-31
Subeucalanus subcrassus Vol.2, 32 Xystonellidae Vol.1, 86
Synchaetidae Vol.1, 164-165 Xystonella Vol.1, 87
Xystonella treforti Vol.1, 87
Tadpole Larva Vol.2, 178 Xystonellopsis Vol.1, 88
Temoridae Vol.2, 51 Xystonellopsis gaussi Vol.1, 88
Temora Vol.2, 51
Temora discaudata Vol.2, 52-53 Zoothamnium sp. Vol.1, 120-121

Photography by:
D. Altukhov: figs. 119; 120 b; 124; 130; 133; 143; 152 c, d; 171; 172; 174 c; 184; 186 a; 193 b; 194; 196;
197; 200 d; 201 a, c; 206; 211 b; 212 b; 214; 215; 223; 224; 225; 227; 228; 236; 239; 241; 246.
Dr. I. Drapun: figs. 112; 113; 120 a; 136; 137; 140; 141; 146; 147; 149; 152 a, b; 156; 157; 161; 162; 164
b; 166; 167; 169; 170; 174 a, b; 175; 177; 178; 180; 181; 183; 200 a, b, c; 201 b; 204; 205; 218-219; 221;
222; 230; 231; 237; 243; 247; 250; 251; 253; 255; 256; 258; 261; 262; 264; 266; 268; 270; 271; 273; 275;
277.
Dr. S. Khvorov: figs. 78-93; 98; 99; 101; 103 a; 109 a; 110 a; 111 a; 278, 279-321; 322; 324; 325; 327 a, b.
Dr. I. Prusova: figs. 126 a, b; 164 a, c; 186 b; 193 a; 211 a; 212 a.
Dr. M. Saburova: fig. 107 d.
Dr. V. Skryabin: figs. 2-77; 94-97; 100 a-d, 103 b, e, f; 104-107a, b, c; 108; 109 b, c, 110 b, c; 111 b, c;
126 c, d; 159; 189; 190; 208; 323; 326; 327 c.

MARINE ZOOPLANKTON PRACTICAL GUIDE for the Northwestern Arabian Gulf
MARINE ZOOPLANKTON
PRACTICAL GUIDE
ISBN 978-99966-95-07-0
Kuwait Institute for Scienatific Research
P.O. Box 24885, Safat - 13109, Kuwait
Tel: (965) 24989000 Fax: (965) 24989399
www.kisr.edu.kw
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Marine Zooplankton Practical Guide (Volumes 1 and 2) for the Northwestern

Book · July 2011

Faiza Al-Yamani Alexandra Gubanova

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for the Northwestern Arabian Gulf

Faiza Yousif Al-Yamani

Faiza Yousif Al-Yamani

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

General Characteristics, Morphology, Terminology and Abbreviations 2

List of Copepods Taxa 8

Systematic account of Copepod Species from the Arabian Gulf 10

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

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

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

2.14. Mysida 126

2.15. Decapoda 128

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

2.16. Chaetognatha 169

2.17. Urochordata 173

2.18. Cephalochordata 180

Index of Scientific Names 193

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

Zooplankton research is an essential component

The samples used to produce this guide were

This illustrated zooplankton guide is an

Information about methods on zooplankton

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

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

The authors are grateful to the Kuwait Institute

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

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

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

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

In the podoplean type of tagmosis (in this document, Cyclopoida, Harpacticoida,

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

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

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

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

Fig. 115. Cont’d.

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

Fig. 116. Plan of body organization in a. gymnopleans

Fig. 117. Differences between copepod Orders:

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

Phylum Class Susclass Order Family Genus / Species

Arthropoda Crustacea Copepoda Calanoida Calanidae Canthocalanus pauper

Eucalanidae Subeucalanus flemingeri

Clausocalanidae Clausocalanus minor

Euchaetidae Euchaeta concinna

Centropagidae Centropages furcatus

Pseudodiaptomidae Pseudodiaptomus arabicus

Temoridae Temora discaudata

Candaciidae Candacia bradyi

Pontellidae Calanopia elliptica

Acartiidae Acartia (Acanthacartia) fossae

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

Tortanidae Tortanus barbatus

Cyclopoida Oithonidae Oithona attenuata

Sapphirinidae Copilia mirabilis