Identification guide to Antarctic Copepods

(From: South Atlantic Zooplankton, by: Demetrio Boltovskoy)

Phylum Arthropoda (von Siebold, 1848)

An arthropod is an invertebrate animal having an exoskeleton, a segmented body, and jointed
appendages. Arthropods are characterized by their jointed limbs and cuticle made of chitin,
often mineralized with calcium carbonate. The arthropod body plan consists of segments,
each with a pair of appendages. The rigid cuticle inhibits growth ,so arthropods replace it
periodically by moulting. Their versatility has enabled them to become the most species-rich
members of all ecological guilds in most environments. They have over a million described
species, making up more than 80% of all described living animal species, some of which,
unlike most animals, are very successful in dry environments.

Subphylum Crustacea (Brünnich, 1722)

Crustaceans form a large, diverse arthropod taxon. They are distinguished from other groups
of arthropods by the possession of biramous (two-parted) limbs, and by their larval forms,
such as the nauplius stage of branchiopods and copepods.

Superclass Multicrustacea (Regier et al, 2010)

Class Hexanauplia (Oakley et al, 2013)

Subclass Copepoda (H.Mline-Edwards, 1840)

Marine copepods are small crustaceans usually between 0.2 – 12mm in length.
The classification of the copepods is largely based on body tagmosis (division into functional
regions) and segmentation and armature of the various limbs. The major body articulation
subdivides the body into an anterior part – the prosome (Pr)- and a posterior part – the
urosome (Ur). 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), respectively, and the
first thoracic segment bearing the maxillipeds (Mxp). (Figure 1).
Figure 1. Structure of calanoid body and mouthparts.
There are seven post-cephalic thoracic somites: the maxilliped-bearing somite, five
pedigerous somites (Pd1-5), each bearing a pair of swimming legs (P1-5) (Figure 2), and the
genital somite. The seventh thoracic or genital somite (Gns) bears the sixth pair of legs (P6)
which are highly modified and form the opercula that close off the paired female genital
apertures. Each pair of swimming legs is joined by an intercoxal sclerite which may be fused
to the coxae. The swimming legs usually have two basal segments: coxa (C) and basis (B).
Two rami are articulated to the basis: an outer exopod (Exp) and inner endopod(Enp). The
exopod and endopod are boardered by spines and/or setae which are denoted in the family
descriptions according to the formula system of Sewell(1949) (Figure 2).

Whitin 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 Figure 3). In the gymnoplean tagmosis (Platycopioida and Calanoida) the major body
articulation is located between the fifth pedigerous somite (Pd5), primitively bearing the P5,
and the genital somite (Gns). In the podoplean type of tagmosis (all reemaining copepod
orders) the major articulation primitively lies between Pd4 and Pd5 (between fifth and sixth
thoracic somites).

In gymnopleans, the urosome includes the anterior somite corresponding to the seventh
thoracic somite (= genital somite). In females of Calanoida it is usually fused with the first
abdominal somite, forming a genital double-somite. The abdomen is the limbless post-genital
region of the body. In podopleans the urosome comprise the sixth thoracic somite bearing the
P5, the seventh thoracic somite and five abdominal somites. In the majority of Harpacticoida,
Poecilostomatoida and Cyclopoida the female second and third urosomites (= seventh
thoracic segment and first abdominal somites) are usually fused to form a genital double-
somite. The difference between gymnopleans and podopleans may be observed even in the
early copepodite development (Fig 3). The last somite of the urosome is the anal somite
bearing a pair of caudal remi (CR).

Antarctic copepods that are large and dominate biomass: Calanoides acutus, Rhincalanus gigas,
Calanus simillimus, Calanus propinquus and Metridia gerlachei (Andrews 1996; Ommanney 1936;
Voronina 1970).
Calanus acutus and Rhincalanus gigas are the most herbivorous species of the group (Hopkins
and Torres 1989; Atkinson et al 1996a; Schmidt et al 2003), and suspension feeders that ingest
both motile and non-motile particles broadly according to their abundance (Schnack 1985;
Atkinson 1996a).
Calanus simillimus, C. propinquus and M.gerlachei appear to be slightly more omnivorous,
possibly able to switch between suspension and raptorial feeding as does Acartia tonsa
(Kiorboe 1996), and ingest a higher proportion of motile prey (Hopkins and Torres 1989;
Atkinson 1995; Burghart et al 1999; Schmidt et al 2003).
Figure 2. Structure of calanoid swimming legs. Basic copepod swimming leg, showing the
maximum setation of a second leg. Sewell’s system of spine and setal description is given in
the box. In the keys P1-P4 segmentation is denoted as, e.g., 3/3 (number of Exp
segments/number of Enp segments). The spine (roman numerals) and seta (Arabic numerals)
formula for the swimming legs is given in mode of Sewell (1949)
Figure 3. Comparison of the developmental pattern in podoplean and gymnoplean copepods.
CI-CV: first copepodid to fifth copepodid developmental stages: solid stars indicate flexure
planes, hollow stars indicates poorly defined flexure plane, heavy arrows indicate the stage at
which the definitive tagmosis is attained and specialisation of the joint commences.
Abbreviations used in the figures and keys (Fig. 1 and Fig. 2)
A1: antennule
A2: antenna
B: basis
C: coxa
Ce: cephalosome (head region when this includes only the somite of the maxillipeds)
CR: caudal rami
Enp: endopod; Enp1-3 – endopod segments 1-3
Exp: exopodo; Exp 1-3 – exopod segments 1-3
Gn: gnathobase
Gns: genital somite
Md: mandible
Mdp: mandibular palp
Mx1: maxillule
Le1: coxal epipodite – first external lobe
Le2: basal exite – second external lobe
Li 1: praecoxal arthrite – first internal lobe
Li2: coxal endite – second internal lobe
Li 3, 4: basal endites – third and fourth internal lobes
Mx2: maxilla
Li 1, 2: praecoxal endites – first and second lobes
Li 3, 4: coxal endites – third and fourth lobes
Li 5: basal endite – fifth lobe
Li 6: (if present) – sixth lobe
Mxp: maxilliped
Li 1: praecoxal endite
Li 2-4: syncoxal endites
P1-5: swimming legs 1-5. The outer elements are given first an are separated by a hyphen
from the inner elements. In the keys P1-P4 segmentation is denoted as, e.g., 3/3 (number of
Exp segments/number of Enp segments). The spine (roman numerals) and seta (Arabic
numerals) formula for the swimming legs is given in mode of Sewell (1949). The terminal
segment of each ramus has 3 components, sepparated by commas: outer, terminal, inner.
Pc: praecoxa
Pcs: praecoxal sclerite
Pd1-5: pedigerous somites 1-5
Pr: prosome (the anterior region of the body limited by a major articulation)
Sc: syncoxa (fused praecoxa and coxa)
Url-5: urosomites 1-5, last somite being the anal somite
ven: ventral view.
Subclass Copepoda (Huys et al, 1996)
Copepods at some stage in their life-cycle possess at least 2 pairs of swimming legs with rami
connected by an intercoxal sclerite; a Ce into which the Mxp bearing somite is incorporated;
uniramous A1 with up to 27 segments, although in many copepods (harpacticoids) the
segmentation of A1 is secondarily reduced; possess egg-sacs, although some important
groups lack true egg-sacs and they are secondarily lost in some highly derived parasitic forms
(Huys et al., 1996).

