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.

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.

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

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.

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.

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.