Occ Pub 15

GUIDE TO THE
COASTAL AND
SURFACE
ZOOPLANKTON OF
THE SOUTH-WESTERN
INDIAN OCEAN
David VP Conway
Rowena G White
Joanna Hugues-Dit-Ciles
Christopher P Gallienne
David B Robins
DEFRA Darwin Initiative Zooplankton Programme

Version 1 June 2003
Marine Biological Association of the United Kingdom

Occasional Publication No 15
GUIDE TO THE COASTAL AND SURFACE ZOOPLANKTON

OF THE SOUTH-WESTERN INDIAN OCEAN
David VP Conway
Marine Biological Association Plymouth
Rowena G White
University of Wales Bangor
Joanna Hugues-Dit-Ciles, Christopher P Gallienne and David B Robins
Plymouth Marine Laboratory
UK-DEFRA Darwin Initiative Project 162/09/004

Zooplankton of the Mascarene Plateau
Version 1 June 2003
Marine Biological Association of the United Kingdom

Occasional Publication No 15
General disclaimer
The authors, the Marine Biological Association and the Plymouth Marine Laboratory do not guarantee
that this publication is without flaw of any kind and disclaims all liability for any error, loss, or other
consequence which may arise from you relying on any information in this publication.
Citation
Conway, D.V.P., White, R.G., Hugues-Dit-Ciles, J., Gallienne, C.P., Robins, D.B. (2003). Guide to the
coastal and surface zooplankton of the south-western Indian Ocean, Occasional Publication of the
Marine Biological Association of the United Kingdom, No 15, Plymouth, UK.
Electronic copies
This guide is available for download, without charge, from the Plymouth Marine Laboratory Website at
http://www.pml.ac.uk/sharing/zooplankton.htm.
2003 by the Marine Biological Association of the United Kingdom and the Plymouth Marine
Laboratory, Plymouth, UK. No part of this publication may be reproduced in any form without
permission of the authors.
ISSN 0260-2784
CONTENTS
Introduction
Background to the Zooplankton programme
Sampling programme
Scope of this guide
Acknowledgements
Page 4
4
4
5
6
Phytoplankton and cyanobacteria
Protozoan Zooplankton
Phylum Sarcomastigophora
Radiolaria
Acantharia
Foraminifera
Phylum Ciliophora
Tintinnida
8
8
8
9
10
10
10
Eumetazoan Zooplankton
Phylum Cnidaria
Class Anthozoa
Class Scyphozoa
Class Hydrozoa
Sub-class Hydroidomedusae
Sub-class Siphonophorae
Phylum Ctenophora
Phylum Platyhelminthes
Phylum Nemertina
Phylum Rotifera
Phylum Mollusca
Phylum Annelida
Phylum Arthropoda
Subphylum Uniramia
Class Insecta
Subphylum Chelicerata
Class Pycnogonida
Subphylum Crustacea
Class Brachiopoda
Suborder Cladocera
Class Maxillipoda
Infraclass Facetotecta
Infraclass Cirripedia
Subclass Copepoda
Order Calanoida
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11
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12
13
13
25
38
39
40
41
42
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82
Order Cyclopoida
Order Harpacticoida
Order Poicilostomatoida
Order Monstrilloida
Order Siphonostomatoida
Class Ostracoda
Class Malacostraca
Order Stomatopoda
Order Mysida
Order Amphipoda
Order Isopoda
Order Cumacea
Order Euphausiacea
Order Decapoda
Phylum Phoronida
Phylum Bryozoa
Phylum Brachiopoda
Phylum Echinodermata
Phylum Chaetognatha
Phylum Hemichordata
Phylum Urochordata
Class Ascidiacea
Class Larvacea
Class Thalliacea
Subphylum Cephalochordata
195
208
218
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263
264
265
265
266
266
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320
320
321
Bibliography
322
The background to the cover of the guide is page 100, Pontellid copepods, from Dakin, W.J. &
Colefax, A.N. 1940. The plankton of the Australian coastal waters off New South Wales. Part
1. Publications of the University of Sydney, Department of Zoology.
INTRODUCTION
This guide has been compiled, as part of a UK DEFRA Darwin Initiative Project, as an aid to
researchers, particularly new researchers, in the identification of south-western Indian Ocean
zooplankton. It has been compiled because much of the identification literature covering this area
of amazing biodiversity is currently spread through the scientific literature and not easily
accessible without extensive library resources. The guide was produced in a limited time period,
so some specialist groups have not been dealt with in great detail. However, we hope to
gradually extend the guide to fill any gaps. We would welcome any comments and corrections
(please contact Dr Dave Conway at dvpc@mba.ac.uk).
Background to the DEFRA Darwin Initiative, Zooplankton Project
The Mascarene Plateau is a 2,200 km crescent-shaped ridge that runs roughly north-south from
the Seychelles Bank to the island of Mauritius. The ridges islands, banks and shoals form a
barrier modifying predominantly westward passage of the South Equatorial Current and waves
across the Indian Ocean. The ridge shallows to less than 30m in many areas of the banks at the
north of the plateau and places, where the depth of the plateau is less than 200m can be more
than 250km wide. The Mascarene Plateau is a rare example of an extensive shallow-shelf sea
completely detached from land boundaries, and is a large, almost completely unexplored marine
ecosystem.
Because it is a little studied region, the Royal geographical Society (with the institute of British
geographers) and Royal Society set up the Shoals of Capricorn marine programme. The
purpose of the research, coupled with education and training, was to help with sustainable
development and conservation of the marine and coastal environments of the Seychelles and
Mauritius, including the off lying island of Rodrigues. A second aim was to develop a better
picture of how the Mascarene ridge affects water flows and ecological processes in a key global
ocean-climate system the western Indian Ocean.
In April 2000, the UK (DEFRA) Darwin Initiative funded a joint training and research
programme between local agencies in Mauritius and Seychelles, the Shoals of Capricorn
programme and Plymouth Marine Laboratory with the Marine Biological Association. The
programme, called Biodiversity in the basement of the marine food web, was to focus on
the zooplankton of the Mascarene Plateau. One of the projects primary objectives was, for the
first time, to identify important zooplankton species around the Seychelles and Mauritius and start
local training and research programmes to monitor plankton diversity and abundance.
When the Shoals of Capricorn field programme formally came to an end in November 2001,
the Shoals bases were handed over to the host nations. In the Seychelles this was undertaken by
the new Seychelles Centre for Marine Research and Technology Marine Parks Authority
(SCMRT MPA) under the Ministry of Environment, and in Rodrigues through a new nongovernmental organisation, Shoals Rodrigues, associated with the Mauritius Oceanography
Institute (MOI). At the same time the management of the Darwin project was passed to the
Plymouth Marine Laboratory (PML).
The zooplankton Darwin Initiative project ended in 2003 with a regional conference and
workshop in the Seychelles to consolidate the three years of research and training, as well as to
help map out strategies for each regional team. The workshop was used to launch this Guide to
the coastal and surface zooplankton of in the South Western Indian Ocean.
Sampling programme
A weekly zooplankton sampling programme was started in September 2000 in both the
Seychelles and Rodrigues. The sampling equipment used is a conical 0.4 m diameter hand net of
125 m mesh aperture, fitted with a General Oceanics (GO) flowmeter. Because the stations
are shallow and over coral, the net is towed on a horizontal haul for at least 5 minutes at
approximately 5 m depth. Whenever possible, temperature and salinity measurements are taken
in association with the hauls and records kept of meteorological conditions. Zooplankton samples
are preserved in 4%, borax buffered formaldehyde. Zooplankton is being identified to as precise
a taxonomic detail as possible and counted using sub-sampling techniques.
Counting of samples is also being carried out at both sites using an Optical Plankton Counter
(OPC), which also gives zooplankton volumes, which can then be used to estimate biomass.
The plankton net mesh size of 125m was specifically chosen, as it samples organisms to the
lower end of the size discrimination of the OPC. This net also efficiently samples the small
copepods, which compose the bulk of the zooplankton biomass at the surface and in the coastal
regions of these tropical waters.
In the Seychelles three stations are sampled for zooplankton, one off the capital Victoria (outside
the immediate influence of the harbour), the second off the island of St Anne and the third in
more open water, outside the small island of Seche. In Rodrigues one station is sampled, in
water of 12 m deep, just beyond the reef edge, outside the shipping channel leading into the
capital, Port Mathurin. Sampling stations were chosen to be accessible, except in extreme
weather, and which were under the influence of both the reefs and open sea. Sampling is carried
out, for safety reasons, only during the day and close to high water, to standardise sampling
conditions as closely as possible.
Scope of this guide
This guide attempts to cover all pelagic zooplankton species sampled in the Seychelles and
Rodrigues. However, it also includes information from ancillary sampling carried out using a
Continuous Plankton Recorder (CPR) towed from HMS Beagle, south-east of the Seychelles,
from 0858'S, 5941'E to 0528'S, 5545'E in February 1999, as well as samples taken
between Seychelles and Rodrigues on the research yacht Zuza in April 2001 and from some
samples taken close to Aldabra in February 2001. We have also included species recorded from
inshore zooplankton studies in Madagascar (Furnestin & Radiguet, 1964; Gaudy, 1967; Binet &
Dessier, 1968; Patriti, 1970). This guide therefore includes a high proportion of the islandcoastal and surface water zooplankton of the south-western Indian Ocean. The location where a
particular species has been sampled has been noted. Some species, that have not been sampled
but that are known to occur in the region, have been included.
For some specialist groups, which we did not have time to fully identify, a selection of
representative members have been given, so that organisms can at least be assigned to perhaps a
particular genus within the main group. This particularly applies to some of the meroplankton
groups such as decapod larvae, which have many morphological changes between phases and
also between stages. For brevity, comprehensive taxonomic information has not been presented,
but we feel sufficient information is contained to identify each species. Keys have not yet been
included for genera, as further species will be added.
Classification of organisms is always in a state of flux, but for convenience here generally follows
the scheme set out by Brands (1989-2002). For crustacea, we have followed the classification
of Martin & Davis (2001).
Bibiography
A bibliography of relevant plankton references useful for identification of Indian Ocean species
has been included, separated where possible into different groups. Some references are not
specific to the Indian Ocean, but contain information on species that are recorded there.
ACKNOWLEDGEMENTS
We would like to acknowledge the UK-DEFRA Darwin Initiative (Project No. 162/09/004)
who provided funding for this work, the Royal Geographical Society (with the Institute of British
Geographers) and the Plymouth Marine Laboratory who have administered it, the original Shoals
of Capricorn team, particularly Juliet Burnett, Jessica Kavanagh, Iain Watt, Jan Robinson,
Martin Callow, Caroline Laughton, Tara Lynch, Sabrina Meunier, Tom Hooper and Jane Dodd,
who have all contributed to the project in different ways. We would also like to thank the
Seychelles Centre for Marine Research and Technology - Marine Parks Authority (SCMRTMPA), Shoals Rodrigues, the Mauritius Oceanographic Institute, the University of Mauritius, the
Rodrigues Ministry and the Albion Fisheries Research Centre for their continued support. In the
Seychelles, the work would not have been possible without the support of many of the staff from
the SCMRT-MPA, Coastguard Service, Meteorological Office, Public Utilities Corporation,
Seychelles Bureau of Standards and Ministry of Environment and Transport. We would also like
to acknowledge the National Marine Biological Library at the Marine Biological Association
Laboratory in Plymouth and the Wolfson Library at the University of Wales, Bangor, whose
comprehensive collections of literature and support from staff, greatly facilitated compilation of
this guide.
The guide could not have been prepared without the generous permission of journals, publishers
and individual authors to use the many figures. The source of each figure has been individually
acknowledged, but some sources we have relied on particularly heavily. In particular we would
like to thank Janet Bradford-Grieve, who has prepared and compiled figures and descriptions of
so many previously poorly described calanoid copepods of this southern area, for a large number
of excellent calanoid copepod figures, Shuhei Nishida for cyclopoid and calanoid figures, John
Mauchline for figures of copepod morphology, John Wickstead for figures of many of the smaller
zooplankton groups, Francesc Pags, Josep-Maria Gili and Jean Bouillon for their figures of
medusae and siphonophores from Scientia Marina, which were beautifully drawn by Jordi
Corbera, the University of California, Scripps Institution of Oceanography who gave permission
to use the wonderfully detailed drawings of chaetognaths by Angeles Alvario, the New Zealand
Oceanographic Institute for the drawings of chaetognaths by Sigrid Lutschinger and the
Zoological Museum Copenhagen who gave permission to use the mollusc figures from the Dana
Report. Annelies Pierrot-Bults gave helpful advice on the chaetognath section.
PHYTOPLANKTON AND
CYANOBACTERIA
The single celled plants are placed in the group we call phytoplankton. While phytoplankton
is not being counted in the present work, a few examples are given in case they are noticed in
samples. The two main groups which will be found are dinoflagellates, which have two thin
whip- like flagellae (usually only visible in living specimens) used in swimming and diatoms
which do not have flagellae. Examples of dinoflagellates are Noctiluca and Pyrocystis. These
are both luminescent when disturbed and can cause the silvery trails seen behind boats at night.
Other examples are Ceratium, many species of which are anchor-shaped and Peridinium which
is a stubby, spindle- shaped species. Dinoflagellates, when alive, can vary in colour from
brownish to green or red and can occur in high enough numbers to colour the sea.
Diatoms may be found singly, or because of the way they can reproduce, they can also be
found in long chains. Some chains can be straight as in Chaetoceros, or in a spiral as in
Asterionella. A common diatom called Coscinodiscus appears circular from above, but when
turned on its side is actually a circular box.
Trichodesmium is a blue-green algae which can occur in high numbers, typically found in
samples as a tangle of tiny elongated fibres, which may be in, bunches. It was first described in
Australian waters by Captain Cook in the 1700's who mistook the large brown blooms of
Trichodesmium as sandbanks. They are not "algae" at all, but in fact photosynthetic bacteria,
cyanobacteria. Trichodesmium is globally distributed in tropical and subtropical seas. They
have large gas vacuoles which enable it to regulate its position in the water column.
A
B
Examples of phytoplankton and cyanobacteria. A: Dinoflagellates; B: Diatoms; C:

Cyanobacteria (From Wickstead 1965).
PROTOZOAN ZOOPLANKTON
PHYLUM SARCOMASTIGOPHORA
Radiolaria
Radiolaria are protozoa, distinguished by segregation of their soft body into the central
capsule, containing the endoplasm, and the surrounding ectoplasm and by the siliceous
(opaline) skeletons of most species. They take their name from the radial symmetry, often
marked by radial skeletal spines, characteristic of many forms, although this symmetry is
sometime lacking. Skeletal elements of radiolarians, even the radially symmetrical ones, do not
actually meet at the center of the organism. This distinguishes them from the superficially
similar skeletons of acantharians, which are composed of celestite rather tha n opal. They are
generally found as single structures, although some form colonies. Some radiolarians can reach
up to 5 mm in diameter, which is very large for a single celled organism.
Representative radiolaria (From Haeckel, 1862, 1867).
Acantharia
These planktonic protozoa are now separated from the Radiolaria into a separate class. They
have skeletons of celestite formed by a set of spicules which, unlike Radiolaria, always meet at
the centre of the protoplasmic body.
Representative Acantharians (From Trgouboff & Rose, 1957).
Foraminifera
The shell of foraminifera may be made of organic compounds, sand grains and other particles
cemented together, or crystalline calcite. The skeleton is usually arranged in a series of
chambers of increasing size, the last open to the outside. Some appear like flattened snail shells
but there are a great variety of shapes, the simplest forms being open tubes or hollow sphere.
Globigerina is a very common foraminiferan in the tropics and is usually found as a group of
spheres of different sizes stuck together. They also have spines like a sea urchin, but these can
be missing.
In some species the protoplasm covers the outside of the shell. The largest species can have a
symbiotic relationship with algae. Food ranges from dissolved organic molecules, bacteria,
diatoms and other single celled phytoplankton, to small animals such as copepods. They move
and catch their food with a network of thin extensions of the cytoplasm similar to the
pseudopodia of an amoeba, although much more numerous and thinner.
Assorted foraminiferans
Examples of foraminferans (From Wickstead, 1965)
PHYLUM CILIOPHORA
Tintinnida
The main body is enclosed in a flask-shaped lorica composed of gelatinous material, usually
smooth, but in some groups embedded with fragments, giving a granular external appearance.
Cilia on the body are reduced or absent. In live animals a crown of cilia emerges from the flask,
sometimes interdispersed with small tentacles.
Examples of tintinnid shapes (From Daday, 1886)
10
EUMETAZOAN ZOOPLANKTON
The majority of zooplankton organisms sampled in the plankton nets will be eumetazoans, which are a
subkingdom of the animal kingdom comprising all animals except the sponges and the wormlike
mezozoans. Their bodies are composed of many cells, usually arranged in tissues and organs.
PHYLUM CNIDARIA
The Phylum Cnidaria (sometimes called Coelenterata) obtains its name from the stinging cells, called
cnidobasts which members of this phylum possess. Cnidarians are usually mainly composed of jelly
and includes the group of animals commonly called jellyfish. The planktonic forms generally swim in
pulses, by muscular contractions of part or all of their bodies, pumping water out of a cavity.
Within the Phylum Cnidaria there are three classes which have representatives in the plankton of the
south-west Indian Ocean region. They are separated using distinctive features in their bottom living
polypoid (or hydroid) or free-swimming jellyfish (or medusoid) phases:
Class Anthozoa - includes sea anemones and corals
Class Scyphozoa the large jellyfish
Class Hydrozoa there are five Subclasses in Class Hydrozoa, but only two have many
planktonic members, Subclass Hydroidomedusae which includes the small jellyfish and Subclass
Siphonophora, the siphonophores.
CLASS ANTHOZOA
Anthozoa means the flower animals and included in this group are corals and sea anemones. The
anthozoans have a polyp shape and medusae are never produced. Very few anthozoans occur in the
plankton, but corals have a larva called a planula which may be found for a short time in high numbers
in the plankton at particular times of year. The planula is flat, oval-shaped and may be up to around 1
mm in length. Additionally, some larvae of sea anemones are found in the plankton, particularly a
species called Cerianthus, which has a larva called an aracnactis. This has tentacles and resembles a
tiny sea anemone.
A, planula larva; B, aracnactis larva (From Wickstead, 1965).
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CLASS SCYPHOZOA
This class includes the easily recognisable large jellyfish, which, because they drift with the currents are
still considered to be planktonic. Thankfully, they are rarely caught in plankton nets, as the plankton
can get tangled in their bodies. The life cycle of scyphozoa either lacks a polyp stage, or they have a
larval polyp attached to rocks or other substrate that may develop directly into a single medusa, or
give rise to many medusae by budding off a small larva called an ephyra. The ephyra stage is often
found in plankton samples.
Young fish are often found associated with these large jellyfish (see the illustration of Chrysaora
below), using them for protection. They do not seem to be affected by the stinging cells and the
jellyfish are probably important for their survival and dispersal.
Ephyra larvae (From Wickstead, 1965)
Chrysaora hysoscella (Linnaeus, 1766)

Umbrella flat smooth and thick, 30 cm or wider in diameter. The colour is variable, but is
characterised by 16 v-shaped brown marks on the upper umbrella, radiating from the central region.
There are 24 marginal tentacles, which are easily broken off, and thirty-two pigmented semi-circular
marginal lappets.
Chrysaora hysoscella (Figure by J. Corbera, From Pags et al., 1992)
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CLASS HYDROZOA
These are either single (solitary), or form colonies and may have a life cycle which is exclusively
polypoid, exclusively medusoid, or with both phases.
Subclass Hydroidomedusae
This group includes the small jellyfish (or medusae), which occur in a variety of different shapes and
thickness of jelly, some flat others bell shaped. The commonest ones are budded of asexually from
flower-like hydroid polyps, which are found attached to rocks, seaweed etc. The free swimming
medusoid stage then reproduces sexually. A free swimming planula larva develops from the egg, which
settles and grows into a new hydroid polyp. Some members do not have a sessile bottom living stage,
but have a completely planktonic existence.
The medusoid stages have a central stomach which may be short, or on a stalk (peduncle). Tentacles
are arranged in bunches or spread round, at or close to the edge of the body in different combinations
of numbers and sizes, depending on the species. Ocelli coloured black, brown or red are sometimes
found on the base of the marginal tentacles. These are organs for light reception. Statocysts can also
be found round the umbrella margin. These are probably organs of orientation, informing the medusa
how it is positioned in relation to gravity. They can be internal open pits, closed vesicles or external
club shaped organs. The gonads are usually situated on the stomach, or on the radial canals which run
between the stomach and circumference of the medusa.
It is difficult to identify the medusae in their young stages, before adult characteristics develop. Some
of the most useful publications on hydroida are by Mayer (1910), Pags et al. (1992) and Russell
(1953). Hydroidomedusae have five main Orders, which are all represented in the south western
Indian Ocean - Anthomedusae, Leptomedusae, Limnomedusae, Trachymedusae and
Narcomedusae. Few specimens in good condition have been sampled. However, a guide to some of
the species likely to be present has been obtained from a survey done in the Seychelles of the bottom
living hydroid stages (Millard and Bouillon, 1973). Examples of a variety of species are given, even
though many of them have not yet been found in the samples.
Gonad
Stomach lips
Stomach
Marginal tentacle
Ring canal
Ocellus
Tentacle bulb
Marginal vesicle
Marginal
tentacle
A, Structure of medusae; B, Details of umbrella margin (From Russell, 1953).
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Order Anthomedusae
Usually have a deep, bell-shaped body (umbrella), but very variable in shape. The gonads are usually
situated on the stomach or occasionally both on the stomach and extending a short distance along the
radial canals. They have no statocysts.
Some anthomedusae produce an actinula larva, which can be found in the plankton. The larva has
tentacles and resembles a small sea anemone, although the tentacles are not at the end of the body, but
mid-way down.
Actinula larvae (From Wickstead, 1965).
Family Tubulariidae
Euphysa sp.
The umbrella is barrel-shaped with a rounded apex; jelly thick especially at apex. Stomach large and
cylindrical, two-thirds the length of the umbrella cavity, at full extension never reaching beyond the
umbrella margin. The mouth is simple and circular. There are 4 narrow ring canals, one single short
marginal tentacle and 3 marginal bulbs without tentacles. In good specimens the tentacle can be seen
to be covered in rings of nematocysts. A small medusae, around 1.4-6.0 mm in height.
Recorded: Seychelles (Millard and Bouillon, 1973), as Corymorpha sp. It is uncertain which genera
this specimen belonged to, so Euphysa is given as an example of the Tubulariidae.
Euphysa aurata (Figure by J. Corbera, From Pags et al., 1992)
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Family Zancleidae
Zanclea costata Gegenbaur, 1856.
Umbrella bell-shaped, about as high as wide, with quite thick jelly. 2 or 4 marginal tentacles. Stomach
cylindrical, never reaching beyond the umbrella margin. Mouth simple, 4 straight radial canals. Can
have two opposite marginal tentacles and 2 opposite marginal bulbs without tentacles, or 4 marginal
tentacles. Umbrella height usually 1.5-3.0 mm.
Recorded: Seychelles (Millard and Bouillon, 1973) as Zanclea? costata.
Zanclea costata. Different development stages (From Russell, 1953).
Family Cladonemidae
Cladonema radiatum Dujardin, 1843.
Very distinctive, creeping and swimming medusae. Umbrella bell-shaped, slightly higher than broad.
Stomach spindle-shaped usually with 5, sometimes 4 pouch-like outgrowths in the central section.
Mouth with 5, sometimes 4 protuberances armed with stinging cells. Stomach not extending beyond
umbrella margin. 5 sometimes 4 radial canals, some of which can branch forming 8 or 10 canals.
Usually 8, sometimes 10 marginal tentacles, corresponding to the number of radial canals. Marginal
tentacle branched, with ocelli on tentacle bulb. Height up to about 4 mm.
Recorded: Seychelles (Millard and Bouillon, 1973).
Cladonema radiatum. Different developmental stages (From Russell, 1953).