Key to orders of Copepoda (adapted from Huys et al, 1996).

1 Body showing gymnoplean tagmosis, with Pr-Ur boundary located between the Pd5
and Gns (Fig. 4.A): ………………………………………………………………………….Calanoida

1a Body showing podoplean tagmosis, with Pr-Ur boundary located between Pd4 and 5
(Fig. 4.B)…………………………………………………………………………………….………….. 2

2 A2 Exp usually with 2 or more segments (Fig. 4.C), rarely 1-segmented or absent, in
which case baseoendopod of P5 with 2 or more setae (Fig.4.D): ………………….Harpacticoida

2a A2 Exp 1-segmented or absent (Fig 4.E), P5 baseoendopod represented by at most 1

seta (Fig. 4.F, H)………………………………………………………………………………………...3

3 Male A1 typically geniculate (Fig. 4.H). Female A1 with up to 26-segmented. Mxp

usually similar in both sexes. Female Gns with single midventral copulatory pore (except in
Oithonidae): ………………………………………………………………………………..Cyclopoida

3a Male A1 non-geniculate. Female A1 at most 7-segmented (Fig 4. I). Mxp always

sexually dimorphic (Fig 4.G, J). Female Gns with copulatory pores located within dorso-
lateral genital apertures (Fig. 4 F) (except in Erebonasteridae):…………….. .Poecilostomatoida
Figure 4. Illustration needed in the key to orders of copepoda: Calanoida, Harpacticoida,
Cyclopoida and Poecilostomatoida.
Order Calanoida (Sars, 1903)
The Calanoide 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 C. 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).

Key to the families of Calanoida (modified from Brodsky et al, 1983)