15
Family Clavidae
Turritopsis nutricula McCrady, 1859
Umbrella bell-shaped, slightly higher than wide composed of thin jelly. Stomach cross-shaped in
section, not reaching beyond umbrella margin. 4 radial compact masses of vacuolated cells above the
stomach. Mouth with 4 lips, with many circular masses of stinging cells along its edge. 4 radial canals.
80-90 marginal tentacle, close together in a single row. Small ocelli on tentacle bulbs. Height 4-5 mm.
Turritopsis nutricula. (From Mayer, 1910).
Family Hydraciniidae
Podocoryne sp.
Umbrella bell-shaped with rounded apex. Stomach not extending beyond stomach margin. Gonads on
stomach wall or extending slightly onto radial canals. Four radial canals. Mouth with 4 simple or
branching lips with circular nematocyst clusters. 4, 8 or more marginal tentacles. No ocelli on tentacle
bulbs.
Recorded: Seychelles.
Podocoryne spp. A, (From Mayer, 1910); B, (From Russell, 1953).
16
Family Rathkeidae
Rathkea sp.
Apex of umbrella thick and high. Stomach not extending beyond umbrella margin; with short peduncle.
Mouth with 4 lips with circular clusters of nematocysts on the extension of the lips and also between. 4
radial canals Gonads completely surround the stomach. 8 marginal tentacle bulbs, each with up to 5
perradial tentacles and 3 interradial. No ocelli.
Recorded: Seychelles Rathkea sp. (Millard and Bouillon, 1973).
Rathkea sp. A, view from above; B, oral tentacles; C, marginal tentacle and bulb (From Russell, 1953).
Family Bougainvillidae
Rounded body with thick jelly, especially at the apex. Marginal tentacles arranged in four clusters of
similar structure. Number of tentacles variable depending on species and stage of development. Ocelli
on tentacle bulbs. 4 radially placed oral tentacles which branch several times, depending on species
and stage of development. 4 radial canals. Gonads on the sides of the stomach. Some immature
specimens have been sampled in Seychelles.
Bougainvillia macloviana Lesson, 1830
Cylindrical umbrella with rounded apex and thick jelly. Stomach with pyramidal peduncle and broad
base. The 4 oral tentacle divide 5-7 times. Gonads slightly folded, extending along stomach. 4 radial
canals, wide with slightly ragged edges. 4 broad tentacle bunches, each with up to 60 tentacles. A
dark red ocellus on the base of each tentacle.
Bougainvillia macloviana Different development stages. A, (From Russell, 1953); B, (Figure by J.

Corbera, From Pags et al., 1992)
17
Family Pandeidae
Leukartiara octona (Fleming, 1823)
The umbrella is bell-shaped, up to around 15 mm in height, with a thick apical process that can vary in
size. The stomach is large, occupying up to half of the stomach cavity and is connected to 4 broad
radial canals, which have jagged outlines. The mouth lips are very frilled. The gonads are typically
orange in colour with transverse folds. Up to 23 long, hollow marginal tentacles, each with a spur.
Between each of these tentacles are one to three smaller secondary tentacles. Each tentacle has an
ocellus on the base.
Recorded: None recorded yet, but L. octona is given as an example of this family.
Leukartiara octona (Figure by J. Corbera, From Pags et al., 1992).
Family Porpitidae
Velella velella (Linnaeus, 1758)
The characteristic blue, asexual, polyp bearing colony with its upright sail, is found floating on the sea
surface and may be sampled as the net is at the surface. The chitinous float/base can be up to 37 mm
long and has a triangular thin sail, set perpendicularly.
Recorded: Mauritius, stranded on beach.
Velella velella Colony. (Figure by J. Corbera, From Pags et al., 1992)
18
Order Leptomedusae
These have hemispherical or flattened umbrellas. The gonads are only found on the radial canals,
though sometimes connecting onto the stomach. With or without statocysts.
Family Campanulariidae
Clytia sp.
Jelly thin, almost hemispherical. Stomach short, without peduncle, with four simple or frilled lips. Four
oval or straight gonads, extending over half to three quarters of the radial canals, up to 70 marginal
tentacles with prominent bulbs. 1-3 marginal vesicles between each pair of tentacles.
Recorded: Seychelles Clytia sp. (Millard and Bouillon, 1973).
Clytia sp. (Figure by J. Corbera, From Pags et al., 1992).

Obelia sp.
Circular umbrella, flat and thin. Stomach short and with four simple lips. Four straight radial canals
with oval or circular gonads on the canals, situated towards the outside. Many short marginal tentacles
with basal bulbs. Eight statocysts on umbrella margin.
Obelia sp. (Figure by J. Corbera, From Pags et al., 1992)
19
Family Lovenellidae
Lovenella sp.
Umbrella hemispherical with thin jelly. Small stomach without peduncle. 4 simple radial canals. The
gonads, are sausage shaped in adults, but more circular in young stages, on the radial canals, separate
from the stomach. The marginal tentacles have small cirri beside the tentacle bulbs. Without ocelli.
Recorded: Seychelles Lovenella sp. (Millard and Bouillon, 1973).
Lovenella sp. A, marginal tentacle, cirri and vesicles; B, gonad and marginal tentacle; C, adult medusa
(A, B From Russell, 1953; C Figure by J. Corbera, From Pags et al., 1992).
20
Family Phialellidae
Phialella quadrata
Umbrella nearly hemispherical; jelly fairly thick. Elongated gonads on outer third of the 4 radial canals,
but not reaching the margin. 16-32 marginal tentacles with globular bases. No ocelli. 8 marginal
vesicles. Stomach short with no peduncle. Mouth with 4 short lips, with slightly folded margins. Many
specimens have 4 black spots at the base of the stomach, visible from above, one in each interradius.
Diameter of umbrella up to 13 mm.
A
C
D
Phialella quadrata. A, B, different development stages; C, gonad and marginal tentacle;

D, Marginal tentacle and vesicle (From Russell, 1953).
21
Order Limnomedusae
Most with thick bell-shaped umbrellas, with gonads either only on stomach, on stomach extending for
a short distance along the radial canals, or only on the radial canals. With or without statocysts.
Family Proboscidactylidae
Proboscidactyla stellata (Forbes, 1846)
Umbrella dome-shaped around 8 mm high and 9 mm wide, thick at the apex. Stomach short with a
hexagonal base and six lobes. Mouth with six frilled lips. Six radial canals, each dividing twice, so that
24 canals reach the umbrella margin, although this number is variable. The gonads are located at the
base of the stomach wall and extend over the six stomach pouches. There are twenty-four marginal
tentacles and a broad velum in well preserved specimens.
Recorded: Not yet recorded, but given as an example of the Family.
Proboscidactyla stellata. (Figure by J. Corbera, From Pags et al., 1992)
22
Order Trachymedusae
Hemispherical or deep, bell-shaped umbrella, gonads only on the radial canals, with statocysts.
Family Geryonidae
Liriope tetraphylla (Chamisso and Eysenhardt, 1821)
Hemispherical umbrella, up to 30 mm in diameter, with thick jelly, especially on the apex where it can
be slightly flattened. Small stomach on elongated peduncle of varying length. 4 straight radial canals
with large flattened leaf-like gonads of varying shape, sometimes almost covering the whole under
surface. 4 large marginal tentacle and four small interradial. 8 statocysts and a broad velum in well
preserved specimens.
Recorded: Seychelles, Zuza, Aldabra, Rodrigues.
Liriope tetraphylla (Figure by J. Corbera, From Pags et al., 1992)
Family Rhopalonematidae
Aglaura hemistoma Pron and Lesueur, 1810
The umbrella is conical with a flattened apex in which there is a small depression. The umbrella is
octagonal in cross-section with narrow longitudinal ridges (although this may not be obvious in poorly
preserved specimens), composed of thin jelly and up to around 6 mm in height. The short stomach is
on a thin peduncle. Eight sausage-shaped gonads are suspended from the peduncle above the
stomach. There are eight radial canals running along the ridges on the umbrella and around 50-75
short marginal tentacles, which are easily damaged and usually partially missing. The velum can be very
broad.
Recorded: Rodrigues, Seychelles, Zuza
Aglaura hemistoma (Figure by J. Corbera, From Pags et al., 1992)

23
Order Narcomedusae
With thick upper umbrella, no radial canals and with the gonads only on the broad circular stomach.
With statocysts. The marginal tentacles do not emerge on the edge of the umbrella, but some distance
above.
Family Aeginidae
Solmundella bitentaculata (Quoy and Gaimard, 1833)
The umbrella is bell-shaped, up to 12 mm high by 15 mm wide. The jelly is thick, especially in the
upper half. The stomach is broad, circular and short with a simple circular mouth. There are 8
rectangular stomach pouches on which the gonads are located. There are 2 long tentacles emerging on
either side of the body from above the stomach.
Recorded: Seychelles, Zuza, Aldabra, Madagascar, Rodrigues.
Solmundella bitentaculata. (Figure by J. Corbera, From Pags et al., 1992)
24
Subclass Siphonophorae
The siphonophores are usually composed of clear jelly, but are completely different in structure from
the free-swimming stage of hydrozoan jellyfish. Siphonophores do not have an alternation of
generation between an attached, asexual polyp stage and a free-swimming sexual medusa like most
other hydrozoans, but modifications of both stages are found, attached to the free-swimming organism.
Different siphonophores species can have a very different appearance and they are often made up of
many different parts which usually all fall apart when caught in a plankton net, so it is often difficult to
estimate the number of complete individuals in a sample. Because of their complicated life cycles, the
fragments we find in the samples can often only be accurately identified by a specialist. Thus, in many
instances we can only note down the general group to which they belong, or simply unidentified
siphonophore pieces. Because many of the structures found in siphonophores are only found in these
organisms, a unique terminology has been developed which can be daunting to people trying to identify
them.
Some useful works on siphonophores, which include species from the south western Indian Ocean
area are by Kirkpatrick and Pugh (1984), Mackie et al. (1987), Pags and Gili (1992), Pugh (1999),
Rengarajan (1973) and Totton (1954, 1965).
The Subclass Siphonophorae is divided into three Orders, Cystonectae, Physonectae and
Calycophorae, on the basis of whether or not they have a gas-filled float called the pneumatophore,
or swimming bells called nectophores.
Order Cystonectae
Siphonophores in Sub-order Cystonectae have no nectophores, only a large gas filled
pneumatophore.
Family Physaliidae
Physalia physalis (Linnaeus, 1758)
Commonly known as the Portuguese Man of War, this species with its large float is the only member
of this family likely to be found in the region. It may be familiar to some people who have seen it
floating on the sea surface or stranded on a beach, but hopefully, because of the painful stings it can
give, will not often be caught in plankton nets.
Recorded: Mauritius, stranded on rocks.
Physalia physalis (Figure by J. Corbera,

From Pags et al., 1992)
25
Order Physonectae
There are only a few members in Order Physonectae. They have a pneumatophore and swimming
bells and a long stem forming a tube through which food and body fluids pass and to which the
different body parts are attached in two regions. The nectosome region is situated just below the
pneumatophore and usually only bears asexual (non-reproductive), medusoid nectophores which are
swimming bells used in jet propulsion. The morphology of the nectophores is an important
characteristic used in identification and as most physonectids have the same basic design, they are
easily distinguished to this group. The nectophores never have a somatocyst, an organ that is part of
the digestive system in Calycophoran siphonophores.
Below the nectosome lies the siphosome , which is usually the longest part of the body. Here both
polypoid and medusoid structures are attached to the stem, which have different functions, arranged in
a succession of groups called cormidia. The cormidium consists of a single gastrozooid, which is
used, in feeding and digestion. The gastrozooids usually have a tentacle and several small tentacles
called tentilla, which bear stinging cells, used in capturing prey. Palpons, which are modified
gastrozooids, are also usually present on the cormidia and may have a sensory or excretory function.
These also may have a tentacle attached. Another part is the gonodendron, which bears the
developing sexual medusoids or gonophores, which can be both male and female in the same
cormidium. The cormidia can also bear asexual nectophores and bracts composed of thick jelly, the
latter probably for buoyancy or protection of the other structures.
The complete animal is referred to as the polygastric stage. In most physonectids the cormidia
remains attached to the stem, although the gonophores which produce eggs and sperm can become
detached. In physonectids only one egg is produced in each gonophore and fertilisation is external. A
planuloid larva is produced which eventually develops a float, then progressively the other features.
Pneumatophore
Medusoid nectophores
Stalk
Nectosome
Tentacle
Siphosome
Bract
Physonectid colony, Agalma sp., showing Structure (From Trgouboff & Rose, 1957).
26
Family Agalmatidae
Agalma elegans (Sars, 1846)
This is one of the few physonectid species occurring in the region. Early A. elegans specimens, usually
less than 4 mm in length, are sometimes sampled and can be recognised as a young physonectid by the
presence of a pneumatophore with a cap tinged dark red. They have a ring of 4-6 thick, triangular
serrated bracts and a single gastrozooid and tentacle.
Adult specimens, which will only be represented by fragments in the plankton samples, have a terminal
pneumatophore, approximately 1 mm in height with a red cap. Often the siphosomal stem is found on
its own with the pneumatophore still attached. The nectophores are up to 7 mm in length, v-shaped
and have two prominent lateral wings and an internal t-shaped nectosac. The bracts are triangular,
approximately 9 mm in length with three terminal projections and a convex dorsal surface.
Recorded: Rodrigues, Aldabra.
Agalma elegans. A, nectophore; B, bract (Figures by J. Corbera, From Pags et al., 1992)
27
Order Calycophorae
Calycophoran siphonophores do not have a pneumatophore and are the siphonophores group with by
far the most members. The nectophores are often very specialised and their structure is variable
between different species. The number of nectophores present can be from 1-15, depending on
species, all appearing quite similar. Usually the stem groups of the polygastric phase, the cormidia,
break off successively to form the free swimming sexual eudoxid stage. They have a somatocyst, an
organ associated with digestion.
Family Prayidae
Prayids are large, slow swimming siphonophores, generally rounded with thick jelly. When the jelly is
thick, the small thin somatocyst may branch complexly.
Sub-family Prayinae
Rosacea cymbiformis (Chiaje, 1882).
The asexual polygastric stage is composed of 2 simple rounded nectophores up to 17.5 mm in height,
attached ventrally, although usually broken apart in plankton samples. One nectophore partially
surrounds the other. The jelly is very thick giving buoyancy to the animal. The inner cavity, the
nectosac, which is used in propulsion, is small, so the animal is slow swimming. The nectosac is less
than two-fifths the height of the nectophore. The nectophores have a shallow furrow called the
hydroecium, extending the length of the ventral surface, inside which the siphosomal stem is attached,
so the region of the stem where budding of the cormidia takes place is protected from damage. The
stem can be drawn up into the hydroecium, both for protection and to streamline the body during
swimming. Alongside the hydroecium runs the somatocyst. The polygastric stage buds off and releases
the sexual cormidia, which develop into the eudoxid stage.
Recorded: Aldabra, Madagascar.
Rosacea cymbiformis. (Figure by J. Corbera, From Pags et al., 1992)

28
Family Diphyidae
This group have streamlined bodies with a large nectosac, features related to their generally fast
swimming speed. They usually only have one or two nectophores present at one time, in both the
asexual polygastric and sexual eudoxid stage. The upper nectophore is called the anterior
nectophore. The second nectophore if present has a different structure and is called the posterior
nectophore. The anterior nectophore of the polygastric stage is the most useful for identification and is
the main part described and figured here. It can be identified by the presence of the sac-like
somatocyst. The shape of the somatocyst and hydroecium and the number of ridges (if any) on the
nectophore are particularly useful for identification.
Ridge
Anterior nectophore
Radial canal
Nectosac
Somatocyst
Hydroecium
Posterior nectophore
General Diphydae structure, polygastric, stage, showing nectophores separated (Figures by J.

Corbera, From Pags et al., 1992).
29
Subfamily Sulculeolariinae
The anterior nectophore of the polygastric phase is smooth and rounded, without ridges.
Sulculeolaria turgida (Gegenbaur, 1853)
The polygastric stage, anterior nectophore, is conical with a rounded apex, reaching up to 15mm in
height. There are no teeth on the ostial margin. Tiny somatocyst, one twentieth of the height of the
nectophore. The basal lamella is quite long, divided into two sections by a deep furrow.
Recorded: Zuza, Madagascar.
Sulculeolaria turgida A anterior nectophore; B, posterior nectophore (Figures by J. Corbera, From

Pags et al., 1992).
Sulculeolaria chuni (Lens and van Riemsdijk, 1908)

The anterior nectophore of the polygastric stage has a rounded apex, no ridges no teeth on the ostial
margin and is up to 8mm in height. The somatocyst is quite long and sausage-shaped, two to three
fifths of the height of the nectophore. The basal lamella is short, divided into two sections.
Recorded: Rodrigues, Zuza, Madagascar.
Sulculeolaria chuni. A, anterior nectophore; B, posterior nectophore (Figures by J. Corbera, From

Pags et al., 1992)
30
Subfamily Diphyinae
Nectophores of all but a few species are ridged.
Diphyes bojani (Eschsoltz, 1829)
The anterior nectophore of the polygastric phase has five long ridges, which are serrated, to varying
degrees. Size is up to approximately 10 mm in height and 3 mm in width. There are 3 ostial teeth of
approximately the same length. The nectosac reaches nearly to the top of the nectophore. The
hydroecium is very deep, generally reaching almost half way up the nectophore. The somatocyst is
sausage shaped, tapering towards its apex and reaches almost to the end of the nectosac.
Recorded: Aldabra, Zuza, Madagascar.
Diphyes bojani. Anterior nectophore (Figure by J. Corbera, From Pags et al., 1992)
Diphyes chamissonis Huxley, 1859.
In the polygastric phase there are five serrated ridges on the anterior nectophore, which is
approximately 5 mm in height and 1.6 mm wide. The hydroecium is deep and the somatocyst is club
shaped and approximately half the length of the nectosac. The basal lamella is deeply indented. There
are 3 ostial teeth of the same size. A posterior nectophore does not develop.
Recorded: Rodrigues, Seychelles. Aldabra, Madagascar.
Diphyes chamissonis. Anterior nectophore

(Figure by J. Corbera, From Pags et al., 1992)
31
Diphyes dispar Chamisso and Eysenhardt, 1821

The anterior nectophore of the polygastric phase is large, up to 36 mm in height, by 18 mm wide.
There are 5 ridges, with only one slightly serrated. The teeth on the basal lamella closest to the opening
of the nectosac are larger than the outer ones. There are 3 ostial teeth. The hydroecium is very deep.
The nectosac tapers to a thin process, which then extends almost to the apex of the nectophore. The
somatocyst is sausage shaped and varies in length, but does not reach the end of the nectosac.
Recorded: Rodrigues, Seychelles, Zuza, Madagascar.
Diphyes dispar. A, anterior nectophore; B, posterior nectophore (Figure by J. Corbera,

Lensia subtilis (Chun, 1886).

The anterior nectophore of the polygastric phase is small, fragile, rounded at the apex and flattened
laterally. It does not have any ridges and is up to 10 mm in height. The hydroecium is short and slants
outwards. The somatocyst has a long thin stalk terminating in a small ball. The basal lamella is small
with two wings and runs obliquely outwards.
Recorded: Zuza, Aldabra, Madagascar.
Lensia subtilis. Anterior nectophore (Figure by J. Corbera, From Pags et al., 1992)
32
Lensia subtiloides (Lens and van Riemsdijk, 1908).

The anterior nectophore of the polygastric phase has 5 ridges and is approximately 3.7 mm high and
1.4 mm wide. The hydroecium is very shallow, reaching the level of the base of the nectosac. The
basal lamella slopes slightly upwards towards the outside of the nectophore, with two short rounded
wings. The somatocyst is club-shaped and about one third the length of the nectosac.
Recorded: Seychelles, Aldabra, Madagascar.
Lensia subtiloides. Anterior nectophore (Figure by J. Corbera, From Pags et al., 1992).
Lensia hotspur Totton, 1941

The anterior nectophore of the polygastric phase has 5 complete ridges and is up to 7.2 mm in height
and 2.3 mm in width. The hydroecium is very shallow and does not reach the bottom of the nectosac.
The basal lamella is quite flat, divided into 2 downwardly slanting wings. The somatocyst varies in
shape, but tends to be short and broad with a short stalk and sloping outwards.
Recorded: Aldabra, Madagascar.
Lensia hotspur. Anterior nectophore (Figure by J. Corbera, From Pags et al., 1992)
33
Chelophyes appendiculata (Eschsoltz, 1829).

The anterior nectophore of the polygastric stage is very rigid, up to 20 mm in height. There are 5
ridges but only 3 reach the apex. The hydroecium is quite deep, directed outwards. The basal lamella
is divided into 2 square wings. The somatocyst is long and spindle shaped, varying between narrow
and thin, reaching two-thirds to three-quarters the length of the nectosac.
Recorded: Rodrigues, Zuza, Madagascar.
Chelophyes appendiculata. A, Anterior nectophore; B, Posterior nectophore (Figure by J. Corbera,

Chelophyes contorta (Lens and van Riemsdijk, 1908).
The anterior nectophore of the polygastric phase is similar in shape and rigidity to that of C.
appendiculata. It is up to 4.6 mm in height by 1.7 mm in width. 5 slightly serrated ridges on
nectophore, 1 very short, only 3 reaching the apex. Somatocyst characteristically twisted to the right,
as are also the ridges close to the somatocyst. Somatocyst is around two thirds of the length of the
nectosac. Hydroecium bends outwards, not as deep as in C. appendiculata. Basal lamella divided
into two wings with a small tooth on either side in the lower central part.
Recorded: Rodrigues, Seychelles, Zuza, Aldabra, Madagascar.
Chelophyes contorta. A, anterior nectophore; B, anterior nectophore side view; C, Posterior

nectophore (Figure by J. Corbera, From Pags et al., 1992).
34
Eudoxoides spiralis (Bigelow, 1911).

The polygastric anterior nectophore (no posterior nectophore develops) is rigid and characteristically
twisted. It is up to 12 mm in height. There are 5 toothed ridges, only 4 reaching the apex. The
hydroecium is deep with a rounded apex. Basal lamella divide into 2 pointed wings, one longer than
the other. The somatocyst reaches half way up the nectosac.
Eudoxoides spiralis. Anterior nectophore (Figure by J. Corbera, From Pags et al., 1992).
Eudoxoides mitra Huxley, 1859.
The polygastric stage anterior nectophore is rigid and not spiralled, up to 7.4 mm high by 2.5 mm
wide. There are 5 ridges on the nectophore which all reach the apex, serrated on the lower
nectophore. Hydroecium has a rounded apex. The basal lamella are divided into 2 serrated wings, the
left wing with a central tooth-like projection. The somatocyst is sausage shaped, starting off very
narrow and reaches halfway up the nectosac.
Eudoxoides mitra. A, anterior nectophore; B, posterior nectophore; C, eudoxid stage (Figures by J.