Females
1 Ur with no more than 3 free somites…………………………………………………….…...2
1a Ur with 4 free somites ..………………………………………………………………………13
2 P5 present……………………………………………………………………………………….3
2a P5 absent:………………………….. Eucalanidae (Eucalanus, Subeucalanus, Paraeucalanus)
3 P5 similar in structure to P1-P4. Segmentation of P1-P4: 3/3, 3/3, 3/3, 3/3…………….4
3a P5 very different in structure from P1-P4, P1-P4 segmentation may be different from
3/3……………………………………………………………………………………………………….5
4 P5 Exp2 with finger-like projection instead of internal seta (Fig 5.A). Posterior corners
of Pr often prolonged into points. Oral parts not reduced. Distal setae of Mxp without small
shield-shaped appendages: ………………………………………………………….Centropagidae.
4a P5 Exp2 segment without finger-like projection internally (Fig 5.B). Posterior corners of
Pr usually rounded. Oral parts strongly reduced to different degrees. Distal setae of Mxp
strong, curved and often with small shield-shaped appendages: ………………….Augaptilidae
5 Inner margin of P2 Enp1 with certain excavation which has one or two hooks on its
inner margin (fig 5.C). R with two filaments. Pr in Pleuromamma with small black rounded
organ on one side of the body. Segmentation of P1-P4:3/3:……………………….. Metridinidae
5a Inner margin of P2 Enp1 without excavation or hook. P1-P4 with various segmentation
which often differ from 3/3…………………………………………………………………………...6
6 Segmentation of P1-P4:3/1-3, 3/2, 3/2, 3/2…………………………………………………7
6a Segmentation of P1-P4: 1-3/1-2, 3/2-3, 3/3, 3/3…………………………………………..11
7 Anterior part of head with (sometimes without) one or two pairs of cuticular lenses,
often lens is present in base of bifurcate R (Fig 5.D, E):………………………………..Pontellidae
7a Anterior part of head without cuticular lenses. R not bifurcate; as 2 filaments or
absent……………………………………………………………………………………………………8
8 Anterior part of head with single pigmented eye…………………………………………..9
8a Anterior part of head without single pigmented eye…………………………………..…10
9 Anterior to upper labrum a rounded, densely haired plate present (Fig. 5.F). R absent.
P2-P4 Exp with external spines: ………………………………………………………….Tortanidae
9a Densely haired plate anterior to labrum absent (Fig 5.G) R present. P2-P4 Exp without
external spines, only unarticulated spiniform protrusion of segment present (Fig 5.J):
………………………………………………………………………………………………..Acartiidae
10 Anterior part of head usually rectangular (Fig. 5.I). Mx2 enlarged, with strong distal
setae. Mxp strongly reduced. Outer margin of P2-P4 Exp segments finely serrated. Some
species with body partly coloured by dark pigment. Swimming legs usually heavily
pigmented black or dark-brown:………………………………………………………. Candaciidae
10a Anterior part of head rounded (Fig 5.N). Mx2 not enlarged, Mxp not reduced. Outer
margins of P2-P4 Exp segments not serrated. Species not dark-pigmented:………... Temoridae
11 Segmentation of P1:2-3/1-2, P2:3/3………………………………………………………...12
11a Segmentation of P1: 1-3/1-2, P2: 3/2-3: ……………………………………...Paracalanidae
12 A1 twice as long as body. Spermathecae nearly spherical (Fig. 6.A, B). Segmentation of
P1: 3/1. Posterior margins of Pd2-4 and Gns without spines: …………………...Mecynoceridae
12a A1 not more than 1.5 times longer than body. Spermathecae not spherical.
Segmentation of P1: 2/2. Posterior margins some or all Pd and Gns with two dorsal spines
each and spine-like projections laterally (Fig 6. C): …………………………………..Eucalanidae
13 P5 present……………………………………………………………………………………...14
13a P5 absent……………………………………………………………………………………….29
14 Segmentation of P1-P4: 3/3; rarely P1 Exp or P1 Enp 2-segmented……………………..15
14a P1 Enp 1-3-segmented, P2 Enp 2-3-segmented, segmentation of P3-P4: 3/3, 3/3……...23
15 P5 similar in structure to P1-P4 (Fig 6.M)…………………………………………………..16
15a P5 strongly differing in structure from P1-P4 (Fig. 6 H, M) …………………………...…20
16 Mx1 Le2 present, with a single seta…………………………………………………………17
16a Mx1 lacking L2 with seta…………………………………………………………………….18
17 P2-P4 Exp3 with 3 outer spines (Fig. 5.L). P5 C always smooth on inner margin. Large-
sized copepods (8-17 mm): …………………………………………………………..Megacalanidae
17a P2-P4 Exp3 with 2 outer spines (Fig 5.K). P5 C often serrated on inner margin (Fig 6.M).
Body length less than 10 mm.: …………………………………………………………….Calanidae
18 CR long, usually 3-5 times longer than wide, if shorter then asymmetrical……………19
18a CR short, symmetrical. Md with elongated, narrow gnathobase. Oral parts strongly
reduced to different degrees. Distal setae of Mxp powerful, strongly curved. Usually very
transparent copepods. Sometimes frontal spine present (Fig 6.E):………………..Augaptilidae
19 Anterior surface of P1 B adjacent to Enp with short cylindrical projection with seta (Fig.
6.K). CR long, 3-10 times longer than wide, symmetrical or asymmetrical, their distal ends
with parallel borders (Fig. 6.F):……………………………………………….…………Lucicutiidae
19a Anterior surface of P1 B adjacent to Enp without cylindrical projection. CR only 3-5
times longer than wide; usually asymmetrical, their distal ends cut obliquely (Fig. 7.H):
…………………………………………………………………………………….…..Heterorhabdidae
20 P1 Enp1-2-segmented, P1 Exp1 lacking outer spine, P3 B with outer spine:
……………………………………………………………………………………….....Bathypontiidae
20a P1 Enp 3-segmented; P1 Exp1 with outer spine. P3 B without outer spine…………….21
21 Copepods usually with characteristic eye composed of pigmented spot surrounded by
refractile lens. Mx1 Le1 with 10 setae. Mx2 Li1-Lu2 with 4 and 3 setae respectively (Fig. 5.H):
……………………………………………………………………………………Pseudodiaptomidae
21a Copepods usually without eye. Mx1 Le1 with 5-9 setae or without setae. Mx2 Li1 with
1-3 setae, Li2 with 1-2 setae…………………………………………………………………………..22
22 Ce rounded or pinted at apex (Fig 7.D). Mdp with rudimentary Enp or without Enp. P5
4-segmented penultimate segment with inner projection bearing 1-3 setae (rarely P5 1- or 2-
segmented), P5 may be biramous. Setae of oral parts and CR long, hairy and brightly
coloured:…………………………………………………………………………………...Arietellidae
22a Ce rounded (Fig.6.G). Mdp with well developed Enp. P5 5-segmented, uniramous,
with distolateral teeth (Fig. 5.M):…………………………………………………….Phyllopodidae
23 Segmentation of P1-P2: 3/2, 3/3. Terminal spines of Exp of P2-P4 smooth (Fig. 6.L):
……………………………………………………………………………………………Paracalanidae
23a P1 Enp 1-segmented. P2 Enp 2-segmeted…………………………………………………..24
24 Mx2 with setae modified into sensoty appendages (Fig. 7.A, B) ………………………...26
24a Mx2 without sensory setae…………………………………………………………………..25
25 R or two thin long filaments or two-pointed (Fig. 7.C)…………………..Clausocalanidae
25a R, if present, one-pointed, thin and hardly visible (Fig. 7.D)…………...Mesaiokeratidae
26 Mx2 with 1 worm-like and 7 brush-like sensory appendages distally (Fig.7.A). Mx2 Li1
usually with 5, rarely with 4 setae: ……………………………………………………...Phaennidae
26a Mx2 usually with 3 (rarely 4) worm-lige and 5 (rarely 4) brush-like sensory appendages
distally (Fig. 7.B). Mx2 Li1 with 3-4 setae (5 setae in Landrumius)……………………………….27
27 Posterior corners of Pr extended into 2 points (Fig 6.J):.…Phaennidae (Neoscolecithrix)
27a Posterior corners of Pr not extended into 2 points…………………….…………………..28
28 P1-P4 surfaces weakly spinulose (Fig 7.K). Mx1 Li1 not extending beyond Enp. Mx2 Li5
usually with sensory appendage (Fig. 7.B): ……………………………………....Scolecitrichidae
28a P1-P4 surfaces weakly spinulose. Mx1 Li1 comparatively large, extending or reaching
beyond Enp. Mx2 Li5 without sensoty appendage (Fig.6.I): ………………………....Tharybidae
29 Segmentation of P1-P4: 2-3/1, 3/1, 3/3, 3/3, or 2-3/1, 3/2, 3/3, 3/3…………………...30
29a Segmentation of P1-P4: 1/1, 2-3/1-2, 3/1-2, 3/1-3 or 2-3/1, 2-3/1-2, 2-3/2-3, 2-3/2-
3…………………………………………………………………………………………………………36
30 Segmentation of P1-P4: 2-3/1, 3/1, 3/3, 3/3. Supralabrum (anterior to labrum) is
highly prominent structure. Mxp strong with long, strongly built, slightly curved setae. Many
species with asymmetrical genital field (Fig 7.E, I, J): ………………………………….Euchatidae
30a Segmentation of P1-P4: 2-3/1, 3/2, 3/3, 3/3, (sometimes P2: 3/1 -Euchirella).
Supralabrum (anterior to labrum) not highly prominent. Mxp moderately developed. Genital
field usually symmetrical ……………………………………………………………………………31
31 Mx2 with terminal setae modified into sensory worm-like and brush-like appendages
(Fig.7.A, B) …………………………………………………………………………………………….32
31a Mx2 without terminal sensory appendages………………………………………………..34
32 Mx2 with 1 worm-like and 7 brush-like appendages (Fig.7.A). R 2-pointed:
……………………………………………………………………………………………....Phaennidae
32a Mx2 with more than 1 worm-like appendage. R one-pointed, or with 2 filaments……33
33 Small copepods, less than 1 mm in total length:………………………………….Diaixidae
33a Copepods more than 1 mm in total length: ……………………………….Scolecitrichidae
34 P3-P4 Exp3 with 4 inner setae. Surfaces of P usually smooth……………………………35
34a P3-P4 Exp3 with 5 inner setae. Anterior and usually posterior surfaces of P usually
spinulose (Fig. 7.G): ………………………………………………………………….Spinocalanidae
35 Posterior corners of Pr always rounded. R of 2 filaments:……………….Clausocalanidae
35a Posterior corners of Pr often prolonged into points, 2-pointed without filaments or,
rarely absent. Ce sometimes with frontal spine (Gaetanus); in some species of Euchirella,
Chirundina, Undeuchaeta with crest (Fig. 8.A): …………………………………………..Aetideidae
36 Mx2 with setae on Li1-Li5 strongly transformed (Fig.7.F). Segmentation of P1-P4: 1/1,
2-3/1-2, 3/1-2, 3/1-3. R absent: …………………………………….……..Aetideidae (Chiridiella)
36a Mx2 without strongly transformed setae on Li1-Li5. Segmentation of P1-P4: 2-3/1, 2-
3/1-2, 2-3/2-3, 2-3/2-3. R two-pointed: ………………………………...Euchaetidae (Valdiviella)