Corbera, From Pags et al., 1992)
35
Family Abylidae
The anterior nectophore of the polygastric stage is much smaller than the posterior nectophore.
Subfamily Abylopsinae
Nectophores very rigid and the anterior nectophores usually beautiful box-like structures with many
ridges or facets, like a crystal.
Abylopsis eschscholtzi (Huxley, 1859).
The polygastric stage is rigid with pronounced serrated edges. The wide somatocyst has a short
projection at the top.
Recorded: Rodrigues, Seychelles, Aldabra, Madagascar.
Abylopsis eschscholtzi. A, complete polygastric stage; B, eudoxid stage (Figures by J. Corbera,

36
Bassia bassensis (Quoy and Gaimard, (1833) 1834).

The polygastric anterior nectophore is very similar to that of A. eschscholtzi, but the somatocyst has
no apical projection.
Recorded: Rodrigues, Seychelles, Aldabra, Madagascar, Mauritius.
Bassia bassensis. A, complete polygastric stage; B, eudoxid stage; C, anterior nectophore (Figures by
J. Corbera, From Pags et al., 1992).
37
PHYLUM CTENOPHORA
Ctenophores are also composed of jelly, but because of their very individual structure, have a
phylum all to themselves. They are not common in the tropics but occur regularly in samples.
They do not have stinging cells like cnidarians; many such as Pleurobrachia capture their prey
with sticky adhesive cells on their tentacles, while the bell-shaped Beroe captures prey inside
the bell. They swim by cilia, but the characteristic of this group is that the cilia are joined
together in rows like a comb - which is why they have been given the common name of combjellies. Velamen whose common name is Venus's girdle, is a beautiful ribbon- like ctenophore,
but unfortunately, like many of the ctenophores, it breaks up when caught in the plankton net
and can only really be observed by divers.
Pleurobrachia sp.
Mnemiopsis sp.
Bere sp.
Velamen sp.
Examples of ctenophores (From Wickstead, 1965).
38
PHYLUM PLATYHELMINTHES
Order Turbellaria
The Order Tubellaria are commonly called the flatworms. Only low numbers of a few species
are found in the plankton and most species are less than 5 mm in length. They are generally
flattened and leaf shaped, with rounded corners, although often curled up in the samples. Their
bodies are opaque and cream coloured, certainly in preserved specimens, and the internal
organs are not usually visible. Externally they usually have two clusters of black pigment spots
on their upper surface, associated with light sensitivity. The Turbellaria are free living or
commensal and there are around 3,000 known species, most of which are marine. They are the
most primitive of the Platyhelminthes.
Typical Turbellarian shapes.
39
PHYLUM NEMERTINA
This Phylum has been spelt in different ways by different authors - Nemertini, Nemertinea,
Nemertea or Nemertina.
Nemertines are wormlike organisms, the adults of which are generally found on and in the
sea bottom, although some species are deep-water pelagic species. They are commonly
called ribbon-worms as some are very long, or sometimes proboscis worms, because they
can extend a proboscis to capture their food. Most look like flatworms, but tend to be
larger, thicker bodied, and longer. Most species are less than 20 cm in length, and some just
a few millimetres long, but a few species may measure several metres or more. Most are
pale, but some are brightly coloured with patterns of yellow, orange, red and green.
Their free-swimming, ciliated pilidium larvae can occasionally be common in plankton
samples..
Pilidium larvae (From Wickstead, 1965).
40
PHYLUM ROTIFERA
Rotifers are small (0.1-0.5 mm) flask-shaped organism, with a characteristic crown of fine hairs
or cilia surrounding the mouth area. These cilia beat together in waves to draw water currents
(and thus food) toward their mouth. Rotifers primarily eat phytoplankton, bacteria, and also
detritus. Some will occasionally eat another rotifer if it is small enough. Rotifers are commonly
cultured in fish farms to feed to larval fish as their first food.
Examples of rotifers (From Berzins, 1960a,b).
41
PHYLUM MOLLUSCA
The Phylum Mollusca comprises the chitons, abalones, snails, clams, squid and octopus as well
as many other less well known groups. Molluscs range in size from a few mm through to
several metres in length for the giant squid (the largest living invertebrate). The bodies are soft
and partly or wholly covered by a mantle, a sheet of tissue exclusive to molluscs. The majority
of species have a protective shell, usually external, that is excreted by the mantle. In a few
species the shell is internal or absent altogether. Locomotion is usually achieved using a
muscular foot, this may be modified for burrowing or swimming. The majority of molluscs are
benthic, a few species have adapted to a free-swimming existence (e.g. squid, heteropods,
pteropods, janthinids and a few nudibranchs).
Molluscs may be dioecious (i.e. sexes are separate) where fertilisation can be internal or
external, they may be simultaneous hermaphrodites (i.e. produce eggs and sperm
simultaneously), or protandric hermaphrodites (i.e. change sex with age). Development may be
direct, or through two planktonic larval stages, a trochophore and a veliger larva. The molluscan
trochophore is similar to the that of segmented worms. The typical trochophore is small, topshaped, with a small mouth opening just below an equatorial ring of cilia. The trochophore
stage may be passed through in the egg. The trochophore is relatively short-lived and passes
into the exclusively molluscan larva, the veliger. The veliger characteristically possesses a shell
and a densely ciliated velum of between one and six lobes. The ciliated lobes are used for
locomotion, feeding and respiration and can be withdrawn into the protective shell. The time
the larva spends in the plankton varies with species, but can be anything from a few hours to
over a year for some cone shells and cowries. In addition to the trochophore and veliger larva,
there is an unusual echinospira larva typical of a few families of Gastropoda (e.g.
Lamellariidae, Eratoidae and Capulidae). The individual is housed in a large transparent,
sometimes ornately ridged, spiral shell.
The Phylum Mollusca is large and the taxonomic relationships between many of the species is
under review. The Phylum is subdivided into seven classes:
Class Monoplacophora. A group of small, limpet-like, deep-sea molluscs. The sexes are
separate and fertilisation is usually external. The group is not included in the guide.
Class Aplacophora. A group of primitive, wormlike molluscs without shells. Fertilisation is
usually external. The group is not included in the guide.
Class Polyplacophora. Chitons or coat-of- mail-shells.
Class Scaphopoda. Tusk-shells.
Class Gastropoda. The largest class of snails, sea slugs, sea butterflies. Further subdivided into
three Subclasses, Prosobranchia, Opisthobranchia and Pulmonata.
Class Pelecypoda. Bivalves.
Class Cephalopoda. Squid, cuttlefish, octopus and nautilus.
42
CLASS POLYPLACOPHORA
The chitons, or coat of mail shells. Polyplacophora (in Greek meaning many plate bearing)
have a characteristic shell consisting of a series of eight overlapping plates covering the dorsal
surface. Chitons usually live close inshore on hard surfaces. Sexes are separate and fertilisation
is external. There is no veliger larval stage. The planktonic trochophore develops directly into a
juvenile chiton.
Chiton development stages. A, trochophore; B, later trochophore; C, juvenile; D, adult. Figures

A, B from Christiansen (1925); D from Pilsbry (1892, 1893).
CLASS PELECYPODA
The Pelecypoda have shells of two valves, bivalves. The majority are benthic, either attached to
structures or burrowing in the substratum. The typical bivalve releases eggs and sperm into the
water where external fertilisation takes place. The larva passes through a trochophore and a
veliger stage before developing into a juve nile. The trochophore is similar to the
Polyplacophoran trochophore. The veligers and juveniles are readily recognised as bivalves, as
they have a small flattened shell attached. When preserved the organs are usually retracted
within the shell.
Bivalve development stages. A, trochophore; B, oyster veliger stage, dorsal and lateral; C, later
larva. Figure A from Crofts (1937); B, from Hardy (1972); C, from Rees (1950).
43
CLASS CEPHALOPODA
The Class Cephalopoda comprises the nautilus, cuttle fish, squid and octopus. The cephalopods
have no distinctive larval stage, the juveniles emerge as miniature adults.
Order Sepiodea
This order includes the cuttle fish. The larvae have ten tentacles round the mouth, with one pair
longer than the others. They either have a sac- like body with long narrow lateral fins, or the
body is rounded posteriorly with circular lateral fins.
Examples of Sepioidea larvae (From Muus 1963a)
Order Teuthoidea
This order includes the squids, the larvae of which, as with the Sepioidea have 10 tentacles
round the mouth, one pair longer than the others. Their bodies are sac- like or torpedo-shaped
with terminal or sub-terminal fins
Examples of squid larvae (A,B from Muus 1963b; C from Muus 1963c)
44
Order Octopoda
This order includes the octopus which has eight tentacles, more or less all the same size, round
the mouth. In the Genus Octopus there are no fins on the body.
Examples of the Genus Octopus (From Muus 1963d).
45
CLASS GASTROPODA
The Gastropoda is the largest class of molluscs and includes the truly planktonic forms as well
as the familiar benthic snails and sea slugs. The typical gastropod consists of a head and a
visceral mass (the internal organs) sitting on top of a muscular foot. The visceral mass is usually
protected by a typically coiled shell. During development the visceral mass is rotated through
180 degrees anticlockwise. The Class Gastropoda comprises two Subclasses, Streptoneura (also
known as Prosobranchia) and Euthyneura (comprising the pulmonates and opisthobranchs). We
have followed the molluscan system proposed by Taylor & Sohl (1962) and adopted by Van der
Spoel et al. (1997).
Larval stages
Gastropod larvae are always present in coastal plankton, and can occur in large numbers. Some
gastropods have a trochophore larvae, and the majority have a veliger larvae. The trochophore
larva is very small and usually passes through the mesh used in sampling. The veligers are easy
to pick out as they look like what they are, miniature snails. All veliger larvae have a shell, even
those that have a reduced or lost shell as an adult. The veliger swims using a lobed, ciliated
structure called the velum. Although the velum is often diagnostic, it is usually withdrawn into
the shell in preserved specimens. The size of the larva varies from approximately 0.5 up to 3
mm or more. The veliger larvae are difficult to identify to species and we have not attempted to
do so in this guide.
A
B
D
C
Gastropod larval types: A, veliger with 2- lobed velum; B, veliger with 12- lobed velum; C,
veliger with 4-lobed velum; D, echinspira larva. Figures A from Werner (1955); B,C, from
Dawydoff (1940); D from Dakin & Colefax (1940).
46
Subclass Streptoneura
This group of gastropods is also known as Prosobranchia. Prosobranchs are the most primitive
of the gastropod subclasses. Prosobranch bodies are typically enclosed in a coiled shell (spirals
in a clockwise direction when viewed from above) and the animal has an operculum. The head
has a single pair of tentacles, each bearing an eye at its base. The sexes are separate. There is
usually a planktonic dispersal phase in the life cycle, trochophore, and veliger larvae, and some
species release free-floating eggs. This subclass has considerable functional and anatomical
diversity, and there is no single characteristic by which prosobranchs can be distinguished from
other gastropod groups. The pla nktonic heteropods and the family Janthinidae belong.
Superfamily Atlantacea Heteropoda

The planktonic heteropods show striking modifications to the basic prosobranch body plan and
life style. The shell may be larger than the body (as in Atlanta), may be reduced (as in
Carinaria) or absent (as in Firoloida). The foot is developed into a tool for propulsion. The
body is nearly transparent. Sexes are separate. Heteropods produce eggs in gelatinous egg
masses that may form strings. The embryos develop to the veliger stage before emerging into
the plankton, the velum of the veligers with three or four pairs of lobes. Heteropods are
voracious carnivores, they swim upside down and are restricted to shallow warm waters. The
Heteropoda is composed of the families Atlantidae, Pterotracheidae and Carinariidae.
Family Atlantidae
The family is composed of three genera, Oxygyrus, Protatlanta and Atlanta. All have a
generally flattened, coiled shell, with a keel extending around all or part of the outer edge of the
shell. The keel is characteristic in atlantids, but is not always obvious in younger specimens. In
young specimens the shell is sufficiently transparent for the eyes to be clearly visible. The eye
has a semi-transparent, amber-coloured lens with an area of dense black pigment behind. The
animal is able to withdraw into the shell completely. The species are separated according to the
shape of the coils in the shell, the start and end of the keel, the shape of the shell opening, the
length of the tentacles and the shape of the eyes. The foot is laterally flattened, shaped like a fin
with an operculum and a sucker. Atlantid species range in size from 0.5-11 mm, but the
majority are in the range of 2-3 mm.
Typical atlantid features. Figure from Souleyet (1852).
47
Genus Atlanta
Body large with a prominent proboscis and a swimming fin. Genus comprised of twenty one
valid species. The velum of atlantid veligers is 6- lobed.
Atlanta inflata Souleyet, 1852
Length: body up to 2 mm, shell diameter up to 1.5 mm (keel excluded).
A small atlantid with large eyes and a single swimming fin. The shell is flattened with a darkly
coloured keel. 7 whorls to shell. The spire is low and coloured violet or yellow/brown.
Recorded: Madagascar.
A
B
Atlanta inflata; A dorsal; B lateral. Figures from Tesch (1949).
Atlanta fusca Souleyet, 1852

Length: body up to 2 mm diameter, shell diameter up to 1.7 mm (keel excluded).
A small atlantid with large eyes and a single swimming fin. The shell is flattened with a white
coloured keel. 5 whorls to shell. The spire is high with a buff colour. The apical angle of the
spire is 90 degrees.
A
B
Atlanta fusca; A dorsal; B lateral. Figures from Tesch (1949).
48
Atlanta turriculata dOrbigny, 1836

Length: body up to 3 mm diameter, shell diameter up to 3 mm (keel excluded).
A small atlantid with large eyes and a single swimming fin. The shell is flattened with a keel
that slightly penetrates between the body whorl and spire. 5.5 whorls to shell. The spire is high
with a buff or rosy colour. The apical angle of the spire is 40 degrees.
A
B
Atlanta turriculata; A dorsal; B lateral. Figur es from Tesch (1949).
Atlanta helicinoides Souleyet, 1852

Length: body up to 2 mm diameter, shell diameter up to 2 mm (keel excluded).
A small atlantid with large eyes and a single swimming fin. The shell is flattened with a keel
that slightly penetrates between the body whorl and spire. 7 whorls to shell. The spire is large
with a yellowish-violet colour, the keel is a slightly darker colour.
A
B
Atlanta helicinoides; A dorsal; B lateral. Figures from Tesch (1949).
49
Genus Oxygyrus
Only one species.
Oxygyrus keraudreni (Lesueur, 1817)
Length: body up to 10 mm diameter, shell diameter up to 10 mm.
Atlantid with large eyes and a single swimming fin. The shell is flattened with a keel that ends
near the aperture. The spire and keel are non-calcareous keel. 1 whorl to shell is visible. The
shell is a brown colour.
Recorded: Madagascar
Oxygyrus keraudreni; dorsal. Figure from Tesch (1949).
Genus Protatlanta
Only one species.
Protatlanta souleyeti (Smith, 1888)
Length: body up to 1 mm diameter, shell diameter up to 1.5 mm (keel excluded).
A small atlantid with large eyes and a single swimming fin. The spire and body whorl is
calcareous but the keel is cartilaginous. The shell is flattened and the keel extends over half the
circumference of the shell and ends just before the aperture. 3.5 whorls to shell. The shell is
transparent.
A
B
Protatlanta souleyeti; A dorsal; B lateral. Figures from Tesch (1949).
50
Family Pterotracheidae
Medium to large planktonic animals, from 5-250 mm in length, but usually between 5-15 mm.
Adults have completely lost their shell. Body usually transparent. The eye is conspicuous, a
spherical, semi-transparent, amber-coloured lens with an area of black pigment behind. Two
genera in the family, Pterotrachea and Firoloida.
Genus Pterotrachea
Genus comprises four species (P. coronata, P. hippocampus, P. minuta and P. scutata), all
occur in the western Indian Ocean. The body is transparent and cylindrical with one swimming
fin. The large proboscis is perpendicular to the body.
Pterotrachea hippocampus Philippi 1836
Length: 20-80 mm.
Eyes are triangular with the base of the retinal part widened. The outer surface is scattered with
red dots in live specimens. Eyes and mouth parts are visible through the transparent body. The
female spawns by extruding a long filament of encapsulated fertilised eggs within a transparent
gelatinous matrix. The egg-string extends to over five times the length of the female.
Pterotrachea hippocampus: A, adult; B, veliger. Figures from Thiriot-Quivreux (1973).
51
Genus Firoloida
Only one species. Sexes are separate.
Firoloida desmaresti Lesueur 1817
Length: 15-40 mm.
The body is colourless and transparent. Elongate proboscic anteriorly and a visceral nucleus
posteriorly. Body long, cylindrical with one swimming fin. The female bears an internal
filament nearly as long as the body. This remains attached and receives embryos that pass from
the uterus. The embryos are gradually pushed by successive spawn to the open end where they
emerge as free-swimming veliger larvae.
Recorded: Seychelles, Rodrigues, Madagascar.
A
Firoloida desmaresti: A, adult; B, veliger. Figures A from Franc (1948), B from ThiriotQuivreux (1973).
52
Family Janthinidae
The janthinids are surface dwelling oceanic animals, they have a large float consisting of a
mucous-coated bubble of air. The animals float upside down. The shell is globular and rather
delicate, it is frequently washed ashore but seldom intact. The shells are spiral, the animal and
the shell are a vivid blue or violet colour, often the part under the water is white. Janthina
doesnt have eyes, but nonetheless is a carnivore. Prey consists largely of the similarly drifting
Velella or by-the-wind-sailor. The family is composed of two genera Janthina and Recluzia.
Genus Janthina
The genus is composed of five valid species. All species do not possess swimming fins and live
in the pleuston. The species are viviparous protrandric hermaphrodites. The shell is thin and
fragile.
Janthina exigua Lamarck, 1816
Length: shell up to 17 mm high, aperture height up to 11 mm..
Approximately 5 whorls in the shell. The shell is dark violet in colour. The spire has a welldefined tip. The edge of the shell opening is characterised with an incision in the middle.
Distribution: western Indian Ocean
Janthina exigua. Figures from Laursen (1953).
Janthina janthina Linnaeus, 1758

Length: shell up to 31.5 mm high, aperture up to 21.8 x 20 mm.
Approximately 4 whorls in the shell. The shell is blue to violet in colour. The spire tip fairly
flat. The edge of the shell opening has a small incision.
Distribution: Kenya, Mauritius, western Indian Ocean
Janthina janthina. Figures from Laursen (1953).
53
Janthina pallida Thompson, 1841

Length: shell up to 24 mm high, aperture up to 22 x 13.5 mm.
Approximately 4 whorls in the shell. The shell is a deep violet colour. The aperture is large in
comparison to the shell height. The edge of the shell opening has a small incision.
Janthina pallida. Figures from Laursen (1953).

Janthina prolongata de Blainville, 1822
Approximately 4 whorls in the shell. The shell is a deep violet colour. The spire tip is flattened.
The edge of the shell opening has a small incision.
Janthina prolongata. Figures from Laursen (1953).

Janthina umbilicata DOrbigny 1840
Approximately 4 whorls in the shell. The shell is thin, transparent and a deep violet colour. The
spire has a well-defined tip. The edge of the shell opening has a distinct incision.
Janthina umbilicata. Figures from Laursen (1953).
54
Subclass Euthyneura
The subclass Euthyneura comprises the opisthobranchs and the pulmonates. The opisthobranchs
comprise marine species with reduced shells and the pulmonates comprise the freshwater and
land gastropods. The opisthobranch larvae hatch as veligers, or less commonly, as juveniles
having passed through the trochophore stage within a gelatinous egg mass. All veliger larvae
have a shell, this is lost during metamorphosis for those species without a shell as an adult (e.g.
the nudibranchs). Of the planktonic species, the majority fall under the group of pteropods.
Pteropod adults may have a shell as in the Order Thecosomata, the sea butterflies or may lose
it during metamorphosis as in the Order Gymnosomata.
Order Nudibranchia
Nearly all nudibranchs are benthic, very few have adapted to a pela gic existence.
Glaucus atlanticus Forster, 1777
Length: up to 43 mm.
A surface living nudibranch that is a vivid blue-green colour dorsally and white ventrally. There
are three or four branched processes on either side of the body. The species is hermaphroditic.
Glaucus are carnivorous and are frequently associated with the siphonophore Velella.
Distribution: western Indian Ocean.
Glaucus atlanticus adult. Figure from Woodward (1875).
55
Order Thecosomata
Thecosomata are shelled pteropods.
Family Limacinidae
The family comprises one genus, Limacina. The shell is a left- handed spiral, transparent and the
spire can be high or flattened. The adults usually possess an operculum. The large winged- feet
protrude from the shell opening, they are used in locomotion and move the animal in a spiral
coarse. Most species produce egg masses, larvae hatch out as veligers. The species are
protrandric hermaphrodites. There are eight species of Limacina.
Limacina trochiformis (dOrbigny, 1836)
Length: shell height 1 mm, width to 0.8 mm.
5 whorls in the shell. The shell is white to light-purple in colour and transparent. The outer
surface of the shell is smooth.
Limacina trochiformis. Figure from Tesch (1946).

Limacina bulimoides (dOrbigny, 1836)
Length: shell height 2 mm, width to 1.4 mm.
6 whorls in the shell. The shell is transparent and colourless. The outer surface of the shell is
smooth.
Limacina bulimoides. Figure from Tesch (1946)..

56
Limacina inflata (DOrbigny, 1836)

Shell with 3 whorls coiled in one level, border of the heart-shaped aperture with a tooth
supported by a rib, umbelicus and surture deep and distinct. Shell diameter up to 1.5mm.
Common species in the Indian Ocean.
Distributed: Seychelles, Rodrigues, Zuza.
Limacina inflata lateral and dorsal views (From Van der Spoel, 1972)
57
Family Cavoliniidae
Shells are straight or slightly curved. Cavoliniidae is composed of eight genera Creseis,
Styliola, Hyalocylix, Clio, Cuvierina, Diacria, Diacavolinia and Cavolinia.
Subfamily Clioinae
Clioinae is composed of four genera, Creseis, Styliola, Hyalocylis and Clio. All are
characterised by straight or slightly curved shells. The species are protandric hermaphrodites.
Genus Creseis
Four species of Creseis, three of them have been recorded in the western Indian Ocean.
Creseis acicula (Rang, 1828)
Length: shell is up to 35 mm long and 1.5 mm across.
Shell is transparent, long and straight, and round in cross section. The shell surface is smooth. A
distinctive tentacular lobe on the anterior margin of the fins.
Creseis acicula: A, Adult, B detail of wings. Figure A from Woodward (1875), B from
Meisenheimer (1905).
Creseis virgula (Rang, 1828)
Length: shell length up to 7 mm, opening diameter 1 mm.
Shell is transparent, curved and round in cross section. The shell surface is smooth and
colourless. There are three varieties forma conica, forma constricta (recorded from Rodrigues)
and forma virgula based on characteristics of the shell shape.
Creseis virgula: Adult shell shapes. Figures from Tesch (1946).
58
Genus Styliola
Only one species.
Styliola subula (Quoy & Gaimard, 1827)
Length: shell length up to 13 mm.
Shell is needle- like in shape, transparent and round in cross-section. A characteristic straight
ridge runs obliquely from the opening down to the left. There is no tentacular lobe on the
anterior margin of the fins. The species can occur in swarms.
Styliola subula: Adult shell. Figure from Tesch (1946)

Genus Hyalocylix
Only one species.
Hyalocylix striata (Rang, 1828)
Length: shell length up to 8 mm.
The shell is cone-shaped, slightly curved dorsally and completely transparent. The opening is
oval in transverse section. The surface of the shell has transverse thickened bands that
progressively get wider apart near the aperture. The animal is often separated from its shell
when preserved, but is easily recognised by the very large fins. The species can occur in
swarms.
Hyalocylix striata: Adult shell. Figure from Tesch (1946).
59
Subfamily Cavoliniinae
Cavoliniinae is composed of three genera, Diacria, Cavolinia and Diacavolinia.
Genus Cavolinia
Five species of Cavolinia. A very distinctive shape of shell with a marked bulge on the ventral
plate. The species are protandric hermaphrodites. Several species occur in the western Indian
Ocean.
Cavolinia globulosa (Gray, 1850)
Length: shell up to 6 mm long and 4.5 mm wide.
The shell is globular, transparent with a brownish colour. The anterior section of the shell is
rounded. The anterior section of the ventral side has strong transverse ribs.
A
Cavolinia globulosa: A, dorsal view of shell; B, lateral view of shell. Figures A & B from
Tesch (1946).
Genus Diacavolinia
Twenty two species of Diacavolinia, Diacavolinia species are characterised by the absence of a
caudal spine.
Diacavolinia longirostris (de Blainville, 1821)
Length: shell length 5.2 7.2 mm, width 4.9 6.8 mm.
Dorsal side of the shell is relatively flat whereas the ventral side is deeply vaulted. The shell has
well developed lateral spines. The shell is brown in colour. Different forms have been noted.
Diacavolinia longirostris: A outlines of different shell forms ventral. In decreasing size forma
limbata, longirostris, strangulata and angulosa; B, same in lateral excluding forma strangulata;
C, forma limbata ventral view. Figures from Van der Spoel (1972).
60
Genus Diacria
The genus comprises two species groups, ten species. The species may be globular, with both
dorsal and ventral sides rounded, or bilaterally symmetrical with a long caudal spine. The
species are protandric hermaphrodites.
Diacria trispinosa (de Blainville, 1821)
Length: shell length 8 mm, width 10 mm.
The shell is bilaterally symmetrical. Very long caudal spine and strong lateral spines. There are
five ribs on the dorsal side and three ribs on the ventral side. The shell is slightly transparent
and coloured brown, a darker colour on the ribbed sections.
A
Diacria trispinosa: A, shell, dorsal; B, shell, ventral. Figures A & B from Tesch (1946).
Diacria quadridentata (de Blainville, 1821)

Length: shell length .2.0 3.0 mm, width 1.8 2.5 mm.
The shell is small, globular, with rounded dorsal and ventral sides. There are no caudal or
lateral spines. The dorsal side extends further than the ventral side, this section can be seen
when viewed ventrally.
B
A
Diacria quadridentata: A, ventral; B, dorsal; C lateral. Figures A, B & C from Tesch (1946).
61
Order Gymnosomata
Gymnosomata are pteropods without shells.
Family Desmopteridae
Only one genus, Desmopterus.
Genus Desmopterus
Three species in the genus, two are present in the western Indian Ocean. The species are
protandric hermaphrodites.
Desmopterus papilio Chun, 1889
Length: body length between 0.5 2.0 mm, swimming wings between 2.0- 4.0 mm..
There is no shell. The animal has wings that are disc-shaped and transparent. There are two
long tentacles off the lateral wings. The body is large and situated centrally between the lateral
wings.
Desmopterus papilio: Adult. Figure from Meisenheimer (1905).
62
PHYLUM ANNELIDA
The phylum Annelida comprise segmented invertebrates characterised by the possession of a
coelom (body cavity) and hair-like bristles known as setae or chaetae. The chaetea occur along
the body in various configurations and aid in locomotion, feeding and sometimes protection.
The phylum is subdivided into the Oligochaeta and Polychaeta (latin derived from greek
meaning few and many bristled respectively). The former have few marine representatives and
are not included in the guide. The Polychaeta can be further subdivided into the Errantia, or
free living mobile worms, and the Sedentaria, the tube dwelling or burrowing forms.
A
Polychaete life forms. Errantia: Bottom dwelling: A, polynoid; B, nereid. Pelagic: C,

tomopterid. Sedentaria: D, spionid; E, terebellid. Figures A-C from Fauvel (1923), D and E
from Fauvel (1927).
Polychaetes may be represented in the plankton by larval stages of bottom living species, adults
and larvae of pelagic species, and adults of some species that ascend into the plankton to breed
at specific times of the year. Not all polychaetes have planktonic larvae, some release them
onto the substratum.
Life cycle of typical bottom dwelling

errant polychaete worm. Figure
adapted from Thorson (1946).
63
Larval Stages
Similar to the molluscs, annelids start their larval life as a trochophore. Development begins
with a pronounced extension of the head section of the trochophore, with segmentation of the
body beginning behind (i.e. the anterior segments are the oldest). The larva is known as a
metatrochophore at this stage. Each segment extends laterally and gives rise to flap- like
parapodia, special pits on the parapodia produce the hair-like setae. The setae may be very
long in young individuals, but tend to be replaced by shorter setae before settling on the
seabed. The nektophore larva has further head development and more segmentation of the
body. As segmentation continues, the larva is referred to as a nektochaeta. There are no great
changes in body form between the nektochaeta and the juvenile worm when it finally sinks to
the bottom and begins its benthic life.
Identification of polychaete larvae is difficult, we have not attempted to identify them to family
or genera. Polychaete larvae are usually present in the plankton of tropical waters all year
round in low numbers, however if sampling coincides with the breeding period, the larvae may
make up as much as 10% of the sample.
A
Post trochophore polychaete larvae: A, Myriochele sp.; B, Spionid sp.; C, Chaetopterus sp.; D,
Hediste sp.; E, Spiophanes sp.; F, Lanice sp.; G, Lagis sp.; H, Lepidonotus sp. Figures from
Thorson (1946).
64
Body parts of polychaete worms.