Males
1 A1 on one side geniculate (Fig. 7.H)…………………………………………………………2
1a Neither A1 geniculate………………………………………………………………………...16
2 Inner margin of P2 Enp1 without excavation or hooks…………………………………….3
2a Inner margin of P2 Enp1 with excavation and hook, or pair of hooks (Fig. 5.C):
…………………………………………………………………………………………….Metridinidae
3 Left A1 geniculated ……………………………………………………………………………4
3a Right A1 geniculated …………………………………………………………………………..8
4 P5 of similar segmentation to P1-P4; distal segmented modified compared with those
of P2-P4 (Fig. 8.B, C, D)………………………………………………………………………………..6
4a P5 very different from P1-P4, with 3-segmented Exp and strongly reduced Enp,
sometimes Enp of one leg absent (Fig. 8.F, H)……………………………………………………….5
5 Right P5 without Enp. In both P5, B with long outer seta (Fig. 8.F):
…………………………………………………………………………………………...Phyllopodidae
5a Right P5 with Enp. Outer seta of right P5 B usually longer than left one; latter may be
absent (Fig. 8.H):…………………………………………………………………………..Arietellidae
6 Anterior surface of P1 B adjacent to Enp without short cylindrical protrusion, but seta
present: ………………………………………………………………………………………………….7
6a Anterior surface of P1 B adjacent to Enp with short cylindrical protusion bearing seta
(Fig. 6.K; 8.D): …………………………………………………………………………….Lucicutiidae
7 Both P5 usually 3-segmented but differ from P1-4 as Exp modified and B has inner
projection (Fig. 8.B): …………………………………………………………………Heterohabdidae
7a Both P5 2-3-segmented, nearly like P4, but differs as distal Exp setae short, or
transformed into spines (Fig. 8.C):……………………………………………………..Augaptilidae
8 One of P5 legs chelate (Fig. 8.E, G, I; 19.A, B, D)…………………………………………….9
8a Neither P5 chelate……………………………………………………………………………..15
9 P2-P4 Enp 2-segmented, Exp 3-segmented………………………………………………...11
9a P2-P4 Enp and Exp 3-segmented……………………………………………………………10
10 P3-4 Exp with 3 outers spines ………………………………………………..Centropagidae
10a P3-4 Exp with 2 outer spines …………………………………………..Pseudodiaptomidae
11 Cuticular lenses or pigmented eye present on anterior part of Ce………………………12
11a Cuticular lenses and pigmented eye absent on anterior part of Ce ……………………..14
12 Anterior part of Ce sometimes with 1 or 2 pairs of cuticular lenses, often lens present
near base of strong R (Fig. 5.E): ………………………………………………………….Pontellidae
12a Anterior part of Ce alsways lacks cuticular lenses, single pigmented eye present. R is
not strong or absent ………………………………………………………………………………….13
13 Anteriorly rounded hairy plate present, anterior to upper labrum (Fig. 5.F). P5
uniramous, massive, assymmetrical; 3-segmented on right with large chela, 4-segmented on
left, curved (Fig. 9.B): …………………………………………………………………….Tortanidae
13a Rounded hairy plate, anterior to upper labrum, absent (Fig.5.G). P5 uniramous, right 4-
segmented, distal segment of right, longer leg, curved, left 3-4 segmented (Fig. 9.D):
……………………………………………………………………………………………….Arcartiidae
14 Posterior corners of Pr always asymmetrical. Gns is almost always asymmetrical,
bearing process on right side: …………………………………………………………..Candaciidae
14a Posterior corners of Pr and Gns usually symmetrical. Gns usually without process on
right side: …………………………………………………………………………………..Termoridae
15 P5 differs in structure from P1-P4. Medium sized and small calanoids (1.0 – 5.6 mm):
………………………………………………………………………………………….Bathypontiidae
15a P5 close in structure to P1-4 (Fig. 9.F). Large calanoids (8-10 mm):…….Megacalanidae
16 Ur of 5 free somites. Anterior part of head without elongated prominence anterior to
A1 (Fig.9.I)……………………………………………………………………………………………17
16a Ur of 4 free somites (anal somite fused to CR) (Fig. 9.J, K). Anterior part of head
triangular, sometimes with elongated prominence anterior to A1: …………………Eucalanidae
17 P5 similar in structure to P1-P4 (Fig.10.A, B) ……………………………………………...18
17a P5 strongly differing from P1-P4 ……………………………………………………………19
18 P2-P4 Exp3 with 2 outer spines (Fig.5.K): ………………………………………..Calanidae
18a P2-P4 Exp3 with 3 outer spines (Fig.5.L): ………………………………….Megacalanidae
19 P1-P4 segmentation 2-3/1, 3/2, 3/3, 3/3…………………………………………………..20
19a P1-P4 segmentation different………………………………………………………………..27
20 Distal part of Mx2 with sensory appendages (Fig. 7.A, B) ……………………………….21
20a Distal part of Mx2 without sensory appendages…………………………………………..24
21 P1-P4 surface of segments usually highly spinulose ……………………………………...22
21a P1-P4 surface of segment poorly spinulose, or smooth…………………………………...23
22 P5 usually biramous on both sides, asymmetrical; C and B narrow and elongated on
left, short on right, with B usually very swollen (Fig. 9.M):…………………... Scolecitrichidae
22a P5 usually uniramous, if Enp present they are rudimentary, asymmetrical, longer on
left (Fig. 9.L): ……………………………………………………………………………..Phaennidae
23 Left P5 with long Enp, right usually uniramous (not in Parundinella) (Fig. 9.H):
………………………………………………………………………………………………Tharybidae
23a Both P5 with uniramous (Fig.9.G): .………………………………………………...Diaxidae
24 P2-P4 Exp3 with 4 setae……………………………………………………………………...25
24a P2-P4 Exp3 with 5 setae (Fig. 10.K): ………………………………………..Spinocalanidae
25 R 1-pointed, 2-pointed, or absent, never with filaments ………………………………...26
25a R of 2 filaments or rudimentary, rarely absent. P5 elongated, uniramous, very
asymmetrical (right leg sometimes rudimentary) (Fig.10.F): ………………..…Clausocalanidae
26 P5 biramous with Enp variously developed, or uniramous, or with only one leg
present. Calanoids usually longer than 1 mm (Fig. 10.D, G, L): ………………………Aetideidae
26a P5 always uniramous, asymmetrical: 3-segmented on right, short, nearly as long as first
segment of 5-segmented left leg. Calanoids usually less than 1 mm (Fig.10.C):
……………………………………………………………………………………...…Mesaiokeratidae
27 P1-P4 segmentation: 2-3/1, 3/1, 3/3, 3/3. P5 powerful, nearly as long as Ur;
asymmetrical. Exp 3-segmented on left, Exp2 with (or without Valdiviella) serrated lamella,
sometimes its tip prolonged into thin long seta (Fig.10.I, J, M): …………….……….Euchaetidae
27a P1-P4 segmentation: 3/1-3, 3/3, 3/3, 3/3………………………………...………………...28
28 P1 Enp 3-segmented (Fig 10.E, H): ………………………………...……………...Calanidae
28a P1 Enp 1-2 segmented ………………………………………………………………………..29
29 Both P5 5-segmented of nearly same length (Fig.11.A). P1 Enp always 1-segmented. A2
not described: ……………………………………………………………..…………..Mecynoceridae
29a If both P5 present, only one 5-segmented; strongly asymmetrical length (Fig. 11.B,G).
P1 Enp 2-segmented, rarely 1-segmented. A2 terminal segment very short, without 3 terminal
setae (Fig.11.H): ………………………………………………………………………...Paracalanidae
Figure 5.
Figure 6.
Figure 7.
Figure 8.
Figure 9.
Figure 10.
Figure 11.
Family Calanidae (Dana, 1846)