A
jaws
uniramous parapodia
biramous parapodia
proboscis
dorsal cirrus
prostomium
notopodia
palps
eyes
neuropodia
acicula
cirri
chaetae
ventral cirrus
Errant polychaetes: A, anterior detail of Nereis sp.; B, detail of parapodia, biramous (nereid)
and uniramous (phyllodocid). Figures from Fauvel (1923).
Adult pelagic polychaetes
There are several species of pelagic polychaetes. They show a variety of adaptations to their
pelagic life, these adaptations aid in identifying the polychaetes to family level. It is often
difficult to identify the polychaetes further to genera or species level. Pelagic polychaetes come
under four main families, Tomopteridae, Typhloscolecidae, Phyllodocidae and Alciopidae.
Family Tomopteridae
The Tomopteridae are wholly pelagic and are one of the more easily identified of the pelagic
polychaete families, however further identification to species requires some skill. They are
transparent, theytusually posess a pair of eyes, there are no chaetae and the biramous parapodia
extend laterally into paddle like structures. The prostomium is drawn out into horn like
antennae The parapodial aciculae (supporting chitinous rods) of the second segment are drawn
out into long streamers, which sweep back up to two-thirds or more of the body length. Key
features for identification to species level include 1) the presence/absence of the small cirrus of
the second segment, between the prostomial horns and acicular streamers, 2) the occurrence
and position of various glands in the parapodia and 3) presence/absence of reduced posterior
segments forming a tail.
A
B
Tomopteris kefersteini: A, adult, whole body; B, middle parapodia showing rosette glands.
Figures A and B from Fauvel (1923).
65
Family Typhloscolecidae
The Typhloscolecidae are small pelagic polychaetes. They are slender, cylindrical, transparent
worms, with the prostomium extending forward as a papilla and merging posteriorly into the
anterior segments. Eyes are always absent, but the prostomium has well developed nuchal
organs. The three segments behind the prostomium bear a single pair of large foliacious cirri.
Remaining segments carry two cirri, the dorsal and ventral cirri of the parapodia. Setae may be
present in the posterior segments. Paired anal cirri also show a similar paddle shape. Common
genera include Typhloscolex and Sagitella.
Sagitella kowalevskii Wagner, 1872
Length: 5-15 mm.
Only one species in the genus. Up to 30 body segments. Nuchal organs are not extended as
freely projecting lobes, they are usually hidden by the pair of large foliacious cirri. The
parapodial cirri are easily lost in preserved specimens.
Recorded: Seychelles, Rodrigues.
Sagitella kowalevskii: A, adult; B, anterior detail with some cirri removed to show nuchal
ridges; C, tail showing anal cirri. Figure A from Fauvel (1923), B and C from Dales (1957).
66
Travisiopsis lobifera Levinsen

Length: 25-39 mm.
Body more plump than Sagitella. Nuchal organs prominent and extend as backwardly
projecting lobes, they encircle a raised caruncle. The parapodial cirri are large and foliacious.
Anal cirri large and foliacious, each with a transparent supporting rib.
Recorded: Indian Ocean.
A
B
Travisiopsis lobifera: A, adult; B, anterior detail; C, posterior detail. Figures A from Fauvel
(1923), B and C from Dales (1957).
67
Family Alciopidae
The Alciopidae are all pelagic. The worms are often small and usually transparent, with
varying amounts of brown pigmentation at the base of the parapodia segmental glands. The
main feature is the presence of a pair of large bulging eyes, which are red in life, but turn
brown in preserved specimens. Unlike phyllodocids the prostomium usually has five small
antennae. The papillate proboscis is eversible, and is sometimes found with two anterior cirri
and occasionally teeth. Parapodia are uniramous with dorsal and ventral cirri. Alciopids are
slender fragile worms, which fragment easily on collection.
Torrea candida (Delle Chiaje, 1828)
Length: 150-250 mm.
Characterised by dark segmental glands in each segment with a line of dark pigment dorsally.
Prostomium not projecting in front of eyes. Prostomium with two pairs of frontal antennae and
a single median antenna between the eyes. Proboscis long, typically with two long horns and
papillae on the rim. Parapodia with foliacious dorsal and ventral cirri.
Recorded: Indian Ocean
Torrea candida: A, adult; B, detail of parapodia. Figures from Dales (1957).
68
Rhynchonerella angelini Kinberg, 1866

Length: 60-20 mm.
Body with 150 segments. Coloured yellowish or brown in preserved specimens. Body
segments and prostomium darkly pigmented dorsally. Four short fat antennae arise from the
prostomium, a rounded lobe projects in front of the eyes. Parapodia with large foliacious dorsal
and ventral cirri.
Recorded: Indian Ocean
A
B
C
Rhynchonerella angelini: A, adult; B, detail of head; C, detail of parapodia. Figures from

Fauvel (1923).
Alciopa reynaudii Audouin and Milne Edwards, 1829
Length: 10-15 mm.
A long, slender, delicate pelagic polychaete. A pair of large lensed eyes. Body segments
terminate in two cirriform appendages. The prostomium with five short antennae, doesnt
extend in front of the eyes.
A
Alciopa reynaudii: A, adult; B, detail of parapodia. Figures from Fauvel (1923).
69
Vanadis formosa Claparde, 1870

Length: 200- 300 mm.
Body long and many segmented. Proboscis long, typically with two long horns. Three pairs of
tentacluar cirri. Prostomium with two pairs of frontal antennae and a single median antenna
between the eyes. Prostomium not projecting in front of eyes. Parapodia with foliacious dorsal
and ventral cirri. Parapodia with pigmented glands except in 3rd and 4th parapodia. Specimens
are frequently broken when sampled.
A
Vanadis formosa: A, adult; B, detail of parapodia. Figures from Dales (1957).

Vanadis longissima (Levinsen, 1885)
Length: 120 200 mm.
Eyes very large, lens not visible dorsally. Proboscis without horns. Three pairs tentacluar cirri.
Prostomium with two pairs of frontal antennae and single long median antenna between eyes.
Pigmented segmented glands in intervals from the fifth parapodium creating brown bands.
A
Vanadis longissima: A, adult; B, detail of parapodia. Figures from Fauvel (1923).
70
Family Phyllodocidae
The Phyllodocidae are not exclusively pelagic. The pelagic phyllodocids are small, usually
between 1 and 30 mm in length, with few segments and a short wide body. The head bears
four small antennae and a single pair of eyes, which may be reduced in some species. The
body is often flattened. The first three segments carry finger like extensions known as cirri, the
following segments bear a pair of paddle shaped parapodia with dorsal and ventral foliacious
(leaf like) cirri. The cylindrical proboscis may be everted beyond the prostomium in preserved
individuals. The proboscis may be smooth or bear papillae.
Lopadorhynchus uncinatus Fauvel 1915
Length: 5-25 mm, typically 12 mm.
Prostomium broad with two pairs of antennae, one long and one short. Three pairs of tentacular
cirri. First two segments are swollen with simple chaetae, the following parapodia with paddleshaped chaetae. The first two segments have dorsal cirri only, following segments have dorsal
and ventral cirri.
Lopadorhynchus uncinatus: A, adult; B, posterior parapodia showing paddle-shaped chaetae.

Figures A and B from Dales (1957).
71
Pelagobia longicirrata Greeff 1879

Length: up to 12 mm.
Body with 15-25 segments. Prostomium with two pairs of short antennae and a single pair of
eyes. Two pairs of tentacular cirri. Dorsal cirri absent from first two segments, following
segments with long dorsal and ventral cirri. Chaetae with serrated tips.
Pelagobia longicirrata: Adult. Figure from Dales (1957).
72
PHYLUM ARTHROPODA
SUBPHYLUM UNIRAMIA
CLASS INSECTA
There is only one true marine insect, Halobates. It belongs to the Family Gerridae, the pond
skaters or water striders. It has long legs and lives on the sea surface and do not dive. While it
cannot be considered planktonic, it is often caught in plankton nets as they sample through the
surface layer.
The genus Halobates contains over 40 species. Like other gerrids, the sea skaters have a life
cycle consisting of the egg, 5 nymphal instars, and the adults. Females are usually larger than
the males.
They are predators, typically feeding by liquefying the muscles and the internal organs of their
prey. The diet of the open ocean species is not fully known. They have been observed feeding
on zooplankton, dead jellyfish, fish eggs and larval fish trapped on the ocean surface and have
never been observed breaking the water surface to feed.
Although they spend their entire lives on the ocean, Halobates requires floating objects on
which to deposit their eggs. The objects include sea bird feathers, floating seashells, pieces of
wood, plastic and lumps of tar. The eggs are oval in shape and quite large (~1 mm x 0.2 mm).
Other insects are often found in plankton samples, but these will have been blown into the sea
by the wind.
Recorded: Seychelles
Halobates sp. (from Cheng, 1975)
73
SUBPHYLUM CHELICERATA
CLASS PYCNOGONODA
Unconventional looking arthropods, sometimes called sea-spiders, and certainly resemble
spiders in appearance. The body is reduced and they usually have 4 pairs of legs (rarely 5 or 6).
They are found crawling over the bottom, feeding on hydroids and other fleshy animals. They
are nor really planktonic, but can turn up in plankton samples in shallow water. Most are small
but the largest have a leg spread of 40 cm.
Example of pycnogonid (From Wickstead, 1965).
SUBPHYLUM CRUSTACEA
CLASS BRACHIOPODA
Sub-order Cladocera
Cladocera are small crustaceans with a large compound eye with the limbs originating just
behind. Some species have a transparent triangular brood pouch, which can contain eggs or
many young larvae, small replicas of the adult e.g. Evadne and Penilia.
B
A
A: Penilia avirostris; B: Evadne tergestina (From Wickstead, 1965).
74
CLASS MAXILLIPODA
Infra class Facetotecta
A tiny unusual larva, considered to be related to the barnacles. Previously recorded from
European and Japanese waters. The carapace has a typical reticulated surface.
Recorded: Rodrigues, Seychelles.
Nauplius y larvae, dorsal and lateral views (Reproduced from

Further records of nauplius y type IV Hansen from Scandinavian
waters by Schram, T.A., from Sarsia, www.tandf.no/sarsia 1972,
Vol 50, p7, by permission Taylor & Francis).
Infra-class Cirripedia
Larvae of the cirripedes or barnacles can be common in the inshore plankton. Their nauplius
larva resembles a copepod nauplius and they also go through a series of moults. They typically
have two horns on the front of the carapace. A cypris stage develops inside the last nauplius,
which then attaches itself to a rock or some other suitable site and develops into the adult. This
cypris stage has a two-section shell and resembles an ostracod. It usually has a dark eyespot on
the side.
A: Cirripede nauplius; B cirripede cypris (From Fraser, 1962).
75
Sub-class Copepoda
Copepods are one of the most numerous, diverse and biologically important zooplankton groups in
the sea, so more background information on them is provided here than for the other groups. There
are ten orders of copepods. The orders with most species are Calanoida, Cyclopoida,
Poecilostomatoida and Harpacticoida and to a much lesser extent Siphonostomatoida and
Monstrilloida. It is likely that Orders Cyclopoida and Poecilostomatoida will be merged in the
near future. The other small orders have not been included in this guide as none have been
sampled. They are either very rare, cave dwelling or live very deep or close to the sea bottom.
Development of the egg and nauplius stages
Typical free- living copepod development is from a small, ge nerally circular or sub-circular egg,
which depending on species is around 30-450m in diameter. The egg surface is smooth in some
species and spiny in others (Fig. 1), so size and surface ornamentation can be useful in
identification. Eggs may be held in an egg sac, matrix of tissue, egg mass without an enclosing
membrane, or be spawned directly into the sea, depending on the copepod order.
Antennule
Antenna
Caudal armature
Fig. 1. Examples of free-spawned calanoid

copepod eggs (From Koga, 1984).
Fig. 2. Calanoid copepod nauplii

(From Koga, 1984).
When the egg hatches, a nauplius stage emerges, the typical crustacean larva (Fig 2). As with
other crustaceans, the nauplius has an external skeleton, which has to be moulted between each
subsequent developmental stage. The nauplii of most free-swimming copepods typically moult six
times (Stages NI-NVI), the sixth time developing into the first copepodite stage (CoI), the stage
from which the adult features start to appear. Each of the nauplius stages can be identified by
changes between moults in body segmentation and particularly by changes in the number of setae
on the terminal segment of the antennule and number and arrangement of spines (caudal armature)
on the posterior end of the body.
Identifying nauplii
It is not difficult to roughly distinguish to at least which of the major groups a nauplius belongs.
Calanoid nauplii are usually reasonably robust, tend to have a humped back and quickly develop
an elongated body, which protrudes well beyond the cephalic shield (Fig. 3). The caudal armature
is usually arranged in pairs of spines but often not of the same length. Cyclopoid nauplii are
shield-shaped and flattened (Fig. 4), the posterior body hardly protruding beyond the cephalic
shield, and as they tend to be small copepods, their nauplii are mainly small. Their limbs also tend
to be much slenderer than calanoid nauplii. Poecilostomatoid na uplii (Fig. 5) are similar to
cyclopoid nauplii (As mentioned above these two orders are in the process of being merged).
76
Harpacticoid nauplii tend to have bizarre shapes and unlike most of the other three main orders,
are often not bilaterally symmetrical (number of spines each side a mirror image of the other).
Some have single spines sticking backwards (Fig. 6), rather than a paired arrangement as found in
most, but not all of the other orders. For some common copepod species, in well-researched
regions of the world, descriptions of each nauplius stage are available, thus, where there is low
species diversity, identification to species can be possible.
Fig. 3. Example of calanoid (Eucalanus)

copepod nauplii stages (From Koga, 1984).
Fig. 5. Example of poecilostomatoid (Oncaea)

Fig. 4. Example of cyclopoid (Oithona)

nauplii stages (From Koga, 1984).
Fig. 6. Example of harpacticoid (Longipedia)

Copepodite body segmentation and limbs

A generalised copepod description is given here; specific differences between copepod orders are
given under the description of their order. Morphological terminology for the copepod body has
varied between authors, but here follows Huys and Boxshall (1991) and Mauchline (1998).
There are typically six copepodite stages, the sixth being either the adult male or female (Fig. 7).
Number of body segments and swimming legs usually changes between moults, so stage of
development (Stages CoI-CoV1) can be distinguished. Segmentation and leg number differences
between species/genera is common. The body is divided into cephalosome, metasome and
urosome (Fig. 8); the cephalosome and metasome together, are termed the prosome. Theoretically,
the adult copepod body consists of 16 segments. In all copepods the cephalosome (sometimes
77
called the head) consists of 6 segments fused together, outwardly appearing as one segment (5
cephalic segments + the first thoracic segment), all bearing limbs. The first pair of limbs, the
antennae 1 (or antennules) are sensory (Fig. 9). In the males of many orders, either one or both
may be modified, sometimes drastically, related to grasping the female during copulation. The
modified A1 is called the prehensile or geniculate A1. In some orders the A1 are small.
CoI
Ce+5
CoII
Ce+6
CoIII
CoIV
CoV
Ce+7
Ce+8
Ce+9
CoVI ?
Ce+9
CoVI ?
Ce+10
Fig. 7. Sequence of development of body segmentation between copepodite stages 1-6 of Calanus
finmarchicus. Arrows indicate the last metasome segment. The shaded segment indicates the segment
added at the previous moult. Number of segments in addition to the cephalosome is noted as Ce+5 etc.
(Based on a figure from Mauchline, 1998 and incorporating figures from Lebour, 1916 and Sars, 1903).
Fig. 8. Diagrammatic illustration of the external morphology and appendages of a calanoid copepod
(From Mauchline, 1998, copyright 1998, with permission from Elsevier).
78
Fig. 9. Diagramatic representations of the appendages of a calanoid copepod. The swimming legs
usually have developed endopods and exopods with up to three segments, here numbered 1-3.
art: arthrite; b: Basis; c: Coxa; e: Endite; en: Endopodite; epi: Epipodite; ex: Exopodite; gnb:
Gnathobase; PC: Praecoxa (From Mauchline, 1998, copyright 1998, with permission from
Elsevier).
The next five limb pairs make up the feeding appendages: antennae 2 (or antennae), mandibles,
maxillae 1 (or maxillules), maxillae 2 (or maxillae) and maxillipedes. One or other of the latter
two limb pairs can be greatly enlarged in some species, associated with carnivorous feeding e.g.
the maxillae 2 in Candacia spp. and the maxillipedes in Euchaeta spp.
Some of the next 5 segments (second to sixth thoracic segments; Fig. 8) may be partially or
completely fused to the cephalosome, or together, so there are interspecific differences in the
number of segments that can outwardly be seen. They generally all bear paired swimming limbs
(P1-P5) and because of this are referred to as pedigerous segments. The fifth legs are often
reduced in size and complexity compared to the other legs, which is typical of many female
copepods, completely missing, which is again typical of many females, or modified. Heavily
modified P5 legs are typical of many male copepods, the leg being adapted to deposit the
spermatophore. The swimming legs are joined at their bases so they beat together. The basal
segment is called the coxa (Fig. 9) and the next segment the basis. From the basis, the limbs
typically become biramous (with two branches), although one branch may be reduced or missing.
The inner branch is the endopodite and the outer the exopodite. The next segment (thoracic
segment seven) bears the genital openings in both sexes and is termed the genital segment. The
sixth legs have become completely reduced to form the structure which closes off the genital
opening in the two sexes. In female copepods the genital segment is usually fused with the
following or more segments, giving the appearance of at least one less segment in the urosome
than found in the males. The segments posterior to the genital segment do not bear any limbs. The
last segment of the urosome has the anus located on the ventral side and is termed the anal
segment. This segment ends in two furca, sometimes called caudal rami, and bears several setae.
Development
While in theory there are 16 segments in the copepod body, because of segment fusion, outwardly
this cannot be seen. The variation in number of prosome and urosome segments between the
adults of species, is also reflected in the 6 (CI-CVI) developmental stages, and this can be useful
in separating species. Additionally, the number of swimming legs changes between the early
79
moults. An example of the sequential development of segmentation in a calanoid copepod

(Calanus) is given in Figure 7. Mauchline (1998) gives a table (page 27) with examples of how
segmentation of the urosome and number of pairs of swimming legs changes between
developmental stages, depending on how many segments or swimming legs are finally developed
in the adult. The table does not include details for calanoid copepods that develop 4 urosome
segments in the adult female, one of the commonest arrangements in calanoid copepods. Between
moults a segment is added, always immediately anterior to the anal segment. In the example in
Figure 7, addition of a segment in CoI and CoII results in a segment being incorporated into the
metasome between each moult. In CoIII the final metasome number is reaches, so after this,
further segments added anterior to the anal segment leads to an increase in the number of
segments in the urosome, till the final number is reached. In this example it is five segments in the
male and would have been five in the female, but the first segment of the urosome fused with the
second segment during the last moult, forming the genital segment. Urosome segment number
thus remains at four between the female CoV and CoVI.
In species which develop an asymmetric P5 in the male, the ultimate sex of the copepodite can be
seen from CIV, because of morphological differences between the female and male P5s (Fig. 10).
In species in which the female does not develop a P5 the male will have a rudimentary P5 in CIV
which will be missing in the female.
IV ?
IV ?
V?
V?
IV ?
V?
IV ?
V?
Fig. 10. Fourth and fifth swimming legs of female and male developmental stages CIV and CV
of A, Temora longicornis and B, Acartia clausi (From Kraefft, 1910).
Morphological differences to distinguish between the main copepod Orders
In copepods the major movable articulation (bending joint) of the body is located between the
prosome and urosome. In calanoids, the articulation is actually between the eleventh and twelfth
segments of the body (Fig. 11), thus all the limbs are on the broader part of the body. In cyclopoid,
poecilostomatoid and harpacticoid copepods, the major articulation of the body occurs between
80
the tenth and eleventh body segments, thus the first segment of the urosome bears a pair of limbs
(usually much reduced). In most poecilostomatoida and cyclopoids, as in calanoids, the urosome is
much narrower than the prosome. In harpacticoids, there is much less of a difference in segment
width between the prosome and urosome e.g. Microsetella.
Fig. 11. A, Calanoid; B: cyclopoid (with twin egg sacs); C: poecilostomatoid; D: harpacticoid
(with single egg sac). All female copepods (A from Sars, 1925; B, C from Sars, 1918; D from
Giesbrecht, 1892). Arrows indicate the genital segments. In the cyclopoid and poecilostomatoid
copepods, rudimentary limbs can be seen immediately after the articulation and in the
harpacticoid copepod there is less clear separation of the cephalosome and urosome.
Points to note
Most identification texts only give descriptions of the adult CVI copepods, so before trying to
identify a copepod to species, it is important to check that it is an adult. In most female copepods
the presence of a swollen genital segment indicates that it is mature. However, a problem with
some females e.g. Acartia is that the genital segment starts to swell in CV, while the body
segmentation is often identical between CV and CVI. However, in both female and male pre-adult
copepods, the segmentation of the P5 and urosome is not as distinct as in the adult stages, which
becomes obvious when you examine them together.
In many figures, for clarity, some of the limbs, or part of the limbs such as setae, are not drawn.
Many copepods have very distinctive features, even in the pre-adult stages; so can often be
identified at least to genera.
There are often far fewer male copepods than females sampled. This is because in many species a
single fertilization is adequate for several batches of eggs, so fewer males are required. In species
where each batch of eggs requires fertilization, the numbers of each sex may be similar.
While most identification texts only give total body lengths (anterior cephalosome to tip of furca),
prosome length is a useful comparative body measurement, as the urosome is often bent at an
angle or damaged, which can make measurement of total length imprecise.
81
Order Calanoida
One of the most numerous and diverse copepod orders. The adults tend to be much larger than
those of other orders. The A1 is long and the A2 is biramous. Males often have a geniculate A1,
sometimes remarkably modified, for grasping the female during copulation e.g. Anomalocera. The
prosome is much broader than the urosome and the body articulates between them i.e. between the
last metasome segment (body segment 11) and the genital segment (body segment 12). In females,
the posterior of the last metasome segment, tends to be bilaterally symmetrical or in most cases
only slightly asymmetric, while in males it is often asymmetric e.g. Anomalocera.
Females usually have 5 pairs of swimming legs, occasionally 4 pairs as in Families Aetideidae
and Euchaetidae, or in individual ge nera such as Acrocalanus. The female P5, when present, are
usually, symmetrical, although often much reduced in size and complexity compared to the other
swimming legs e.g. Paracalanus. Males always have 5 pairs of swimming legs, the P5 usually
greatly modified and asymmetrical, used during copulation e.g. Candacia. Female have between 2
and 4 segments in the urosome (not counting the furca) while in males there are generally 5,
occasionally 4 as in some Labidocera spp. There are usually 5 setae on each fur ca, but sometimes
only 4 or 3. Females either spawn their eggs directly into the sea (e.g. Temora, Acartia), or are
typically held in a single egg sac (e.g. Euchaeta), occasionally paired egg sacs (e.g. some
Pseudodiaptomus spp.), or egg mass, although some deep-water species carry a pair of very large
eggs. Egg sacs are usually knocked off during sampling, but some species e.g. Euchaeta are often
found bearing eggs.
82
Family Augaptiliidae
Cephalosome and first pedigerous segment separate. Urosome in female 3-4 segments, male 4-5
segments, symmetrical in both sexes. Mandible with few teeth. P5 of female similar to the other
legs, biramous. A1 of male geniculate on the left, P5 with a 3-segmented exopodite, endopodite of
1-3 segments. P5 without claws and not very asymmetric.
Genus Euaugaptilus
Euaugaptilus hecticus (Giesbrecht, 1889)
Length: females: 2.45-2.75 mm; males: 2.28-2.40 mm.
Cephalosome appendages of both sexes generally long and setose.
Female: Urosome long and slender, narrowing slightly at the distal end, genital segment twice as
long as the two following segments together. Furca equal in length to the anal segment, around
three times as long as wide. In perfect specimens, the inner furcal seta is as long as the whole
body. A1 much longer than the body. The endopodites and exopodites of P5 both 2-segmented.
Mandible very simple, with only 2 teeth.
Male: Similar in appearance to the female. Urosome long and slender. Genital segment as wide as
long. A1 geniculate on the right. The second segment of the right P5 exopodite, with an internal
process, the last segment terminating in a long seta.
A
B
C
E
D
Euaugaptilus hecticus: Female. A, dorsal; B, P5; C, mandible. Male. D, P5; E, dorsal. Figures A-B
from Bradford-Grieve (1999), C-E from Giesbrecht (1892).
83
Genus Haloptilis
Haloptilis longicornis (Claus, 1863)
Cephalic appendages of both sexes large and plumose.
Female: A knob-like projection on the anterior cephalosome when viewed dorsally. Long A1,
which extends beyond the furca by around the last 9 segments. P5 almost symmetrical, with the
last inner seta on the terminal segment of the exopodite shorter than the other inner setae.
Mandible very simple.
Male: Considerably smaller than the female. A1 reaches the end of furca. The 2 setae on the
terminal segment of the right P5 equal in length. Long spine on the end of the left P5.
Recorded: Madagascar, Aldabra.
B
C
Haloptilis longicornis: Female. A, Dorsal; B, P5; C, mandible. Male. D, dorsal; E, P5. Figures
from Giesbrecht (1892).
84
Haloptilis mucronatus (Claus, 1863)

Female: The cephalosome is tapered anteriorly with a point on the forehead, which curves
downwards. The A1 reaches around the end of the furca. Very short urosome.
Male: Anterior cephalosome rounded. Spination on the P5 terminal exopodite segments weak.
B
D
Haloptilus mucronatus: Female. A, dorsal; B, head lateral; C, mandible. Male. D, dorsal; E, P5.
Figures A-C from G.O. Sars (1925), D-E from Giesbrecht (1892).
85
Haloptilis oxycephalus (Giesbrecht, 1889)

Length: females: 3.0-4.1mm; males: 2.4-2.85 mm.
Female: Body quite slender with a very long projection on forehead, curving slightly downwards.
Urosome short and furca diverge slightly. The A1 reaches slightly beyond the furca. Mandible
simple.
Male: Cephalosome broadly triangular with apex rounded. P5 with glandular pores at the base of
every spine on the exopodite, quite large projection on the basal segment.
B
E
D
Halopsis oxycephalus: Female. A, dorsal; B, head lateral; C, mandible; D, P5. Male. E, dorsal. F,
P5. A-C from G.O. Sars (1925) D-F from Bradford-Grieve (1999).
86
Haloptilis spiniceps (Giesbrecht, 1892)

Female: Body quite robust, anterior cephalosome with a short curved point similar to H.
mucronatus. The A1 reaches slightly beyond the furca.
Male: Cephalosome rounded without point. P5 slightly asymmetric, the third segment of the right
exopodite terminating in 2 quite strong short spines and with a median transverse spine.
Haloptilus spiniceps: Female. A, dorsal; B, head lateral; C, P5. Male. D, doral; E, P5. Figures A-C
from Bradford-Grieve (1999), D from Mori (1937), E from Giesbrecht (1892).
87
Family Heterohabidae
Genus Heterohabdus
Heterohabdus papilliger Claus, 1863
Female: Anterior cephalosome rounded in dorsal view with a small papilla on forehead, but no
point. A1 hardly reaches the end of furca. Genital segment in lateral view with a pronounced
angular protrusion. The inner marginal spine of segment 2 of the exopodite of P5 is around the
same length as the third segment.
Male: P5 is asymmetrical. The proximal portion on the inner margin of segment 2 of the exopodite
of the right P5 has a process with a distal tooth; segment 3 is wider than the corresponding
segment on the left leg; basis has a rounded projection.
B
H
F
C
Heterohabdus papilliger: Female. A, dorsal; B, head lateral; C, P5; D, genital segment dorsal; E
genital segment lateral. Male. E, dorsal. F, P5; G, P5. Figures A-C, F-H from G.O.Sars ( 1925)
and Giesbrecht (1892), D, E, from Bradford-Grieve (1999).
88
Family Lucicutiidae
Males and female have very long urosomes, typically over half the lengt h of their bodies. They
also have long furca. In perfect specimens there is a seta on each furca, much longer than the other
setae.
Genus Lucicutia
Lucicutia flavicornis (Claus, 1863)
Bradford-Grieve (1999; p 98) points out that there are several distinct size classes attributed to this
species, indicating that there may be several different species.
Female: Cephalosome without lateral protrusions. Large genital boss, placed centrally on the
ventral segment. Furca slightly more than 5 times as long as wide; innermost terminal seta small
and slender. A1 reaches middle of furca. P5 endopodite with 3 segments; inner spine on exopodite
segment 2 is long and straight, except for a slight bend at tip, reaching beyond base of first inner
seta on exopodite segment 3; terminal segment on exopodite segment 3 less than half the length of
the segment. Outer margin of exopodite segment 3 with several teeth.
Male: Cephalosome without lateral protrusions. Furca slightly more than 5 times as long as wide;
innermost terminal seta small and slender. A1 reaches middle of furca. P5 right coxa inner margin
with a conspicuous rounded protrusion, right basis with a triangular inner border bearing hairs
distally; left coxa with a ridge on the inner margin, left basis inner distal corner protruding and
ending in a point and with 3-5 extra teeth and sometimes a proximal spinule. Inner margins of
both basis segments without pointed projections.
Lucicutia flavicornis: Female. A, dorsal; B, urosome lateral; C, P5. Male. D, dorsal; E, P5. From
Bradford-Grieve (1999).
89
Lucicutia ovalis (Giesbrecht, 1889)

Female: Cephalosome without lateral protrusions, furca 4 times longer than wide. In lateral view
genital segment with a ventral protrusion on posterior half of segment. In dorsal view urosome
symmetrical. A1 extends to posterior border of anal segment. P5 endopodite with 2 segments.
Male: Cephalosome without lateral protrusions. Furca 1.4 times longer than wide. A1 extends to
posterior border of anal segment. P5 right coxa smooth, left one with small spines; proximal part
of inner margin of both the basis segments with pointed projections; right endopodite 3segmented.
B
C
Lucicutia ovalis: Female. A, dorsal; B, urosome lateral; C, P5. Male. D, urosome dorsal; E, P5. A
Figures A from Bradford-Grieve (1999), B-E from Tanaka (1963).
90
Family Metridinidae
Males and females have a long urosome, over half the length of the body. In lateral view, the
body is generally widest just behind the head and behind this the back may curve in slightly. The
female P5 is small and simple.
Genus Pleuromamma
Typically have a brown or red button on the side of the body around the junction of the first body
segment. Depending on the species this can be on the right or left side.
Pleuromamma abdominalis (Lubbock, 1856)
Female: Anterior cephalosome without pointed process. The brown button can be on either side in
the female. The first two segments of the A1 of the female bears two strong spines of varying size
and appearance, one on first segment stronger and usually more curved than that on second
segment. Genital swelling centrally placed on segment, genital pore region has a dark protruding
cap. P5 with 3 free segments, terminating in three unequal spines, the innermost the longest.
Male: The brown button is on the left side. Only small spines at base of A1. The male urosome is
very asymmetric with tufts of hairs. Right A1 geniculate. P5 left with a wide terminal segment.
Recorded: Rodrigues, Seychelles, Zuza, Madagascar, CPR.
B
A
D
Pleuromamma abdominalis: Female. A, dorsal; B, genital segment lateral; C, P5. Male. D, dorsal;
E, P5. From Bradford-Grieve (1999).
91
Pleuromamma gracilis (Claus, 1863)

Female: Brown button on right side of body. First segment of A1 with only small, low teeth. P5
with only one free segment, terminating in 3 short spines with 2 thin spines on outer side.
Male: Pigment spot on right side. Urosome almost symmetrical, except that segment 3 has a patch
of hairs on the left rear border. Left A1 geniculate.
Recorded: Zuza, Madagascar, CPR.
A
D
B
C
Pleuromamma gracilis: Female. A, dorsal; B, urosome lateral; C, P5. Male. D, dorsal; E, P5. From
92
Pleuromamma indica Wolfenden, 1905

Length: females: 2.06-2.72 mm; males 1.62-2.38mm.
Female: Quite like P. robusta but has a small projection on the genital pore. Small spines on first 2
segments of the A1. The 3 terminal spines on the P5 reduce in size by 50% each time from the
longest inner spine.
Male: Button on right hand side. Symmetrical abdomen. Divergent furca.
Recorded: Zuza.
F
C
G
I
Pleuromamma indica: Female. A, urosome dorsal; B, urosome lateral; C, P5; D, base A1. Male.
E, urosome; F, dorsal and lateral; G, Geniculate A1, segments 17-21; H, geniculate A1; I P5
ventral; J, P5 dorsal. Figure F from Grice (1962), other figures from Steuer (1933).
93
Pleuromamma piseki Farran, 1929

Female: Slightly larger than P. gracilis. In dorsal view, the genital segment has a marked groove
or constriction on the left side, parallel to the posterior margin of the segment and a large blackpigmented area around the genital pore. P5 is imperfectly segmented, terminating in 3 short
spines, shorter than the width of the segment.
Male: Geniculate A1 with a double-toothed row on segment 18, as well as on proximal part of
segments 19-21, segment 17 is naked. Left P5 segment 3 with process with a knob, separated from
the process by a slight notch.
Recorded: Zuza, CPR.
E
F
B
Pleuromamma piseki: Female. A, dorsal; B, urosome lateral; C, P5. Male. D, dorsal; E, geniculate
A1; F, P5. From Bradford-Grieve (1999).
94
Pleuromamma robusta (F. Dahl, 1893)

Length: females: 3.0-4.7mm; males 3.0-4.0 mm.
Female: Genital swelling on anterior part of genital segment. Teeth on basal segments of A1 small
and low. P5 has 4 segments and 3 terminal setae, the innermost considerably longer than the
others. Pigment button on right side.
Male: Urosome almost symmetrical. Pigment button on right side. Left A1 geniculate. Right
exopodite segment 2 with a long thin process.
A
C
Pleuromamma robusta: Female. A, dorsal; B, urosome lateral; C, P5. Male. D, dorsal; E, P5. From
95
Pleuromamma xiphias (Giesbrecht, 1889)

Female: resemb les P. abdominalis but differs in outline of anterior cephalosome, which extends
forwards into a pointed process, bent slightly downwards. Large curved spine on basal segment of
A1 and small spine on second segment
Male: Cephalosome pointed as in female, but pointed forwards rather than downwards. Urosome
asymmetric and A1 geniculate
Recorded: Zuza.
B
D
Pleuromamma xiphias: Female. A, lateral; B, dorsal; C, P5. Male. D, lateral; E, dorsal; F, P5.
Figures A, B, D, F from Bradford-Grieve (1999), figures C, E from Sars (1925).
96
Family Acartiidae
In this family, only the Genus Acartia has been sampled in the region.
Genus Acartia
Typically both sexes have slender bodies, often coffin-shaped. When freshly preserved they are
usually quite transparent, sometimes tinged blue, often with a single prominent eyespot. The A1
typically bears many quite long fine setae. The furca each bear a fan of fine setae, which have a
characteristic spiky appearance. These setae are often damaged or broken off in other preserved
copepods, but are usually intact in Acartia, even in young stages. No rostrum, but some species
have paired fine rostral filaments. In the female the urosome is 3-segmented. The tiny P5 is
usually uniramous, 3-segmented, with the last segment modified into a long slender spine. In the
male the urosome is 5-segmented. The A1 is usually geniculate on the right side only, with
expanded segments in the middle. The tiny P5 is uniramous and asymmetric.
97
Acartia amboinensis Carl, 1907

Length: females: 1.34-1.51 mm; males: 1.30 mm.
Female: Corners of posterior metasome segment produced into a strong spine on either side
laterally and with 2 small spines dorsolaterally. The first segment of the A1 has 2 strong spines
and a minute spine on the posterior margin. The second segment has a moderate sized spine on the
distal posterior margin and 4 minute spines on the posterior margin. The third and fourth segments
each have a small spine on the posterior dorsal margin and the fifth segment has 4 small spines.
Ueda (1986) gives an A1 spine arrangement as 2 stout spines on the first segment, one small spine
on the second segment and one small spine on each of segments 3 and 4. He appears to disregard
some of the smaller spines described by Tanaka (1965). The genital segment has 2 spines on the
dorsal distal margin. The following segment has 4 tiny spines on the distal margin. In the P5, the
terminal spiny segment has 2 slight swellings near the base and bends abruptly inwards in the
distal half. The external seta is over 1.5 times as long as the terminal spine.
Male: Posterior metasome segment with 2 strong lateral spines. A pair of small spines on each side
of the dorsal posterior metasome margin. Strong spines on the posterior margin of urosome
segment 2 with a smaller lateral spine each side; third segment with 2-4 small marginal spines and
the fourth segments with 4-6 small marginal spines. P5 basis segments with long strong external
setae.
Recorded: Madagascar by Binet and Dessier (1968), but their illustrations differs slightly from
those of Carl (1907) and Tanaka (1965), especially in the structure of the female P5, so this record
may be invalid.
B
A
Acartia amboinensis: Female. A, dorsal; B, urosome lateral; C, proximal A1; D, P5. Male. E, P5;
F, urosome dorsal; G, thorax and urosome lateral. Figures A-D from Tanaka (1965), figures E-G
from Carl (1907).
98
Acartia bispinosa Carl, 1907

Length: females 1.32 mm; males: 1.30 mm.
Female: Rostral filaments present. Corners of posterior metasome segment with strong spines; 2
smaller spines dorsal and medially each with a tiny spinules just behind them. Posterior edge of
the genital segment with a pair of strong spines dorsolaterally. First segment of the A1 with 2
strong spines and a smaller spine near the base of these spines. The second segment with a
characteristic, obvious, strong curved spine. The terminal slender spines of the P5 curved strongly
from around the middle.
Male: Each corner of the posterior metasome segment with a pairs of spines, one slightly shorter
than the other. A pair of spine one each side of the same segment dorsolaterally, one much smaller
than the other and with a fine setule just behind them. Carl (1907), in the original description, only
draws a single dorsal curved spine on either side. The second urosome segment has a strong
dorsolateral spine on each side of the posterior margin and smaller ones on each side
ventrolaterally. Terminal segment of P5 left with 2 fine terminal spines, a stout spine on the
midanterior surface with fine setae at its base, tufts of hairs proximal to the medial spine, row of
spinules along inner margin, and several spinules along outer margin.
E
F
A
C
D
Acartia bispinosa: Female. A, rostrum and base of A1; B, P5; C, thorax and urosome dorsal. Male.
D, thorax and urosome dorsal; E P5 right; F P5 left. From Nishida (1985).
99
Acartia danae Giesbrecht, 1889

Female: Rostral filaments present. Spine present on the basal segment of the A1. A strong spine
on each posterior corner of the metasome. The urosome does not bear strong spines, but the first
two segments have small hairs on the posterior dorsal edge. The P5 terminal spiny segment is
coarsely serrated on both sides, twice as long as the previous segment, while the outer marginal
seta is three times the length of the spine.
Male: Posterior metasome as in female. Urosome segment 1 with lateral hairs, and a pair of small
spines on the rear edge of segment 2.
Recorded: Seychelles, Zuza, Madagascar.
F
C
Acartia danae: Female. A, dorsal; B, head lateral; C, urosome lateral; D, P5. Male. E, dorsal; F,
urosome dorsal; G, P5. From Bradford-Grieve (1999).
100
Acartia erythraea Giesbrecht, 1889

Length: females: 1.4 mm; males: 1.2 mm.
Female: Rostral filament present; 2 strong spines on segment 1 of the A1, a smaller spine on the
second segment, 3 small spines on the third and 2 larger spines on the fourth segment. A very
strong spine on each corner of the posterior metasome segment and a pair dorsally, towards the
posterior edge of the same segment. A pair of spines on the dorsal rear edge of the genital segment
and a tiny pair on the following segment. The P5 is simple, with a curved terminal spiny segment
(Giesbrecht (1889) drew the spine gently curved, while Mori (1937) drew a much stronger curve);
the outer plumose seta is one and a half times the length of the spiny segment.
Male: No strong spines at the base of the A1. The same spine arrangement on the last metasome
segment as the female. The second urosome segment is wider than long, with a pair of prominent
spines either side on the posterior margin, of the same size. Two tiny dorsal medial spines on the
rear edges of urosome segments 3 and 4. Ueda (1986) draws 4 spines on segment 3.
Recorded: Seychelles region
E
F
Acartia erythraea: Female. A, dorsal; B, A1; C, P5; D, base of A1. Male. E, thorax and urosome
dorsal; F, P5. Figures A, E, F from Giesbrecht (1892), B, C from Mori (1937), D from Ueda
(1986).
101
Acartia fossae Gurney, 1927

Female: 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 segment 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.
Bradford-Grieve (1999b) illustrates the P5 with the outer seta only slightly longer than the
terminal spine.
Male: Posterior metasome segment with row of tiny spinules on dorsoposterior margin as in the
female. First urosome segment with lateral tufts of hair. Second urosome segment with
dorsolateral rows of tiny spinules arranged across and on the posterior edge of the segment. Third
and fourth segments also have tiny spinules on posterior dorsal edge. The left P5 has a terminal
segment with a thick naked terminal spine, one modified spine and one seta on the midanterior
surface. The inner margin has a row of fine hairs.
Acartia fossae: Female. A, urosome dorsal; B, P5. Male. C, urosome dorsal; D, P5. From Nishida
(1985).
102
Acartia negligens Dana, 1849

Female: Rostral filaments present. Small species, with one or sometimes more small spines on the
posterior corners of the last metasome segment. A row of tiny hairs may also be present dorsally,
although difficult to see except under very high power. A1 reaches to furca, its first segment with
a small slender spine. The first two urosome segments have a dorsal row of fine spinules on the
posterior border. The P5 is very simple, similar to A. danae, terminating in a very short, straight,
spiny segment with serrated outer edges and a very long outer seta at least 5 times longer than the
spine.
Male: A1 shorter than body. The male is also reported to have a tiny spine on either side of the last
body segment, although some authors suggest tufts of hairs. Urosome segments 1 and 2 hairy
laterally, segments 2, 3 and 4 with tiny dorso-posterior spinules. P5 similar to A. danae.
B
A
C
D
G
Acartia negligens: Female. A, dorsal; B, anterior urosome dorsal; C, urosome lateral; D, P5. Male.
E, dorsal; F, urosome dorsal; G, P5. From Bradford-Grieve (1999).
103
Family Candaciidae
Typically medium sized robust copepods, with a very square front to the cephalosome and
pronounced shoulder shape anteriorly. In almost all species the last segment of the metasome, in
both males and females, ends in widely spaced spines. Both the male and female P5 are small and
simple. Many species, even in the young stages, have tinges of black pigment on the ends of the
swimming legs or on other parts of the body. The genital segment in both sexes often bears
prominent spines. Maxilla 2 large, associated with their carnivorous diet. Female P5 uniramous,
usually symmetrical. Male posterior segment of the urosome always asymmetrical and genital
segment almost always asymmetrical. A1 geniculate on right; 23 segments, segments 17-18 and
19-20 fused, with or without teeth at bend. P5 4-segmented on left and 3-segmented on right; may
be chelate (have a claw- like process) on right or ending in a feather- like seta.
Genus Candacia
Female P5 terminal segments may end in one or more spine like processes, a finger-like process or
a single long setae; setae may or may not be present on the inner lateral margins. Right A1 of male
with teeth present on one or more segments at the bend in the geniculate region. Male right P5 is
chelate.
104
Candacia bradyi A. Scott

Length: females: 1.4-2.1 mm; males 1.4-1.8 mm.
Female: Posterior metasome segment corners each end in a short spine. Genital segment broad and
almost symmetrical in dorsal view, with a slight anterior protrusion on right side. Urosome
segment 2 has a pointed protrusion on the mid ventral surface. P5 segment 3 curved slightly
inwards, with 2 seta on inner margin and three spines on the distal outer edge with an additional
spine further back.
Male: Genital segment produced into a small toothed process on right side. Left P5 segment 3 is
produced at outer distal angle into a short stout pigmented tooth- like process, which is divided into
3 blunt points, segment 4 is elongated and narrow with 3 small terminal spines.
D
E
H
G
Candacia bradyi: Female. A, urosome ventral; B, urosome lateral; C, P5; D, urosome dorsal.
Male. E, urosome dorsal; F, urosome segments 1 and 2; G, P5; H, maxilla 2. Figures A-C, E-H
from Greenwood (1978), figure D from Lawson (1973).
105
Candacia catula (Giesbrecht, 1889)

Female: Posterior metasome symmetrical and pointed. Genital segment symmetrical with both
sides swollen. No spines on genital segme nt, but with a backward projecting protuberance on the
ventral surface, situated towards the rear edge. Segment 3 of P5 is long and has 2 outer marginal
spines and 3 inner marginal setae and 3 teeth at the end.
Male: Last segment of the metasome pointed, but not prominently. Points are slightly asymmetric.
Unusual among Candacia males in having no protuberances on the genital segment.
Recorded: Rodrigues, Seychelles, Zuza, Madagascar, Aldabra.
D
E
Candacia catula: Female. A, dorsal and lateral; B, urosome lateral; C, urosome dorsal; D, P5.
Male. E, dorsal and lateral; F, maxilla 2; G, P5: H, urosome. Figures B, C, F, G from Greenwood
(1978), A, D, E, H from Grice (1962).
106
Candacia curta (Dana, 1849)

Length: female s: 1.82-2.70 mm; males: 1.90-2.65 mm.
Female: The body is broad and the spines on the last metasome segment are set wide apart. Has an
asymmetric genital segment with a sharp downward projecting spine on the ventral surface,
originating from the right side of the segment. There are 2 points on the end of the P5 with a
further point set further back
Male: One of the spines on the last metasome segment is very crooked, and there is a very obvious
curved sharp protuberance on the genital segment.
Recorded: Zuza, Aldabra, Madagascar, CPR.
A
B
E
F
Candacia curta: Female. A, dorsal; B, lateral; C, urosome ventral; D, P5. Male. E, dorsal; F, P5;
G, corner of last metasome segment and urosome segment 1. From Grice (1962).
107
Candacia discaudata A. Scott, 1909

Female: The spines on the last metasome segment of the female are not as obvious as in most of
the other Candacia species and project forwards slightly in lateral view. The genital segment is
quite symmetrical, with a slight protuberance on the front. The second urosome segment has a
ventral rounded projection in lateral view. The anal segment is short and asymmetrical. The P5 are
almost symmetrical.
Male: The genital segment is asymmetrical and bulges on the right side. Viewed from the right the
inflated region bears a small tooth at each end. The anal segment is asymmetrical.
Recorded: Rodrigues, Aldabra, Seychelles.
D
A
E
C
B
Candacia discaudata: Female. A, dorsal; B, urosome lateral; C, P5; D maxilla 2. Male. E, dorsal;
F, urosome dorsal; G, P5. Figures A-D, F, G from Scott (1909), E from Mori (1937).
108
Candacia ethiopica (Dana, 1849)

Female: The prosome is often very darkly pigmented. Easily distinguished from the other species
of the genus by the small lateral and ventral spiny processes of the genital segment and by the
terminal segment of the P5, which has 7 spines. In the female the last cephalosome segment rises
to a small crest on the back at the rear edge. The genital segment is slightly asymmetric, with
protrusions on each side, one larger than the other.
Male: Recognised by the curved, spiny projection of the right posterolateral corner of the
metasome. The genital segment has 2 triangular processes on the right margin. There is also a
small crest on the last cephalosome segment, which is also found in some other Candacia spp. The
genital segment has protuberances on one side, consisting of a rounded knob in front of which is a
pointed projection.
G
D
Candacia ethiopica: Female. A, dorsal; B, urosome ventral; C, urosome lateral; D, P5. Male. E,
dorsal. F, urosome ventral;G, P5. From Bradford-Grieve (1999).
109
Candacia longimana (Claus, 1863)