Females

Ce and Pd1 may be fused or separate, Pd4 and 5 always separate. R of 2 filaments. CR with 4
subequal, terminal setae and 1 outer seta. Ur of 4 somites. A1 25-segmented, generally
segments 8-9 partly fused; segments 23 and 24 with elongate seta. A2 C 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. Exp and Enp 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, Enp with 14-15 setae; Exp with 11 setae; Le1 with 9 setae; Le2 with 1 seta. Mx2
Li1-Li5 with 4-6, 3,3,3 and 4 setae, respectively. Terminal part with about 10 setae. Mxp C with
1 proximal seta, then with groups of 2, 4, and 4 setae from proximal to distal; Enp2-6 with 4, 4,
¾, 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). Spine and seta formula:
Males

Ur 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 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 P, left leg variously modified; Enp sometimes reduced and devoid of segmentation on
one or both sides.

Key to genera of Calanidae (Bradford and Jillet, 1974)

Females

1 P5 C inner edge with teeth…..………………………………………………………………..2

1a P5 C inner edge without teeth (naked)………..……………………………………………..4
2 P2-3 B with posterior surface spinules:………..………………….Cosmocalanus darwinii
2a P2-3 B without surface spinules (naked): ……...…………………………………………….3
3 Ce and Pd1 fused: ……………………………….……………….…….Nannocalanus minor
3a Ce and Pd1 partly fused: ……………………….…………………………………….Calanus
4 P2 Exp1 with recurved spine (hook) adjacent to outer spine: .……………….Neocalanus
4a P2 Exp1 without recurved (hook) adjacent to outer spine……..…………………………..5
5 Posterior corners of Pr pointed. P2 Exp2 with outer proximal edge invaginate:
……………………………………………………………………………….………Undinula vulgaris
5a Posterior corners of Pr rounded. P2 Exp2 outer edge without invagination ..…………...6
6 P1 Enp1 without inner seta. P5 Enp with 7 setae: ……………………………..…………... 7
6a P1 Enp1 with 1 inner seta. P5 Enp with 6 setae: ………………………………..Calanoides
7 Mx2 Li1 with 6 setae. P1 without modification: ……………….Mesocalanus tenuicornis
7a Mx2 Li1 with 4 setae. P1 C anterior margin terminates in well-defined projection; B
with inner marginal seta modified into proximally thickened spine:…. Canthocalanus pauper

Males

1 Left P5 highly modified, of complex structure (prehensile)……………………………….2

1a Left P5, if modified, then slightly (not prehensile)………………………………………….3
2 P5 C inner edge with teeth. P2 Exp2 without invaginated outer proximal border:
……………………………………………………………………………….Cosmocalanus darwinii
2a P5 C inner edge without teeth. P2 Exp2 with invaginated outer proximal border:
…………………………………………………………………………………...….Undinula vulgaris
3 P5 C inner edge with teeth…………………………………………………………………….4
3a P5 C inner edge without teeth……………………………………………………………...…5
4 Right P5 Exp3 with inner setae: ……………………………………....Nannocalanus minor
4a Right P5 Exp3 without inner setae: …………………………………………………Calanus
5 P2 Exp1 with recurved spine (hook) adjacent to outer spine: …….……….....Neocalanus
5a P2 Exp1 without recurved spine (hook) adjacent to outer spine…..………………..……..6
6 Both P5 Enp with 7 setae each: ………………………………..…Mesocalanus tenuicornis
6a P5 Enp with different number of setae……………………….………………………...…….7
7 Mouth parts not very reduced. Right P5 Enp with 8 setae, left with 2 setae:
……………………………………………………………………………..…...Canthocalanus pauper
7a Mouth parts very reduced. Right P5 Enp with different number of setae (not more than
6), often left Enp lacking setae:…………………………………………..……………….Calanoides

Key to species of Calanoides

Females

1 A1 extends beyond CR. Specimens 3.50-5.70 mm in total

length…………………………………………………………………….Calanoides acutus (Fig. 12).
1a A1 shorter. Specimens 2.25-3.90 mm in total length ……………………………………..…2
2 Spermatheca consists of 2 parts, proximal part is massive, in lateral view spermatheca
short:……………………………………………………………………..Calanoides macrocarinatus
2a Spermatheca is of different shape, in lateral view spermatheca long:
…………………………………………………………………………………..Calanoides carinatus

Males

1 Right P5 Enp1 and Enp2 nakes ……………………………………………………………….2

1a Right P5 Enp1 and Enp2 with 1 inner seta each: ………….…Calanoides acutus (Fig. 12)
2 Distal border of right Exp3 extends about 0.75 of way along left Exp2. Left Enp 2-
segmented:……………………………………………………………...Calanoides macrocarinatus
2a Distal border of right Exp3 reaching, or exceeding distal border of left Exp2. Left Enp 1-
segmented: …………………………………………………………………….Calanoides carinatus
Figure 12. Calanoides acutus. Female length (mm): 3.50-5.70. Male length (mm): 5.00-5.50.
Vertical distribution: Epipelagic and Mesopelagic. Inshore/offshore distribution: Oceanic.
Latitudinal distribution: Antarctic
Key to species of Calanus