Length: females 2.86-3.90 mm; males: 2.40-3.55 mm.
Female: Posterior metasome segment with small points. Genital segment swollen and
symmetrical. Segment 3 of P5 terminated by 3 short teeth of similar length and a further tooth set
half way back along the segment.
Male: Can be identified by the spiny process on the right posterolateral margin of the asymmetric
metasome, the knob- like process on the right side of the genital segment, and the structure of the
P5. There is a fine-toothed comb on the right geniculate A1.
F
D
C
G
Candacia longimana: Female. A, urosome dorsal; B, urosome lateral; C, P5. Male. D, urosome
dorsal. E, urosome lateral;F, geniculate A; G, P5. From Bradford-Grieve (1999).
110
Candacia pachydactyla (Dana, 1849)

Both the female and male have small ridges on the back at the rear of the last cepha losome
segment.
Female: Posterior metasome segment pointed and asymmetrical. A markedly asymmetric genital
segment with characteristic large pointed protuberances extending obliquely Segment 3 of P5
thickened with 3 strong terminal spines, three seta on the inner side and a strong spine on the outer
edge.
Male: Posterior metasome asymmetrical. Genital segment with a large process on the right,
consisting of a single, broad, rounded projection. Has a large characteristic protuberance on the
genital segment.
B
A
G
H
Candacia pachydactyla: Female. A, dorsal; B, urosome ventral; C, urosome lateral; D, P5. Male.
E, dorsal and lateral; F, urosome ventral;G, urosome lateral; H, P5. Figures B-D and F-H from
Goesbrecht (1892), A, E from Grice, 1962).
111
Genus Paracandacia
Female P5 terminal segments end in a fingerlike process which may be finely serrated on one or
both margins. There are 2 setae on the inner lateral margin of these segments. Geniculate right A1
of male with no teeth at the bend. Male right P5 never chelate, ending in a feather- like seta.
Paracandacia bispinosa (Claus, 1863)
Length: females: 1.74-2.01 mm; males: 1.88-2.11mm.
Female: The posterior metasome is produced into spines either side. The genital segment is nearly
triangular when viewed from above. Each side of the segment is produced into a small spine; the
spine of the left side is longer and directed backward. These spines, particularly the one on the left
are variable in shape and size. There is a small rounded knob arising from the ventral side of the
genital segment. Segment 3 of the P5 is wide and terminates in a broad spine, finely serrated on
both sides. There are 2 sharper spines slightly further back on the outside, with a further spine half
way back and 2 setae on the inside.
Male: No diagrams of the male body seem to be available, but presumably the posterior metasome
is produced into spines as in the female. Can be distinguished by the swollen segment 20 of the
geniculate portion of the right A1. There are 3 outer edge spines on the distal segment of the right
P5
Recorded: Rodrigues, Zuza, Madagascar, Aldabra.
Paracandacia bispinosa : Female. A, dorsal; B, urosome lateral; C, P5. Male. D, geniculate A1

segments 16-21; E, P5. From Grice, (1962).
112
Paracandacia simplex Giesbrecht

Length: females 1.67-1.93 mm; males 1.73-1.80 mm.
Female: Posterior corners of the metasome pointed and directed backwards. Genital segment
almost symmetrical and without lateral spine- like protrusions. Dorsal side of genital segment with
pointed protrusion. P5 segment 3 with a broad fingerlike terminal processes, finely serrated
proximally. There are two inner edge spines, the distal one twice the length of the proximal one.
Male: Posterior corners of the metasome pointed but not prominently so. Geniculate A1 segments
19-20 fused, but without any protrusions.
H
C
B
D
I
E
Paracandacia simplex: Female. A, dorsal; B, urosome dorsal; C, urosome ventral; D, urosome
lateral left; E, urosome lateral right; F, P5. Male. G, dorsal and lateral; H, geniculate A1 segments
17-21; I, P5. Figures B-F Bradford-Grieve (1999), A, G, H, from Grice (1962), I from Giesbrecht
(1892).
113
Paracandacia truncata (Dana, 1849)

Length; females: 1.84-2.10 mm; males: 1.87-2.11 mm.
Female: A very characteristic square end to the last metasome segment when viewed laterally. The
corners of the metasome are pointed and directed forwards, so the points are not visible in dorsal
view. The urosome is quite symmetrical, with no protuberances. P5 has 3 terminal fingerlike
processes, finely serrated distally and 2 terminal setae of slightly different lengths.
Male: Last metasome segment symmetrical, with sharp points. The urosome is symmetrical with
no projections. The P5 is quite similar to the P5 of P. bispinosa but can be distinguished as it has a
small spine on the external distal side of the left leg segment 2. There is another spine on the
external distal corner of segment 3, and 4 long spines of almost equal length on the end of segment
4. Segment 3 of this limb is also much narrower than in P. bispinosa. The geniculate A1 has a
series of stout proximal segments followed by a thin section, then a very broad club section.
Beyond the club section, segment 16 has a fingerlike protrusion which is difficult to observe
clearly. Fused segments 17 and 18 are characteristically curved.
Recorded: Rodrigues, Zuza, Aldabra, Madagascar, CPR
Paracandacia truncata: Female. A, dorsal; B, P5. Male. C, dorsal and lateral; D, geniculate A1
segments 15-20; E, P5. Figures A, B, E from Dakin and Colefax (1940), C, D from Grice (1962).
114
Family Centropagidae
A wide range of Centropages spp are recorded from the Indian Ocean. They are small to medium
sized copepods and tend to have quite square cephalosomes. Some have the lateral corners of the
posterior metasome segment ending in a spine, with a characteristic undulating edge on the section
between the spine and the urosome. The males have a complex P5 with one side forming a large
claw. One side of the male A1 is typically very thickened along part of its length.
Genus Centropages
Centropages calaninus (Dana, 1849)
Female: The last metasome segment is rounded. Both the female genital segment and furca are
slightly asymmetric. The genital segment swells laterally in dorsal view. The anal segment is
almost twice as long as urosome segment 2. A1 extends beyond furca by its last 2 segments. P5
exopodite segment 2 inner edge spine is straight and longer than exopodite segment 3. There is a
notch on the proximal inner margin of exopodite segment 1.
Male: The last metasome segment is rounded. Right P5 exopodite segment 3 claw is longer than
the inner extension of exopodite segment 2 and is sharply bent.
Recorded: Rodrigues, Seychelles, Madagascar, Aldabra.
E
A
Centropages calaninus: Female. A, dorsal; B, urosome dorsal; C, exopod of P5. Male. D, dorsal;
E, P5. Figures B-D from Giesbrecht (1892), A, E from Mori (1937).
115
Centropages elongatus Giesbrecht, 1896

Female: Last metasome segment is quite rounded; genital segment almost symmetrical. A1 passes
the furca by the last 2 segments. The spine- like extensions of the exopodite segment 2 of the P5
exceed the distal end of exopodite segment 3. Notch in proximal internal part of segment 1 of P5.
The furca are as long as the 2 preceding segments together.
Male: Last metasome segment quite rounded. Long external spine terminating the right exopodite;
distinctive shape to end of left exopodite.
B
E
Centropages elongatus: Female. A, dorsal; B, urosome lateral; C, P5. Male. D, dorsal; E,

urosome dorsal; F, P5; G, terminal segments of exopodite of P5 left; H, terminal exopodite of left
P5 other side; I, geniculate A1 segments 17-21. From Grice (1962).
116
Centropages furcatus (Dana, 1849)

Female: Quite transparent species. Cephalosome has a conspicuous ventral ball- like eye, obvious
even in young stages. Prominent second spines between the lateral spines on the last metasome
segment and the urosome. Strong spines on segments 1, 2 and 5 of the A1. P5 exopodite segment
2 inner edge spine does not reach the distal border of exopodite segment 3. No spines on genital
segment. Anal segment not quite symmetrical, long, around twice the length of urosome segment
2. The furca are long and slender
Male: Posterior metasome border asymmetrical. Prominent second spines between the lateral
spines on the last metasome segment and the urosome. P5 right extension on exopodite segment 2
has a rounded protrusion on the proximal portion; exopodite segment 3 claw is stout and has a
spine on inner margin and 2 spines on outer margin; left exopodite segments 2 and 3 with long
terminal fixed appendages. The furca are very long and slender.
D
B
A
G
C
J
I
E
H
F
Centropages furcatus: Female. A, dorsal; B, head lateral; C, urosome ventral; D, anterior A1: E,
exopodite segments 2 and 3 of P5. Male. F, dorsal ; G, urosome dorsal: H, P5; I, right P5
exopodite segments 2 and 3; J, left exopodite segments 2 and 3 of P5. Figures A-E, G, I, J from
Giesbrecht (1892), F, H from Mori (1937).
117
Centropages gracilis (Dana, 1849)

Female: The lateral angles of the last metasome segment are rounded. The second urosome
segment has lateral knobs covered in short spines. The furca is symmetrical. The A1 extends
beyond the furca by about the last 5 segments. The inner marginal spine on exopodite 2 of the P5
does not reach the end of segment 3.
Male: The terminal claw of the grasping right P5 is longer than the thumb and has a pronounced
triangular protrusion, absent in other species of the genus.
Centropages gracilis Female. A, dorsal; B, urosome ventral; C, P5. Male. D, dorsal ; E, P5.
Figures A, B. D from Mori (1937), C, E from Grice (1962).
118
Centropages orsinii Giesbrecht, 1889

Female: The last metasome segment corners are pointed, but not sharply. The furca are around
twice as long as wide. The genital segment is symmetrical and has an offset spine on the ventral
surface, which points backwards. The A1 does not have spines on segments 1, 2 and 5 and reaches
the end of the furca. P5 are nearly symmetrical and the inner marginal spine on exopodite 2 is
curved, is usually covered in small spines and does not reach the end of segment 3.
Male: The last metasome segment corners are weakly pointed. The terminal claw of the grasping
right P5 is longer than the thumb. The terminal portion of exopodite segment 3 of the left P5
projects into a spine.
Recorded: Rodrigues, Seychelles, Madagascar.
B
E
D
A
C
G
H
F
Centropages orsinii: Female. A, dorsal; B, urosome ventral; C, P5; D, exopodite segments 2 and 3
of P5. Male. E, dorsal ; F, P5; G, left exopodite segments of P5; H, right exopodite of P5. Figures
B, D, E, G, H from Giesbrecht (1892), A, C, F from Mori (1937).
119
Centropages violaceus (Claus, 1863)

Female: Posterior metasome segments rounded. Genital segment moderately swollen, without seta
or spines but with a patch of spinules on right side and a ventral downward facing projection.
Urosome segment 2 with a centrally placed ventral protrusion. A1 without spines on proximal
segments; extends beyond the furca. P5 exopodite segment 2 does not reach distal border of
exopodite segment 3.
Male: Right geniculate A1 segments 15 and 16 without spines. Right P5 exopodite segment 2
bulbous, with an inner process with a narrow base, which is situated at mid-segment; exopodit e
segment 3 long and s-shaped, 3 times the length of the inner process on exopodite segment 2. The
second segment of the exopodite of the left P5 has 2 outer marginal appendages.
E
B
D
A
F
A
Centropages violaceus: Female. A, dorsal; B, urosome ventral; C, urosome lateral; D, P5. Male. E,
dorsal ; F, P5. Figures B-D from Giesbrecht (1892), A, E, F from Mori (1937).
120
Family Pontellidae
Many have prominent hooks on the sides of the cephalosome and paired lenses on the dorsal
cephalosome. P5 of the female is small and simple, but robust.
Genus Calanopia
The P5 of the female has a 1or 2-jointed exopodite. The endopodite is completely missing. The
male P5 has a 2-jointed exopodite. The exopodite of the right foot is in the form of a claw.
Calanopia elliptica (Dana, 1846, 1849)
Female: 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 segment. Furca nearly 3 times as long
as broad. P5 asymmetrical, each limb uniramous of 4 segments. Left leg longest, exopodite of 2
segments.
Male: Urosome segment 2 right side distal border produced into a well defined tooth. P5 terminal
segment of the le ft exopodite 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 exopodite
segment 1 with 3 blunt teeth, while the claw- like third segment has 3 small pointed teeth.
E
A
Calanopia elliptica: Female. A, dorsal; B, urosome lateral; C, P5. Male. D, dorsal ; E, urosome;
F, P5. Figures A-C, E, F from Scott (1909), D from Mori, (1937).
121
Calanopia minor A. Scott, 1902

Female: Superficially resembles a small C. elliptica. The last metasome segment produced on each
side into an acute spine; the rostrum pointed. Urosome 2-segment, the second segment distinctly
longer than the genital segment. 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: The second urosome segment has no spiny process. A1 extends to the end of the third
thoracic segment; the right A1 is geniculate. The basis of the left P5 is swollen towards the
proximal end of the inner margin, the swollen part produced into a small tooth- like spine. The
basis of the right exopodite is also swollen. Flattened joint of the first exopodite segment, with a
seta on the inner margin; outer thumb is short and naked. The claw- like second segment is spoonshaped and without any teeth, but with 2 inne r marginal seta.
Recorded: Seychelles, Madagascar.
B
A
F
D
C
Calanopia minor: Female. A, dorsal; B, P5. Male. C, metasome and urosome dorsal; D, urosome;
E, P5. Figures A, B, D, E from Scott, (1909), C, F from Tanaka (1964).
122
Calanopia parathompsoni Gaudy, 1969

Female: Lateral hooks on cephalosome, spines on last metasome segment reaching the middle of
the genital segment. Urosome 2-segmented as in other members of this genus. Genital segment
clearly asymmetric in dorsal view, twice as long as anal segment and is not produced on the
ventral surface into 2 lobes as in C. thompsoni. The furca are symmetrical. The A1 reaches half
way down the genital segment. P5 are very similar to those of C. thompsoni.
Male: Urosome symmetrical with the exception of 2 fine spines situated ventrally on the second
segment, on the left side. These laterally directed spines have not been reported in the males of
other species. The geniculate A1 and the P5 are similar to those of C. thompsoni. However, C.
parathompsoni has on the median part of the second segment of the right P5, a small spine,
swollen at the base. The second segment of the left P5 is also straighter. The terminal segment has
a curved spine and a rounded knob with small projections.
H
D
I
J
Calanopia parathompsoni: Female. A, dorsal; B, urosome lateral; C, urosome ventral; D, P5; E,
distal segment of P5. Male. F, urosome ventral; G, geniculate A1; H, P5; I, Right P5; J distal
portion of left P5. From Gaudy (1969).
123
Calanopia thompsoni A. Scott, 1909

Length: females: 2.62 mm; males: 2.19-2.31mm.
Female: Lateral hooks on cephalosome. Strong spines on last metasome segment, reaching half
way down ge nital segment. Genital segment slightly longer than the second urosome segment.
Genital segment ventrally is produced anteriorly and posteriorly into 2 blunt lobes. Furca slightly
asymmetrical, longer on left. P5 symmetrical, endopodite of 2 segments, first one moderately long
with distal portion of outer margin with 2 very strong spines. Second segment narrow, terminating
in a long stout spine, serrated on both edges, segment also with 2 outer marginal spines and a short
inner one.
Male: Lateral hooks on cephalosome, forehead angular in outline. Posterior metasome segment
sharply pointed. No spine on urosome segment 2. Right A1 geniculate. Furca symmetrical. P5
asymmetric, exopodite of the right leg claw-like.
C
A
Calanopia thompsoni: Female. A, dorsal; B, urosome lateral; C, P5. Male. D, dorsal; E, geniculate
A1; F, P5. Figures A-C, E, F from Scott (1909), D from Mori (1937).
124
Genus Labidocera
Cephalosome with or without hooks and with 1 pair of cuticular lenses and a ventral eye which
extends anterioventrally between the deeply bifurcate rostrum. Posterior of metasome with large
pointed processes. Female urosome 2 or 3 segmented, male urosome 4 or 5 segmented.
Labidocera acuta (Dana, 1849)
Female: Sharp spine on front of cephalosome, directed slightly downwards. No lateral cephalic
hooks; posterior metasome symmetrical with large lateral points. Urosome 3-segmented, genital
segment asymmetrical, with a projection on the posterior right side which extends half way along
the next segment, visible from the dorsal side. P5 with a rather variable exopodite, with relatively
large spines on the outer border; endopodite claw- like.
Male: Posterior metasome border asymmetric, right side longest and bent outwards. Urosome
segment 1 with a small spine on the right. Left P5 with 3 terminal and 1 outer spine; right P5
without a thumb on the claw, but with a curved triangular flap.
A
B
E
C
D
Labidocera acuta: Female. A, dorsal; B, urosome dorsal; C, head lateral; D, P5. Male. E, dorsal;
F, P5. Figures A from Giesbrecht (1892); B-D, F from Greenwood (1979); E from Mori (1937).
125
Labidocera acutifrons (Dana, 1849)

Female: Anterior cephalosome with dorsal lenses and median spine, without lateral hooks;
posterior metasome segment produced into points on each side, the corners of which are slightly
divergent. Urosome 3-segmented, genital segment slightly asymmetrical with a small lump on the
right side; second segment with a dorsoposterior spine on the right side, pointing towards a large
anal flap. P5 asymmetrical, left leg slightly larger, exopodite with 3 spines, endopodite reduced to
a spiny process.
Male: Anterior cephalosome with a median spine as in female, dorsal lenses well developed and
place close together; urosome 5-segmented; posterior metasome symmetrical. Left P5 with a
rudimentary endopodite, exopodite segment 2 with 3 terminal spines; right P5 exopodite claw with
an extra small segment bearing 2 setae and with a spine.
A
E
F
B
Labidocera acutifrons: Female. A, dorsal; B, urosome dorsal; C, P5. Male. D, dorsal; E, P5 left; F,
P5 right. Figures A from Giesbrecht (1892) B-C, E-F from Bradford-Grieve (1999), D from
Wilson (1932).
126
Labidocera bataviae A. Scott, 1909

Female: No lateral cepha lic hooks, rounded cephalosome, dorsal eye lenses spaced apart. Posterior
metasome segment almost symmetrical and produced into 2 divergent points; urosome short with
2 segments, genital segment asymmetrical and large, produced on the right side; well deve loped
lamina on the anal segment; furca very asymmetrical, left ramus almost 2 times wider than long
and arranged at right angles to the urosome. P5 asymmetrical, left leg longer, exopodite with 2
outer marginal spines and a minute spine on inner margin; terminating in 3 unequal spines, middle
spine longer and serrated along both edges. Endopodite much reduced with bifurcated tip.
Male: Body resembles female, but posterior metasome quite asymmetrical, right point longer than
left; urosome 5-segmented, the first segment with a notch on left side. Furca slightly asymmetric,
right ramus rounder than left. Right A1 geniculate. Right P5 claw like, thumb at proximal end of
exopodite segment 1, long and robust, with a distinct spine and seta close to base of thumb;
terminal segment curved inwards, with a conical protrusion on its inner mid- margin. Left P5
terminal segment with 1 outer mid- marginal spine and 3 terminal curved spines of which the
outermost is longest.
Recorded: Rodrigues
F
C
Labidocera bataviae: Female. A, dorsal; B, urosome dorsal; C, P5. Male. D, urosome dorsal; E,
P5; F, terminal segment of left P5 enlarged. Figures A, D, E from Scott (1909), B, C, F from Silas
and Pillai (1973).
127
Labidocera detruncata (Dana, 1849)

Length: females: 2.42-2.72 mm; males: 2.20-2.70mm.
Female: Forehead anterior prosome rounded, dorsal eyes spaced apart, no lateral cephalic hooks;
last metasome segment asymmetric, extending further back on left, with small lateral points;
internal to the points the segment curves outwards to the level of the points. Urosome 3-segmented
with left dorsal surface uneven, lined and notched, furca broad and almost symmetrical. P5
symmetrical, exopodite with 3 outer marginal spines, terminating in a bifurcated end, endopodite
small.
Male: In dorsal view posterior metasome segment pointed, rounded in lateral view; urosome 5segmented. Right A1 geniculate. P5 asymmetrical, the right leg claw like with a short triangular
thumb on the first exopodite segment, terminating in a spine which is as long as the preceding
segment. Left P5 terminal segment with 4 characteristic long spines, one of which is longer.
Recorded: Seychelles, Zuza, Madagascar, CPR.
B
G
C
D
Labidocera detruncata: Female. A, dorsal; B, urosome ventral; C, urosome lateral; D, urosome

dorsal; E, P5. Male. F, body lateral; G, P5. Figures B-E, G from Bradford Grieve (1999), A, F
from Mori (1937).
128
Labidocera kryeri (Brady, 1883)

Female: Cephalosome with lateral cephalic hooks, lateral wings of last metasome segment pointed
and symmetrical. Urosome 3-segmented; anal segment short, genital segment and urosome
segment 2 asymmetrical, with many spines and hooks (variable in appearance); furca symmetrical.
P5 endopodite reduced and bifurcate at the end.
Male: Lateral cephalic hooks on cephalosome. Lateral corners of the last metasome segment
asymmetrical, left one pointed the right one bifurcated. 5-segmented urosome, genital segment
swollen on left side. Right A1 geniculate. Right P5 exopodite segment 1 claw- like, with curved
thumb, middle portion of the external margin with 2 processes.
C
A
Labidocera kryeri: Female. A, dorsal; B, urosome lateral; C, urosome dorsal; D, P5. Male. E,
body dorsal; F, urosome lateral; G, urosome dorsal; H, P5. Figures from Greenwood (1979).
129
Labidocera laevidentata (Brady, 1883)

Length: females: 1.60-1.95 mm; males: 1.70 mm. (Silas and Pillai (1973) give lengths of 2.202.36 mm for females).
Females: Lateral cephalic hooks on cephalosome closer to anterior than in most other species;
posterior metasome symmetrical with divergent pointed wings. Urosome 3-segmented, genital
segment almost symmetrical, with a pair of posterior dorsal spines. Urosome segment 2 and anal
segment asymmetrical; anal segment shorter on right side; furca asymmetrical, right furca broader
and longer than left. Dorsal posterior margin of urosome segment 2 with a bifurcated spine,
ventral lateral margin with a fringe of small spines. P5 slightly asymmetrical, right exopod shorter
and stouter than left. Exopodites with 3 outer and 2 inner spines, with a curved spine on end.
Endopodites reduced, short with a weakly bifurcate tip.
Males: Elongate body, posterior metasome asymmetrical, one spine on right, 2 on left. Cephalic
hooks present. P5 with hook on right side; left P5 terminating in 3 unequal crooked spines, the
largest at right angles to the other 2 spines.
B
C
Labidocera laevidentata: Female. A, dorsal; B, urosome lateral; C, P5. Male. D urosome dorsal;
E, P5. Figures from Scott (1909).
130
Labidocera madurae A. Scott, 1909

Females: Prosome elongated and narrow, rounded anteriorly, without cephalic hooks; dorsal eye
lenses feeble and small. Last metasome segment pointed laterally and symmetrical. Short 2segmented urosome; genital segment quite large and asymmetrical, bulging on the right side.
Furca symmetrical, longer than broad. P5 nearly symmetrical, exopod with 2 outer spines,
terminating in 3 unequal spines. Endopodite simple and not bifurcated.
Males: Resembles female, but eye lenses slightly larger, urosome 5-segmented, the fourth and
fifth shorter than the others. Right A1 geniculate but only slightly swollen. Thumb-like process on
the first segment of the right exopodite, short and curved, no tooth at the base of the process as in
L. bataviae. Terminal joint long and narrow. Terminal joint of left P5 elongate and oval, twice as
long as broad, outer margin with 3 spines, inner with a fringe of hairs.
B
C
Labidocera madurae: Female. A, dorsal; B, P5. Male. C, urosome dorsal; D, geniculate A1; E, P5.
Figures from Scott (1909).
131
Labidocera minuta Giesbrecht, 1889

Length: females: 1.76-2.26 mm; males: 146-1.83 mm.
Females: Cephalosome narrow and rounded, small dorsal eye lenses; small lateral hooks present.
Lateral posterior margin of metasome rounded, but extended on the right side with a ventrally
pointing spine; left side without any spine; urosome 3-segmented, genital segment elongated and
half the length of urosome; right anterior and also posterior corner of ge nital segment with lateral
swellings, in the proximal centre of the ventral surface is a small chitinous outgrowth. Right
ventral surface of urosome and second urosome segment with chitinous knobs along right margin;
furca asymmetrical, right side larger. P5 symmetrical, exopodite bifurcated at tip; the endopod is
normally bent outwards over the exopodite, 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).
Males: 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 5-segmented, genital segment 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. Recorded: Seychelles,
Madagascar.
A
B
D
Labidocera minuta: Female. A, dorsal; B, urosome dorsal; C. urosome lateral; D, P5. Male. E,
body dorsal; F, P5; G, terminal segment of P5 right; H, P5. Figures A from Giesbrecht (1892); B,
D-G from Greenwood (1979), C, H from Silas and Pillai (1973).
132
Labidocera pavo Giesbrecht, 1889