Females

1 Posterior corners of Pr triangular, pointed distally ……………………………………….2

1a Posterior corners of Pr rounded:……………... Calanus agulhensis or Calanus australis
2 Total length 4.75-6.00 mm. Seta of A1 sefment 23 longer than last 8 segments:
………………………………………………………….………………..Calanus propinquus (Fig. 13)
2a Total length 2.65-3.80 mm. Seta of A1 segment 23 shorter than last 7 segments:
…………………………………………………………………………….Calanus simillimus (Fig.14)

Males

1 Right P5 Exp extends beyond distal border of left Exp1 …………………………………..2

1a Right P5 Exp does not extend as far as distal border of left Exp1:.
…………………………………………………………………………….Calanus simillimus (Fig.14)
2 Right P5 Exp extends more than half way along left Exp2; right P5 about 1.5-1.6 times
shorter than left P5:.....Calanus australis (2.90-3.47 mm) or Calanus agulhensis (2.74-3.00 mm)
2a Right P5 Exp extends less than half way along left Exp2; right P5 about twice shorter
than left:…………………………………………………………………Calanus propinquus (Fig.13)
Figure 13. Calanus propinquus. Female length (mm): 4.75-6.00. Male length (mm): 4.75-4.95.
Vertical distribution: Epipelagic / Mesopelagic. Inshore/offshore distribution: Oceanic.
Latitudinal distribution: Antarctic / Subantarctic
Figure 14. Calanus simillimus. Female length (mm): 2.65-3.80. Male length (mm): 2.62-3.42.
Vertical distribution: Epipelagic / Mesopelagic. Inshore/offshore distribution: Oceanic.
Latitudinal distribution: Subantarctic / antarctic.
Family Eucalanidae (Giesbrecht, 1892)

Females

Body elongate, especially anterior to A2, more or less transparent, often with triangular
anterior part of Ce. Ce and Pd1 fused, pd4 and 5 partly fused. CR sometimes asymmetrical;
usually left ramus slightly and second seta much longer than others. R of 2 long, slender
filaments. Ur of 3 or 4 somites with CR 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, B and Enp elongate; Exp with reduced number of setae; Le 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. Spine and seta formula:

Males

Ur of 5 somites, with CR usually completely fused to anal somite. A1 24-segmented. Oral

parts and P similar to females. P5 uniramous on both sides (Eucalanus, Pareucalanus),
biramous on left (Rhincalanus), or absent on right (Subeucalanus).
Key to genera of Eucalanidae

Females

1 P5 present. Dorsal surface of at least some of Pd2-4 and Gns with 2 spines: 1 on right
and 1 on left. Posterior corners of at least some Pd2-4 extended into spines: ……..Rhincalanus
1a P5 abstent. Dorsal surface of Pd2-4 and of Gns without spines. Posterior corners of
Pd2-4 lacking spines …………………………………………………………………………………..2
2 Female Ur of 3 free somites (somite 4 fused with CR): ………………Eucalanus hyalinus
2a Female Ur of 2 free somites (somite 3 fused with CR) ……………………………………..3
3 A2 Exp1 and 2 fused. Mx1 with 2 inner lobes………………………………Subeuncalanus
3a A2 Exp1 and 2 separate. Mx1 with 3 inner lobes: ……………………..…….Pareucalanus

Males

1 At least Gns with 2 dorsal spines. Left P5 biramous: ………………………...Rhincalanus

1a Gns without dorsal spines. P5 uniramous ………………………………………………….2
2 Right P5 4-segmented: …………………………………………………..Eucalanus hyalinus
2a Right P5 3-segmented, or absent……………………………………………………………..3
3 Right P5 absent:………………………………………………………………....Subeucalanus
3a Right P5 present, 3-segmented: ……………………………………………….Pareucalanus

Key to species of Rhincalanus

Females

1 P5 terminal segment with smooth outer terminal spine, shorter than its segment:
…………………………………………………………………………………..Rhincalanus cornutus
1a P5 terminal segment with plumose outer setae (or spine), much longer than its
segment………………………………………………………………………………………………….2
2 Dorsal spines absent from Pd3 and 4: ……………………...…Rhincalanus gigas (Fig. 15)
2a Dorsal spines present on Pd3 and 4: ………………………………....Rhincalanus nasutus

Males

1 Specimens are over 6.5 mm: ……………………………..…….Rhincalanus gigas (Fig. 15)

1a Specimens are less than 6.5 m length: ……………………………….Rhincalanus nasutus
Figure 15. Rhincalanus gigas. Female length (mm): 6.49-9.30. Male length (mm): 6.90-7.20.
Vertical distribution: Epipelagic / Mesopelagic. Inshore/offshore distribution: Oceanic.
Latitudinal distribution: Antarctic (to 50°S).
Family Metridinidae (Sars, 1092)