Female: Body robust, no lateral hooks on cephalosome; dorsal eyes quite well developed;
posterior metasome segment pointed and symmetrical; urosome short, 2-segmented; right side of
genital segment has a process with a spine on the end, ventrally the posterior margin produced into
a long lobe, second urosome segment short. Some specimens with a transparent elliptical plate on
ventral surface of urosome. Furca nearly symmetrical; P5 almost symmetrical, exopod with 2
outer marginal spines and terminates in 3 spines, endopod is simple and not bifurcated, left
endopod quite rounded and right ending in a sharp point.
Male: Dorsal eye lenses arranged close together. Posterior metasome segment symmetrical with
lateral points. Urosome 5-segmented. Urosome and furca quite symmetrical. Right A1 geniculate.
P5 asymmetrical; right exopod claw- like with well developed thumb, towards inner base of thumb
is a robust seta; terminal segment elongated, curved, with a blunt conical projection on proximal
inner margin, with 2 inner mid- marginal seta, terminates in a point with one terminal seta; left leg
terminal segment with 1 outer marginal spine and 3 terminal sub-equal spines, all curving inwards.
D
C
A
B
Labidocera pavo: Female. A, dorsal; B, P5. Male. C, body dorsal; D, P5. Figures A-C from Mori,
(1937); D from Silas and Pillai (1973).
133
Genus Pontella
Cepalosome with lateral hooks, usually without a crest; with one pair of cuticular lenses and
usually with an additional lens in rostrum; last metasome segment usually with pointed lateral
lobes; female urosome 2 or 3-segmented and asymmetrical, male urosome 4 or 5-segmented,
symmetrical. Right male A1 geniculate. Female P5 biramous with exopodites and endopodites of
1 segment, male P5 uniramous.
Pontella fera Dana, 1849
Female: Rostrum slender and pointed with weakly developed rostral lens, dorsal eye lenses
moderately large; posterior corners of last metasome segment asymmetrically produced into winglike processes which extend beyond middle of genital segment; urosome 2-segmented, genital
segment asymmetrical, bulging laterally on left side; ventrally the genital segment has 2 knob- like
processes, varying in size and shape. Furca asymmetrical, left furca larger. P5 symmetrical,
endopodite short and bifurcated at tip, variable in appearance. Exopodite elongated, with 3 spines
on both inner and outer margins, terminating in another spine, outer marginal spines more widely
spaced.
Male: Cephalosome as in female; last metasome segment ending in bluntly rounded posterior
projections of more or less equal size. A1 geniculate; urosome 5-segmented, genital segment with
a small lateral bulge on right side, furca asymmetrical slightly broader on right. Right P5 with
claw. Base of claw with a large projecting thumb with a small process at its base and a short and a
large process between this and the terminal segment. Terminal segment initially almost straight,
but the distal half curves backwards, the tip bearing a small seta. Left P5 terminating in 3 unequal
processes.
Recorded: Zuza.
D
G
Pontella fera: Female. A, dorsal; B urosome dorsal; C, urosome lateral; D, rostrum; E, P5. Male.
F, geniculate A1; G, P5. Figures A from Giesbrecht (1892), B-F from Silas and Pillai (1973).
134
Genus Pontellina
Pontellina plumata (Dana, 1849)
Female: Prosome in dorsal view quite oval; posterior metasome segments almost symmetrical and
conspicuously pointed in lateral view; cephalosome without lateral hooks, without cuticular
lenses, with inconspicuous ventral eye without a lens. The A1, A2 and furca bear very long
plumose setae. Urosome 2-segmented, right furca fused to anal segment; P5 with 1-segmented
exopodite, bearing one lateral and 3 terminal seta, as well as 1 medial seta; endopodite is
bifurcated, or may come to a single point.
Male: Posterior metasome segments symmetrical, less pointed than in the female when viewed
laterally; one pair of cuticular lenses. A1 geniculate. P5 uniramous; right P5 ending in a claw, base
of claw broad, with a conical tooth- like elevation on inner margin and a seta. Left leg distal
segment with prominent outer marginal spine and 3 terminal subequal spines, inner margin with a
tuft of hairs.
Recorded: Seychelles, Zuza, Madagascar, Aldabra, CPR.
F
D
Pontellina plumata: Female. A, dorsal; B, P5. Male. C, body dorsal; D, body lateral; E, P5; F, P5
left terminal segments. Figures A, B from Bradford-Grieve (1999); C-F from Fleminger and
Hlsemann (1974).
135
Genus Pontellopsis
Head without lateral hooks, dorsal or rostral lenses. Last metasome segment with lateral points,
usually asymmetrical in the male. Urosome asymmetrical in both sexes, 1 or 2-segmented in the
female, 5-segmented in the male. The male right A1 is geniculate, the terminal section with 2
segments. Male urosome segment 3 with a projection on the right.
Pontellopsis armata (Giesbrecht, 1889)
Female: Cephalosome broadly rounded anteriorly; posterior metasome symmetrical, produced into
long lateral, sharply pointed projections. Urosome 2-segmented, genital segment with a
backwardly pointed spine above the genital opening and a dorsal swelling on the right side. Anal
segment symmetrical, dorsally overlapping the furca. P5 asymmetrical, exopodite stout and curved
inwards, with 3 minute external marginal spines and 3 apical spines, the middle the longest.
Endopodite bifurcated apically.
Male: Cephalosome resembles female. Lateral angles of the last metasome segment are
asymmetrical, the left side produced into a pointed straight spine reaching the third urosome
segment, the right side bluntly rounded, bearing a thin pointed curved process, longer than the
spine on the left. Urosome 5-segmented, with spine on the dorsal right genital segment; right side
of segment 3 with large protuberance with spinules on tip. P5 asymmetrical, the right leg with a
claw, basis broad in middle and with 2 sub-equal seta. Short thumb on claw segment, with a long
seta at the base; terminal segment slightly bent, with 2 setae at proximal inner end and one distal
seta, a short spine at outer base of segment.
Recorded: Zuza.
Pontellopsis armata: Female. A, dorsal; B, urosome lateral; C, P5. Male. D, body dorsal; E, P5; F,
P5 left, terminal segments. Figures A-C, E, F from Silas and Pillai (1973); D from Mori (1937).
136
Pontellopsis macronyx A. Scott, 1909

Female: Last metasome segment with 2 robust points. Genital segment long, bearing four dorsal
spines, the posterior left one larger than the others. (Silas and Pillai (1973) only found 3 spines on
the genital segment, the largest one was missing). Furca short and symmetrical. P5 is small, robust
and asymmetrical, longer on right. Right exopodite has 2 outer marginal spines, 2 small and 1
large terminal spine and a very long spine on the inside proximal margin. The left exopodite
resembles the right, but does not have the large inner spine. Endopodites are bifurcated at tip.
Male: Cephalosome is similar to female. Last metasome segment pointed on the left, reaching the
posterior of urosome segment 1, with a long thin process on the right, reaching the fourth urosome
segment. Urosome 5-segmented, segment 1 with a small spine on right, segment 3 with a well
defined blunt process on the right lateral margin. Right A1 geniculate. P5 right terminal segment
claw- like; thumb slender, curved and very elongated, with a seta at the base; terminal segment of
leg short and curved inwards, with 2 outer marginal setae and one inner mid- marginal seta situated
on a raised process. Left P5 with 3 terminal spines.
B
D
Pontellopsis macronyx: Female. A, dorsal; B, urosome lateral; C, P5. Male. D, urosome dorsal; E,
geniculate A1; F, P5. Figures A, D-F from Scott (1909); figure B from Silas and Pillai (1973).
137
Family Temoridae
Genus Temora
The cephalosome in Temora is typically the widest and deepest part of the prosome and they
have very long furca. P5 of female small, uniramous, 3-segmented; P5 male asymmetrical, left
leg larger and 4-segmented, segment 2 produced on inner edge into a long curved thumb-like
process; right P5 3-segmented, terminal segment curved inwards, claw-like.
Temora discaudata Giesbrecht, 1889
Female: Quite a large robust species; posterior metasome segments produced into spines. This
species bears some resemblance to T. stylifera and several authors have recorded this latter
species as being present in the Indian Ocean. However, as Arcos and Fleminger (1991)
suggested, the records are misidentifications of immature specimens. T. discaudata immature
specimens have an extension of the cephalosome laterally over the metasome segments, which is
similar to the segment arrangement found in adult T. stylifera. However, in adult T. discaudata
this cephalosome extension is not present.
In T. discaudata the anal segment and long slender furca are asymmetrical. P5 uniramous and
symmetrical, inner spine of segment 3 longer than the 2 terminal spines, which are almost equal
in length.
Male: Lateral angles of the last metasome segment pointed and slightly asymmetrical; right A1
geniculate. Urosome of 5 segments, almost symmetrical, with long slender furca. 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.
D
E
C
Temora discaudata: Female. A, dorsal; B, P5. Male. C, body dorsal; D urosome dorsal; E, P5.
Figures A, from Giesbrecht (1892); B, D, E from Greenwood (1978); C from Mori (1937).
138
Temora turbinata (Dana, 1849)

Female: Body widest at cephalosome, tapering anteriorly to urosome; last metasome segment
rounded. Urosome 3-segmented; anal segment symmetrical, shorter in length than previous
segment; furca almost symmetrical, very long and slender. P5 symmetrical, inner marginal spine
of the 3 terminal spines shorter than the other two.
Male: body shape similar to female; Urosome quite symmetrical; furca long and slender; P5
asymmetrical, the left with a claw; thumb of claw is slender and gradually curved.
E
F
G
Temora turbinata: Female. A, dorsal; B, lateral; C, urosome; D, P5. Male. E, body dorsal; F
body lateral; G, P5. Figures from Bradford-Grieve (1999).
139
Family Tortanidae
Genus Tortanus
Copepods with a median eye, but no lateral hooks or cuticular lenses; no rostrum. Well
developed maxilliped associated with their carnivorous diet. Right A1 of male geniculate with
swollen middle section. Urosome long, 2 or 3-segmented in female, 5-segmented in male. Furca
sometimes asymmetrical. P5 uniramous in both sexes; the male right leg forms a claw.
Tortanus barbatus (Brady, 1883)
Female: Urosome 3-segmented; anal segment and furca fused, asymmetrical; segment 3 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: Anal segment and furca of urosome fused, slightly asymmetric. Right P5 second segment
with a triangular projection with a seta half way along inner margin; left P5 penultimate segment
with no setae, terminal segment with a long outer seta, numerous spinules on the inner and outer
distal edges.
C
F
I
J
H
Tortanus barbatus: Female. A, dorsal; B, urosome dorsal; C, urosome dorsal of broad ramus
form; D, urosome lateral of broad ramus form; E, P5 with left leg variant. Male. F, dorsal; G,
urosome dorsal; H urosome lateral; I, P5 posterior view; J, P5 anterior view. Figures A, F from
Steuer (1926); B-E, G-J from Greenwood (1978).
140
Tortanus forcipatus (Giesbrecht, 1889)

Female: Urosome 3-segmented, anal segment and furca fused, asymmetrical. In la teral view the
third urosome segment 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: Anal segment and furca of urosome fused, slightly asymmetric. Right P5 second segment
with a pointed triangular projection; left P5 terminal segment curved, with an inner and outer
marginal spine.
B
A
D
E
Tortanus forcipatus: Female. A, dorsal; B, lateral; C, P5. Male. D, body dorsal; E, P5. Figures AC from Giesbrecht (1892); D, E from Steuer (1926).
141
Tortanus gracilis (Brady, 1883)

Female: Posterior of last metasome segment rounded, knob- like. Urosome 3-segmented; anal
segment and long furca fused, asymmetrical; anal segment very narrow in dorsal and lateral
view, reminiscent but differing from T. forcipatus. P5 slightly asymmetrical, longer on left,
terminal segments with 2 small spines on distal outer margin. Steuer (1926) drew a seta on the
outer middle margin of each basis, not figured in Bradys original drawings reproduced here.
Male: Urosome slightly asymmetric. Right P5 second segment with a pointed triangular
projection with tooth just internal to the point; left P5 longer than right, terminal segment curved,
inner margin with a fringe of fine hairs and 2 small spines. Steuer (1926) drew this outer
segment with 1 inner and 1 outer marginal spine.
A
D
C
F
Tortanus gracilis: Female. A, lateral; B, urosome dorsal; C, urosome lateral; D, P5. Male. E,
dorsal; F, P5. Figures from Brady (1883).
142
Tortanus insularis Ohtsuka and Conway, 2003

Large robust species, with a bulbous forehead that is tinged red in fresh specimens.
Female: Prosome approximately three times longer than urosome. Posterior metasome nearly
symmetrical in dorsal view. Urosome 3-segmented, genital segment as long as remainder of
urosome, slightly asymmetrical, bulging anterolaterally on both sides, with a small spine on tip
of bulges, downward pointing projection on ventral surface. Anal segment very asymmetric,
protruding posterolaterally on left side; furca very asymmetric with proximal half of left ramus
swollen dorso-ventrally. P5 slightly asymmetric, with right leg longer than left; 1-segmented
exopodite bearing 1 inner and 2 terminal pointed spines; outer terminal spine with 2 minute
spines on anterior surface.
Male: Body narrower than the female; posterior metasome rounded and symmetrical. Genital
segment slightly asymmetrical; right A1 geniculate. Right P5 very robust; basis massive, bearing
large inner bilobed process at mid- length. Left P5 with elongated basis, slightly curved inwards,
with 1 inner and 1 outer setule; exopodite 2-segmented, proximal segment with inner proximal
round process bearing a setule and with tiny spinule on outer distal edge; outer segment with
dense patch of setules along proximal half of outer margin, row of finer setules along distal half,
terminating in a sharp point.
Recorded: Rodrigues
K
J
H
Tortanus insularis: Female. A, dorsal; B, lateral; C, urosome lateral; D, P5. Male. E, dorsal; F,
urosome lateral, G, A1; H, right furca; I, P5; J, right P5; K, distal exopodite segment of left P5.
Figures from Ohtsuka and Conway (2003).
143
Family Calanidae
Head and first pedigerous segment usually fused, pedigerous segments 4 and 5 always separate.
Rostrum composed of 2 filaments. Generally with two characteristic, strong, backwardly
directed setae on the third and second last segments of the A1. The various genera are mainly
based on characters of the male P5. Females have 4 segments in the urosome, 5 swimming legs
of similar size and appearance with both rami 3-segmented. Exopodal joints 3 of P2-P4 each
with 2 spines on outer edge and 5 setae on inner edge. Male A1 are alike and only slightly
transformed, never exhibiting any geniculate structure, 25 segmented with segments 1 and 2
always fused. Males usually with a hump on mid-dorsal region. P1-P4 as in the female, the right
P5 is usually similar in appearance and size to the other swimming legs (less so in Undinula)
while the left varies from being similar (Calanus) to drastically modified (Cosmocalanus).
Urosome 5-segmented.
Genus Canthocalanus
Only one species.
Canthocalanus pauper (Giesbrecht, 1888)
Female: Similar in shape to Nannocalanus minor, but urosome is not as deeply inset into
metasome, and the inner margins of the P5 coxa are not denticulate. Head and pedigerous
segment 1 fused. Anterior cephalosome and posterior metasome rounded. Described as having a
slightly asymmetrical last metasome segment, however this does not appear to be the case with
specimens sampled. Urosome 4-segmented. Strong setae on furca.
Male: Right P5 asymmetric, exopodite no inner marginal spines; left endopodite with 2 terminal
setae; left exopodite with elongated segments with long outer distal setae on segments 2 and 3.
Inner margins of the P5 coxa are not denticulate
D
A
E
C
B
Canthocalanus pauper: Female. A, dorsal; B, P1 endopod removed; C, P5. Male. D, body dorsal;
E, P5. Figures from Mori (1937).
144
Genus Cosmocalanus
Only 2 species, although the separate validity of C. caroli is in some doubt.
Cosmocalanus darwini (Lubbock, 1860)
The furcal setae sometimes branch. Inner margin of P5 coxa finely serrated.
Female: The 5 swimming legs are the same size and similar structure. The last metasome
segment comes to a blunt point when viewed laterally. Short spinules round the posterior
margins of the genital and second urosome segments, with a prominent tuft of longer spinules on
the dorsal edge of the second urosome segment. Sometimes spinules are only present along the
posterior left side of the genital segment. Genital segment in lateral view bulges quite strongly
and comes to a blunt point towards the anterior segment. A1 reaches almost to the end of the
urosome. The posterolateral metasome borders may take 3 forms (symmetrical, forma
symmetrica; very asymmetrical with left border produced into a squared-off lappet, forma typica;
an intermediate form which is slightly asymmetrical, forma intermedia).
Male: Extremely large P5, very asymmetrical, sometimes projecting from the side of the body.
Similar in size to the male P5 of Undinula vulgaris but different structure. The tooth on the inner
edge of the external spine, which is part of the last segment of the left P5, is placed about one
third of the length from the base of the spine.
Recorded: Rodrigues, Seychelles, Zuza, Madagascar, Aldabra, CPR.
D
F
Cosmocalanus darwini: Female. A, lateral; B, urosome lateral; C, urosome lateral of forma

typica; D, P3; E, P5. Male. F, body dorsal; G, P5; H, Distal portion of left P5; I, right P5. Figures
from Bradford-Grieve (1994).
145
Cosmocalanus caroli (Giesbrecht, 1888)

Length: females: ? mm; males:? mm.
The separate status of this species is in some doubt (Bradford-Grieve, 1994), as it is very similar
to C. darwini. There appear to be a difference between males, which is described.
Female: Very similar to C. darwini. Wolfenden (1906) found a female which differed, as the
denticulation of the last joint of the exopodite of the P2 and P3 (external margin) was absent.
This specimen also had 3-4 fine teeth on the inner margin of the coxa of the P1. He suggested it
might be C. caroli.
Male: The position of the tooth on the inner edge of the external spine, which is part of the last
segment of the left P5, is the specific character distinguishing this species from C. darwini. In C.
caroli the tooth is placed about one sixth of the length of the spine from the base of the spine and
in C. darwini the tooth is one third from the base of the spine.
Cosmocalanus caroli: Male. Terminal portion of left P5. From Wolfenden (1906).
146
Genus Mesocalanus
In both sexes pedigerous segment 1 separate; P5 coxa with a smooth inner margin, endopodite
with a total of 7 setae. Male P5 with both rami 3-segmented. Only 2 species.
Mesocalanus tenuicornis (Dana, 1849)
Female: Prosome quite slender. The lateral angles of the last metasome segment are rounded.
The furca and seta are symmetrical. The A1 are around twice the length of the prosome. The
internal margin of the P5 coxa is not denticulate.
Male: The A1 are around twice the length of the prosome. The P5 are only slightly asymmetric,
left P5 terminal segment with spine almost as long as the segment. Furca divergent.
Recorded: Rodrigues, Madagascar.
Mesocalanus tenuicornis: Female. A, dorsal; B, lateral; C, P5. Male. D, body dorsal; E, P5.
Figures from Bradford-Grieve (1994).
147
Genus Nannocalanus
There was only one species but a new species, N. elegans, has just been described by Andronov
(2001).
Nannocalanus minor (Claus, 1863)
Length: females: 1.70-2.30 mm; males: 1.20-2.01 mm
Female: Short stubby urosome, which is characteristically inset into the metasome, the rounded
last metasome segment extending almost halfway down the genital segment. The genital segment
bulges strongly when viewed from the side, with a small projection low on the surface. In fresh
specimens the edges of the prosome segments are sometimes tinged with red. The female has 5
swimming legs which are all the same size and of similar structure. The P5 coxa has serrations
on the inner surface. These serrations separate it from a closely related species that also occurs in
this region, Canthocalanus pauper, which does not have these serrations. The A1 reach almost to
the end of the urosome.
Male: Similar in appearance to the female but with a longer urosome. Furca divergent in dorsal
view. P5 slightly asymmetric, right P5 similar to other swimming legs, with spines on the inner
border of the exopodite; left endopodite with no spines on internal edge, left exopod with
elongated outer edge spines. The P5 coxa has serrations on the inner surface.
Recorded: Rodrigues, Seychelles, Zuza, Madagascar, Aldabra, CPR, Mauritius.
A
B
Nannocalanus minor: Female. A, lateral; B, P5. Male. C, body lateral; D, P5. Figures from
148
Genus Neocalanus
Head and pedigerous segment 1 usually fused in female, separate in male. Swimming le g 2 in
both sexes with a recurved spine at the outer distal border or exopodite segment 1. Coxa of P5
inner border without serrations in both sexes. Male leg 5 with both exopodite and endopodite 3segmented; left leg modified, endopodite usually with 8 setae; right leg unmodified, or with
inner edge setae of exopodite absent.
Neocalanus gracilis (Dana, 1849)
Length: females: 2.43-4.00 mm; male: 2.3-3.10 mm.
Female: Body robust; head and pedigerous segment 1 fused; 5 pairs of swimming legs, all of
similar structure and size. A1 is approximately twice the length of the prosome. The urosome is
approximately one third the length of the prosome; genital segment, in lateral view, rounded and
protruding on distal portion of the segment; furca short, second inner marginal seta on left side
very long (if not damaged). Basis of P1 each with a prominent hook on anterior surface.
Male: Head and pedigerous segment 1 separate. Swimming legs are all very similar; right P5
same general structure as other swimming legs, terminal exopodite segment with setae on the
inner edge; left P5 with first 2 exopodite segments elongated, the terminal segment with 2
spines; the 3-segmented endopodite is not well developed and in some specimens the number of
seta can be greatly reduced. A1 much shorter than in the female, reaching halfway down
urosome. Urosome approximately one third length of prosome. Small process close to dorsal
posterior margin of the cephalosome, further forward than the same process in N. robustior.
Recorded: Rodrigues (offshore), Zuza, Madagascar, CPR.
C
B
E
Neocalanus gracilis: Female. A, dorsal; B, urosome lateral; C, P1. Male. D, body dorsal; E, P5.
Figures from Giesbrecht (1892).
149
Neocalanus robustior (Giesbrecht, 1888)