Females

Body elongate. Ce and Pd1 separate, Pd4 and 5 fused, Posterolateral corners of Pr usually
rounded, expanded in Gaussia; Pd1 with dark-pigmented spot on one side in Pleuromamma. Ur
of 3 somites, CR often asymmetrical with up to 6 setae; genital apparatus comprising a
comoon genital aperture located medially on ventral surface of Gns; copulatory pore
contained within median genital aperture. R with paired filaments. A1 23-24-segmented
segments 1 and 2 fused, segments 3 to 7 separate, segments 8 and 9 fused, segments 10-22
separate, apical segment double. A2 with C and B separate with 1 and 2 setae respectively;
Enp 2-segmented, bilobed distal segment with 1-2, 8-9+ 6-7 setae; Exp 7- segmented with 1, 2,
1, 1, 1, 1, 4 setae respectively. Md with well developed gnathobase, palp B with 4 setae; Enp 2-
segmented with 4 and 10 setae respectively; Exp 5 segmented with 1, 1, 1, 1, 2 setae
respectively. Mx1 Li1 with about 15 spines and setae, Li2-4 with 4-5, 4, 5-7 setae; Enp 2-
segmented with 6,9-12 setae; Exp with 11 setae; Le1 and 2 with 9 and 1 setae respectively. Mx2
Li1-5 with 5 (9 in P. xiphias), 3, 3, 3, 4 setae respectively; Enp 4-segmented with 3-4, 3, 2, 2
setae, or reduced. Mxp 7-segmented, C with 1, 2, 4, 4 setae; B with 3 setae plus 2 setae on the
incorporated Enp segment 1; free Enp 5-segmented with 4, 4, 3, 3, 4 setae respectively. P1-4
with both rami 3-segmented; Exp of P2-4 very large and flat, uter margin spines small. P1 B
inner seta situated on its anterior surface and passing across face of Enp segment 1. Enp
segment 1 of P2 typically incised and ornamented with 1 or more hook-like spinous
processes; Enp typically small, cylindrical. Spine and seta formula typically as follow but
sometimes reduced:
Males
Ur of 5 somites, sometimes strongly asymmetrical, single genital aperture located
ventrolaterally at posterior margin of Gns, usually on right side, sometimes on left. A1 usually
prehensile on left side only, occasionally on right side only. Mouthparts similar to those of
female. P5 asymmetrical, attached to plate formed from fusion of C and intercoxal sclerite;
right leg comprising B with an outer seta and 2-3 segmented Exp, Exp segment 2 with an
inner spinous process in some genera, Exp segment 3 with 1 or 2 minute distal setae; left leg
comprising B with outer seta and 2 or 3-segment Exp: Exp segment 1 bearing curved inner
process, distal segment swollen, often curved or claw-like.

Key to genera and species of Metridinidae

Females and males

1 Pd1 with anterolateral dark-pigmented spot on one side:………………....Pleuromamma

1a Pd1 without antenolateral dark-pigmented spot……………………………………………2
2 Posterior corners of Pr produced into long points; Ur of female symmetrically or
asymmetrically swollen; anal somite with every long posterolateral provesses:……….Gaussia
2a Posterior corners of Pr not produced into long points; Ur of female symmetrical, not
swollen; anal somite with short posterolateral processes: ……………………………….Metridia

Key to species of Metridia

Females

1 P5 2-segmented, asymmetrical: …………………………………………...Metridia venusta

1a P5 3- or 4-segmented…………………………………………………………………………...2
2 P5 3- segmented………………………………………………………………………………...3
2a P5 4-segmented…………………………………………………………………………………5
3 P5 segment 3 very small with only 2 terminal setae; posterior Pr rounded:
……………………………………………………………………………………Metridia brevicauda
3a P5 segment 3 larger and with 4 setae; posterior Pr rounded or pointed………………….4
4 Posterior Pr corners rounded; CR twice as long as wide:……..………….Metridia lucens
4a Posterior Pr corners rounded; CR about 3.5 times as long as wide:
…………………………………………………………………………..….Metridia gerlachei (Fig.16)
5 P5 C with row of long hairs; anterior Ce without anterior horns; CR about 5 times as
long as wide: ……………………………………………………………………….Metridia princeps
5a P5 C naked………………………………………………………………………………………6
6 P5 B with outer distal plumose seta; CR more than twice as long as wide………………7
6a P5 B with outer distal seta naked; CR less than twice as long as wide:
………………………………………………………………………………………..Metridia discreta
7 P5 segment 4 1/3 length of segment 3; segment 3 often without outer edge spine:
…………………………………………………………………………………….Metridia curticauda
7a P5 segment 4 more than ¾ length of segment 3; segment 3 with outer edge spine:
…………………………………………………………………………………………..Metridia longa

Males

1 Terminal segment of right P5 with long appendix, originating proximally and lying
parallel to outer border:…………………………………………………………...Metridia princeps
1a Terminal segment of right P5 without long appendix……………………………………..2
2 Right Exp1 of P5 without long, pointed inner extension:…..…………..Metridia venusta
2a Right Exp1 of P5 with long, pointed inner extension………………………………………3
3 Left P5 naroowed distally; left P5 rounded distally: ………………………Metridia longa
3a Both P5 rounded distally………………………………………………………………………4
4 Posterior corners of Pr rounded………………………………………………………………5
4a Posterior corners of Pr pointed or with small knob………………………………………...7
5 Right P5 with 1 thick, short inner spine on penultimate segment:...Metridia brevicauda
5a Left P5 with more than 1 short spine in penultimate segment…………………………….6
6 Left P5 with 3 inner spines on penultimate segment:……...…Metridia gerlachei (Fig.16)
6a Left P5 with 2 inner spines (one thick an one fine) on peultimate segment:
……………………………………………………………………………………….Metridia calypsoi
7 Posterior corners of Pr bearing minor knob:…………………..……...Metridia curticauda
7a Posterior corners of Pr pointed:.…………………………………………….Metridia lucens
Figure 16. Metridia gerlachei. Female length (mm): 3.38 – 4.25. Male length (mm): 2.16- 3.00.
Vertical distribution: Mesopelagic. Inshore/offshore distribution: Oceanic. Latitudinal
distribution: Antarctic.
Attachment

Calanoides acutus, Copepodite CII

Calanoides Acutus, Copepodite CIII.
Calanoides acutus, Copepodite CIV
Calanoides acutus, Copepodite CIV
Calanoides acutus, Copepodite CV
Rhincalanus gigas, Copepodite CIII
Rhincalanus gigas, Copepodite CIV