Female: Distinguishable from Neocalanus gracilis by its usually larger size, the shape of the
genital segment which is squarer, convex and more prominent in lateral view. Head and
pedigerous segment 1 fused. The basis of the P1 has a process on the anterior surface, similar to
that of N. gracilis.
Male: Head and pedigerous segment 1 separate. Small process close to the dorsal poster margin
of the cephalosome. The left P5 with a rudimental 3-segmented endopodite, the distal segment of
which in some specimens has 2 or 3 setae; the exopod with very elongated segments; right P5
similar to the other swimming legs.
Recorded: Seychelles, Rodrigues, CPR.
E
C
Neocalanus robustior: Female. A, dorsal; B, lateral; C, urosome lateral; D, P1; E, P5. Male. F,
body dorsal; G, P5. Figure A from, Wolfenden (1906); B- E, Mori (1937); F, G, Park (1968).
150
Genus Undinula
Only one species
Undinula vulgaris (Dana, 1849)
Female: Head and pedigerous segment 1 fused. 5 swimming legs, all the same size and also
similar structure. Similar to Cosmocalanus darwini but the last metasome segment comes to a
sharp point in lateral view. Some specimens have two points on the left side. The A1 reaches
almost to the end of the urosome. P2 with a notch on the external margin of the second segment
of the exopodite. Coxa of the P5 without serrations on the internal margin, endopodite with 7
setae, those on segments 1 and 2 spine- like.
Male: Head and pedigerous segment 1 fused. P2 with a notch on the external margin of the
second segment of the exopodite. Has an extremely large P5, similar in size to that of C. darwini
and very asymmetric. Coxa of the P5 without serrations on the internal margin. P5 left is highly
modified, without endopodite and bears two large spikes on exopod segments 1 and 2 and a
structure like a wrinkled trunk on the end of the limb. Right P5 is tiny and has an endopodite.
Recorded: Rodrigues, Seychelles, Zuza, Aldabra, CPR.
A
B
Undinula vulgaris: Female. A, lateral; B, P2; C, P5. Male. D, body lateral; E, P5. Figures A, B
from Bradford-Grieve (1994); D from Mori (1937); E, from Dakin and Colefax (1940).
151
Family Mecynoceridae
Genus Mecynocera
Mecynocera clausi Thompson, 1888
Female: This is a very small species that initially may be confused with Calocalanus, but the
prosome is 6 segmented while in Calocalanus it is 4 or 5-segmented. The A1 is over twice the
length of the whole body and plumose, but is often broken off. The limbs seem also easily
damaged. The prosome is separate from the segment bearing the P1. The posterior metasome
segments are rounded. The urosome is 3-segmented; the genital segment is symmetrical and very
swollen in lateral view. P5 are very simple, uniramous with straight symmetrical limbs, with 5
segments on each side, with 1 seta on the fourth and 3 inner and 2 terminal setae on the fifth
segment.
Male: Dorsal hump on cephalosome. A1 shorter than in female, extending slightly beyond the
end of the urosome. Urosome 5-segmented; P5 are very simple quite straight limbs, with 5
segments on each side, asymmetrical, longer on right. Right P5 inner border hairy; terminal
segments on both legs with a stout terminal spine and a small disto- lateral spinule. End of left P5
reaches to middle of segment 4 of right P5.
A
G
C
Mecynocera clausi: Female. A, dorsal; B, lateral; C, P5. Male. D, body lateral; E, body dorsal; F,
urosome; G, P5. Figures A, B from Bradford-Grieve (1994); D-G from Corral (1972); C from
Giesbrecht (1888).
152
Family Paracalanidae
Cephalosome and first pedigerous segment us ually at least partially fused, pedigerous segments
4 and 5 fused or separate. In the female the urosome is 2-4 segmented, the anal segment usually
much longer than any segment between it and the genital segment. Exopodites of P1-4 3segmented, external border of P1-P4 usually fringed with small spines; endopodite of P1 usually
2-segmented, those of P2-4 3-segmented. Distal spine of exopods with smooth blade-like border.
P5 when present is uniramous and symmetrical. Male urosome 5-segmented; dorsal hump
usually present on cephalosome. P1-4 similar to female, P5 uniramous.
Genus: Acrocalanus
Small copepods, rostrum with long slender filaments. Pedigerous segments 4 and 5 incompletely
separated. Obvious small spines on outer edge of exopodite segments of P2-4. Females with a
short urosome and superficially resemble Paracalanus, but they only have four pairs of
swimming leg, which are all of similar size and structure, although sometimes there is a
rudimentary fifth pair present. The male P5 is only present on the left. Males of the different
species are difficult to tell apart.
Acrocalanus gibber Giesbrecht, 1888
Female: Deep body in lateral view, with a pronounced hump on the dorsal cephalosome; distinct
though partial line of separation of the cephalosome from the first pedigerous segment is
obvious; A1 reaches just to the furca, or slightly beyond. 4 pairs of swimming legs, the distal
toothed outer border of exopodite segment 3 of P4 is about 60% of the length of the proximal
part of the toothed border; the distal teeth are strong compared to the fine teeth in A. longicornis.
Genital segment longer than any of the others, with a prominent ventral swelling.
Male: Distinct hump on the dorsal cephalosome; 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.
Acrocalanus gibber: Female. A, lateral; B, P4. Male. C, lateral; D, P4. Figures from Sewell
(1929).
153
Acrocalanus gracilis Giesbrecht, 1888

Female : The anterior cephalosome is evenly round in lateral view and the line of separation of
the cephalosome from the first pedigerous segment is very slight, if it can be seen at all; no
obvious hump on cephalosome; A1 exceeds the end of the furca by 2-4 terminal segments; the
distal toothed outer border of exopodite segment 3 of the P4 is about 80% of the length of the
proximal part of the toothed border. The coxa of P4 in more convex than in A. longicornis and
there are more spines distally.
Male: The anterior cephalosome is evenly round in lateral view and there is no hump. The P5 is
asymmetrical, when fully extended reaches the end of urosome segment 2, the right foot is
vestigial or absent, the left P5 is of 5 segments, the distal the largest. A1 reaches just beyond the
furca, although the drawing by Mori (1937) suggests it being shorter than this?
Recorded: Seychelles, Zuza, Madagascar.
A
B
Acrocalanus gracilis: Female. A, lateral; B, P4. Male. C, lateral; D, P4; E, P5. Figures A, B, D
from Sewell (1929); C, E from Mori (1937).
154
Acrocalanus longicornis Giesbrecht, 1888

Female: Can be distinguished from other species as the A1 extends further beyond the furca than
in other species, by its last 5 segments. The anterior cephalosome is rounded in lateral view but
the dorsal surface is quite strongly arched at the level of the mouth, but less so than in A. gibber.
There is reported to be a partial segmentation between the cephalosome and the first pedigerous
segment (Wolfenden, 1906; Sewell, 1929). Genital segment broader than long, laterally swollen
and projecting ventrally. Outer distal border of exopodite segment 3 of P4 with over 18 very
small teeth and with coarse larger teeth on the proximal border.
Male: P5 only present on the left; 5-segmented. A1 only extends just beyond the posterior
metasome. Cephalosome partially separated from the first pedigerous segment.
B
A
Acrocalanus longicornis: Female. A, dorsal; B, P4. Male. C, lateral; D, P5. Figures A, B from
Bradford-Grieve (1994); C from Sewell (1929); D from Wolfenden (1905).
155
Acrocalanus monachus Giesbrecht, 1888

Length: females: 0.90-1.10 mm; males ? mm.
Female: Similar to A. gracilis but in lateral view the cephalosome of the female produced
forward forming a sort of hood, anteriorly with a very characteristic blunt point which slopes
down in a straight line towards the rostrum, immediately distinguishing it from other members of
the genus. The anterior cephalosome in dorsal view is very narrow compared with the rest of the
prosome. No indication of any division between the cephalosome and the first pedigerous
segment and little evidence of a dorsal hump. A1 extends beyond the furca by at most, 2 or 3
segments. Outer distal border of exopod segment 3 of P4 with about 10-12 large teeth, similar to
the 16 on the proximal margin.
Male: Not described.
B
A
C
Acrocalanus monachus: Female. A, dorsal; B, lateral; C, P4. Figures from Grice (1961).
156
Genus Calocalanus
Females generally with a broad, bulbous genital segment when viewed from above. A1 often
much longer than the body. Prosome of 4 or 5 segments, compared to Mecynocera which has 6
segments. Female urosome 2 or 3-segmented, often 2 setae on the furca much thicker than the
others. The P5 is uniramous and simple, symmetrical, 3 or 4-segmented with varying number of
setae on the end of the terminal segment. Male P5 asymmetrical, uniramous, 4-segmented on the
right and 5 on the left.
Calocalanus pavo (Dana, 1849)
Female: Small quite transparent species; in well-preserved specimens have plumose A1 and
furca. Prosome stout, 3 times as long as wide. Cephalosome and first pedigerous segment fused.
Urosome 2-segmented, genital segment onion-shaped, furca symmetrical, elongate and typically
divergent, arranged almost at right angles to the urosome. P5 uniramous and symmetrical, 4segmented; terminal segment with 5 plumose setae and 1 spine on the outer distal border and 2
rows of tiny spinules.
Male: Body quite different from the female, but can also have plumose A1 and furca.
Cephalosome and first pedigerous segment are more or less separate, posterior metasome
extending laterally into almost square corners. Urosome 5-segmented, furca longer than wide,
not divergent. P1-4 as in the female. The P5 is small and biramous, limbs of very unequal
lengths. Both P5 legs terminating in 2 small sub equal spines; right leg does not extend as far as
the distal border of the second segment of the left leg.
Recorded: Seychelles, Madagascar
D
A
F
C
E
G
B
Calocalanus pavo: Female. A, dorsal; B, genital segment lateral; C, P5. Male. D, dorsal; E,
urosome; F, distal A1; G, P5. Figures A-C from Bradford-Grieve (1994); D-G from Corral
(1972).
157
Calocalanus styliremis Giesbrecht, 1888

Female: Wide robust prosome; cephalosome and first pedigerous segment fused; metasome 3segmented. Urosome short, 3-segmented; genital segment swollen like an onion; furca
symmetrical and typically divergent; P5 3-segmented, uniramous, terminal segments with a spine
with 4 or 5 minute spinules at its base.
Male: Very different from the female. Cephalosome and first pedigerous segment fused.
Urosome 5-segmented, furca not divergent. Left P5 may be 4 or 5-segmented, right P5 is much
shorter and may be 3 or 4 segmented. These latter differences may be because closely related
species have been mistakenly examined (Bradford-Grieve, 1994).
B
C
Calocalanus styliremis: Female. A, lateral; B, dorsal; C, genital segment lateral; D genital

segment dorsal; E, P5. Male. F, dorsal; G, urosome; H-J, P5 variations. Figures A-E from
Bradford-Grieve (1994); F-J from Corral (1972).
158
Genus Ischnocalanus
Ischnocalanus plumulosus (Claus, 1863)
Female: Body slender, 4 times as long as wide. Cephalosome and first pedigerous segment fused.
Urosome 3-segmented, genital segment almost cuboidal in shape. Posterior border of genital
segment and urosome segment 2, lined with fine spinules; furca and their setae asymmetrical, in
good specimens a very long, plumose seta on the left. Furca only slightly divergent Terminal
segment of A1 twice the length of the penultimate segment. P5 of 4 segments, the terminal
segment with 1 plumose seta between 2 sharp setae, 1 external spine and a horizontal row of fine
spinules. The last 2 segments of P5 are hairy.
Male: Cephalosome and first pedigerous segment not fused. A1 exceeds the furca by the distal 5
segments. P5 left terminating in a single spine, P5 right terminating in 2 spines and reaching
beyond the second segment of the left leg.
E
B
Ischnocalanus plumulosus: Female. A, dorsal; B, dorsal; C, genital segment lateral; D, P5. Male.
E, dorsal; F, P5. Figures A, C-F from Bradford-Grieve (1994); figure B from Giesbrecht (1892).
159
Genus Paracalanus
Quite small oval copepods; cephalosome fused with first pedigerous segment; rostrum composed
of 2 fine filaments; A1 extends beyond the metasome. Posterior metasome segments rounded.
Exopodites of P3 and P4 with serrated outside margin. Female with 4-segmented urosome, short
genital segment and a tiny, simple P5, which is slender, short, uniramous, symmetrical, 2segmented with 2 sub-equal terminal spines. P5 in the male it is uniramous, asymmetrical, with 5
segments on the left and 2 on the right. The right leg is very similar to the female P5 legs.
Paracalanus aculeatus Giesbrecht, 1888
This species is sometimes sampled in 2 separate size classes.
Female: Urosome roughly a third of the length of the prosome. The longest furcal setae, when
intact, are longer than the urosome. Similar to P. parvus, but the A1 extends beyond the furca.
P5 symmetrical, uniramous, 3-segmented, the distal segment cylindrical, with a terminal spine
longer than it and an outer distal spine almost half its length.
Male: Furca almost as long as wide, anal segment shorter than urosome segment 4. A1 extends
almost as far as the furca. P5 extends beyond the posterior border of urosome segment 3; the 3segmented right leg is similar to the female P5 legs and extends well beyond the distal border of
segment 2 of the 5-segmented left leg.
F
E
Paracalanus aculeatus: Female. A, lateral; B, dorsal; C, P5. Male. D, lateral; E, dorsal; F, P5.
Figures from Vervoort (1963).
160
Paracalanus parvus (Claus, 1863)

Female: Cephalosome fused with the first pedigerous segment. A1 reaches around the middle of
the 4-segmented urosome. Urosome around a third the length of the prosome, genital and anal
segments about the same length. P5 uniramous and symmetrical, 2-segmented (one leg
sometimes 3-segmented), terminal segme nt with a slender spine at the tip and a much smaller
spine outside this.
Male: Slight ridge along the dorsal surface of the prosome. A1 quite robust. Urosome 5segmented, second segment the longest. P5 asymmetrical, uniramous, the right foot 2segmented, similar to the female P5 legs and the left 5-segmented. The right foot reaches the
middle of segment 2 of the left foot. Both feet terminating in 2 sub-equal spines.
Paracalanus parvus : Female. A, lateral; B, P5. Male. C, lateral; D, P5. Figures from Sars
(1903).
161
Family Eucalanidae
Mostly large copepods with elongated and often transparent bodies. Stage V females can have a
swollen genital segment and resemble a stage V, but the segmentation is not clearly defined and
if there is any uncertainty whether the specimen is mature, it probably is not. The female
urosome is very short, 3 or 4-segmented, 5-segmented in the male, the furca usually fused to the
anal segment in both sexes. Females often have a broad genital segment. Most have in dorsal
view, a typical triangular anterior cephalosome. Cephalosome and first pedigerous segment
fused, pedigerous segments 4 and 5 partially fused, rostrum with 2 long filaments. In some
species the first antenna reaches well beyond the urosome. Female P5 missing or only on one
side, or uniramous on the left and biramous on the right.
The number and arrangement of small integumental organs over the body is a useful character in
identification (see Bradford-Grieve, 1994) but requires special preparation of the specimens.
Genus Eucalanus
Female urosome 4-segmented. The posterior margins of pedigerous segments 2-4 and urosome
segment 1 without any spines. P5 missing in female, uniramous on both sides in the male.
162
Eucalanus elongatus (Dana, 1849)

Female: Characterised by the pointed postero-lateral corners of the metasome. Transparent body.
Anterior cephalosome triangular, urosome 4-segmented, genital segment longer than wide. Left
furca wider and longer than the right. A1 extending beyond the furca by around the last 4
segments. Second inner seta of right furca elongated, if not broken off.
Male: Also has lateral points on the posterior metasome. Transparent body, anterior cephalosome
bluntly rounded. A1 extending beyond the furca by around the last 4 segments. Urosome 5segmented, genital segment slightly wider than long. Both P5 legs 4-segmented, terminating in
spines; right leg shorter than the left, terminal segment reaching around the third segment of the
left leg.
Eucalanus elongatus: Female. A, dorsal; B, rostrum ventral. Male. C, dorsal; D, P5. Figures
163
Genus Pareucalanus
Includes the largest of the Eucalanidae. Female urosome 3-segmented and the posterior margins
of pedigerous segments 2-4 and urosome segment 1 without any spines. P5 missing in the
female, uniramous, simple in the male, the right leg shorter than the left, not extending beyond
the distal part of left leg segment 2.
Pareucalanus attenuatus (Dana, 1849)
Length: females: 3.55-7.00 mm; males: 2.95-6.04 mm. There appears to be small and large forms
and great variability in appearance of specimens. It is possible that some of the specimens are P.
sewelli (Fleminger, 1973) but they are very difficult to tell apart without detailed examination of
the integumental organs.
Female: In dorsal view, generally with a triangular, bluntly pointed cephalosome, very elongated
anterior to the A2 and sometimes narrows in this region. In lateral view the anterior cephalosome
is quite pointed. Posterior prosome corners rounded. A1 extends beyond the furca by the last 6-7
joints. No P5 present; genital segment is longer than wide; furca fused to the anal segment, one
of the left furcal setae longer than the others.
Male: Anterior cephalosome triangular. A1 extends beyond the furca by the last 6-7 joints.
Genital segment wider than long; P5 very simple, the left leg longer than the right, both legs of 4
segments and in both the terminal segment bears a spine. Furca fused to anal segment. One of the
left furcal setae longer than the others.
Pareucalanus attenuatus: Female. A, lateral; B, dorsal; C, head lateral; D, genital segment

lateral. Male. E, dorsal; F, P5. Figures A, B, E, F from Mori (1937); figures C, D from Vervoort
(1946).
164
Genus Subeucalanus
Female urosome 3-segmented, the posterior margins of pedigerous segments 2-4 and urosome
segment 1 without any spines. No P5 in female, uniramous in male, with only left leg present.
Subeucalanus crassus (Giesbrecht, 1888)
Female: Body robust, anterior cephalosome rounded and short. A1 reaches just beyond the furca.
Urosome 3-segmented, genital segment wider than long and markedly onion shaped. Furca
slightly asymmetrical, anal segment fused with the furca. The left second terminal seta of the
furca is slightly longer than that of the right.
Male: Does not have distinct secondary sexual characters. A1 reaches around the end of the
furca. Genital segment longer than wide. P5 right foot absent, left foot uniramous, 4-segmented,
with terminal spine.
C
E
Subeucalanus crassus: Female. A, dorsal; B, lateral; C, dorsal; D, genital segment dorsal; E

genital segment lateral. Male. F, dorsal; G, P5. Figures A, D, E from Bradford-Grieve (1994);
figures B, C, F, G from Fleminger (1973).
165
Subeucalanus mucronatus (Giesbrecht, 1888)

Female: Anterior cephalosome triangular and sharply pointed, both dorsally and laterally.
Genital segment longer than wide; furca asymmetrical. Second inner seta on the right furca
thicker than other setae and elongated, if not broken off.
Male: Anterior cephalosome rounded, quite square laterally. Right P5 absent. The terminal spine
on the left P5 is shorter than the terminal segment, which separates it from S. subtenuis. Second
inner seta on the right furca thicker and longer than other setae, if not broken off.
Recorded: Zuza, Madagascar, Aldabra.
Subeucalanus mucronatus: Female. A, dorsal; B, dorsal; C, lateral. Male. D, dorsal; E, P5.

Figures B, C from Fleminger (1973); figures A, D, E from Mori (1937).
166
Subeucalanus pileatus (Giesbrecht, 1888)

Female: Prosome slender, widest at the posterior of the cephalosome; anterior cephalosome
triangular, rounded anteriorly, more pointed than in E. subcrassus, but exact shape can be
variable; urosome short, one sixth the length of the prosome. Posterior metasome more or less
rounded. A1 passes the furca by around 5 segments. Urosome 3-segmented, genital segment
slightly broader than long, moderately swollen in dorsal view, widest in the centre, moderately
swollen in lateral view. The line of fusion between the furca and anal segment obvious. Furca are
almost symmetrical, the second seta on the left greatly lengthened, but only slightly thicker than
the other setae. P5 absent.
Male: Similar to the males of S. crassus but smaller. The P5 is 4-segmented and only present on
the left. The terminal spine is shorter than the terminal segment.
Recorded: Rodrigues, Zuza.
F
I
H
Subeucalanus pileatus: Female. A, lateral; B, dorsal; C-E dorsal and lateral views of head shape
variants; F, urosome ventral. Male. G, dorsal; H, head dorsal; I, P5. Figures A,B from Fleminger
(1973): C-I from Giesbrecht (1892).
167
Subeucalanus subcrassus (Giesbrecht, 1888)

Length: females: 1.93-2.88; males: 2.40 mm.
Female: Similar to S. crassus but slimmer, head more pointed and extends further forward in
lateral view, the second terminal seta on the left furca is much longer than the right seta and the
genital segment is narrower. Anterior of cephalosome rounded; A1 extends beyond the furca by
around 4 segments. Urosome 3-segmented, genital segment broader than long, with widest part
in dorsal view on the posterior half of the segment, compare to E. pileatus where it is wider in
the centre; furca fused with anal segment.
Male: P5 right absent, terminal seta on left P5 longer than the terminal segment. Second terminal
setae of the furca are asymmetrical, as in the female.
Subeucalanus subcrassus: Female. A, dorsal; B, dorsal head profile; C, urosome ventral; D.

anterior urosome and genital segment ventral; E, P3 endopod segment 2. Male. F, dorsal; G, P5.
Figures A-E, G from Greenwood (1976); F from Mori (1937).
168
Subeucalanus subtenuis (Giesbrecht, 1888)

Female: Similar to S. mucronatus, as the anterior of cephalosome triangular, but its apical
portion is less sharply pointed. Furca asymmetrical.
Male: Anterior cephalosome rounded. Resembles S. mucronatus but the terminal spine on the
left P5 is longer than the terminal segment. Also similar to S. crassus but has more developed
asymmetry of the setae of the furca, the left second seta is much longer and thicker than the right
seta.
Recorded: Rodrigues, Seychelles, CPR.
C
D
Subeucalanus subtenuis: Female. A, Lateral; B, dorsal; C, head dorsal; D head lateral; E,

urosome ventral. Male. F, dorsal; G, head dorsal; H, P5. Figures A from Fleminger (1973); B-H
169
Genus Rhincalanus
Long, very slim species with anterior cephalosome extended into a conical process, carrying on
the ventral surface the rostral filaments. Cephalosome and first pedigerous segment fused. A1 is
longer than the whole body. Metasome segments with lateral and dorsal spines. Urosome short,
3-segmented, the distal segment fused with the furca. The female P5 is uniramous, 3-segmented
and very simple. The male P5 is also a very simple limb, biramous on one side.
Rhincalanus nasutus Giesbrecht, 1888
Female: Strongly pointed anterior cephalosome, but less so than in R. rostifrons. Prosome long
and slender with lateral spines and dorsal pairs of spines on the posterior margin of the first 3
metasome segments. A1 around 1.5 times the length of the whole body. Urosome 3-segmented,
with pair of small spines on dorsal surface of genital segment. Rostral filaments situated under
the frontal projection and are hardly visible from above. P5 are uniramous and 3-segmented,
with one seta on the second segment internally and three terminal setae.
Male: Prosome similar to female, but A1 considerably shorter. Spines on the metasome segments
and on the first urosome segment similar to the female. Right P5 is uniramous, 3-segmented with
a slender curved terminal spine. Left P5 biramous, endopodite 3-segmented with segments of
almost equal length, exopodite 1-segmented with a long terminal plumose seta and outside of it a
naked, very short seta.
B
E
F
Rhincalanus nasutus: Female. A, lateral; B, dorsal; C, head ventral; D, head lateral; E, urosome
ventral; F, P5. Male. G, dorsal; H, P5. Figures A, C-F, H from Sars (1903); B, G from Mori
(1937).
170
Rhincalanus rostifrons (Dana, 1849)

Female: Extremely extended point on the anterior cephalosome, with prominent rostral filaments
which protrude laterally, visible in dorsal view. A1 around 1.5 times as long as whole body. P5
3-segmented, without seta on the second segment, the terminal segment extended into a stout
curved point, with an internal toothed spine.
Male: Right P5 uniramous, 3-segmented with the terminal segment extended into a curved spine.
Left P5 biramous with a 1-segmented exopodite bearing a curved terminal spine; a 2-segmented
endopodite, the first segment longer than the second, the endopodite around the same length as
the exopodite.
Recorded: Seychelles, Zuza, Madagascar, Aldabra, CPR.
C
D
Rhincalanus rostifrons: Female. A, dorsal; B, P5. Male. C, dorsal; D, P5. Figures from Mori
(1937).
171
Family Aetideidae
Cephalosome and first pedigerous segment fused. Last metasome segment often prolonged in
points. Rostrum usually strong, simple or bifurcated, rarely absent. Female urosome 4segmented, male 5-segmented. P5 generally absent in the female, of simple structure in the male.
Genus Aetideus
In the female the cephalosome and first pedigerous segment are fused; sometimes with a crest on
the dorsal cephalosome; rostrum bifurcated, strong, widely spaced. Posterior metasome segments
usually symmetrical, prolonged into points. P5 missing, P4 coxa with a few small spinules near
base of the internal setae. In the male the cephalosome and first pedigerous segment are also
fused. No rostrum or crest; posterior metasome segment with lateral points. P5 with only one
uniramous leg, the left, 5-segmented, simple and elongated.
Aetideus bradyi A. Scott, 1909
Length: females: 1.38-1.80 mm; males: ? mm.
Female: Rostrum robust, bifurcated and protruding ventrally. Crest present on dorsal anterior
cephalosome, visible dorsally as a line. Wing- like extensions, pointed at ends, on the posterior
metasome segment, usually reaching the ends of the second urosome segment. A1 almost as long
as the prosome. P5 absent.
Male: Undescribed.
B
A
Aetideus bradyi: Female. A, dorsal; B, lateral; C, urosome lateral; D. rostruml; E, urosome

ventral; F, head lateral. Figures from Markhaseva (1996).
172
Aetideus giesbrechti Cleve, 1904

Female: Rostrum very robust and protruding ventrally, with 2 thickenings between the points.
Crest not very pronounced, but visible in dorsal and ventral view. Wing- like extensions, pointed
at ends, on the posterior metasome segment, usually reaching the ends of the second urosome
segment. A1 reaches to around the third urosome segment. P5 absent
Male: Corners of the last metasome segment much less pointed than in the female, reaching the
posterior edge of the genital segment. A1 reaches the posterior border of the third abdominal
segment. P5 simple, uniramous with only the left leg present, straight and 5-jointed
B
A
D
C
G
H
F
Aetideus giesbrechti: Female. A, dorsal; B, lateral; C, rostrum; D. urosome ventral; E, urosome

lateral. Male. F, dorsal; G, lateral; H, urosome; I, P5. Figures from Markhaseva (1996).
173
Genus Euchirella
Robust copepods. In the female the cephalosome and the first pedigerous segment are
completely or partially fused. A crest on the dorsal cephalosome may be present. Posterior
metasome segments rounded or angular, usually symmetrical. Furca nearly as long as wide. A1
usually longer than the prosome. P5 absent; depending on species, coxa with variable (1-13)
number of spines or spinules at the base of the internal seta, on the posterior surface. In the male
a crest on the dorsal cephalosome may be present. The cephalosome and the first pedigerous
segment are fused. Posterior metasome segments usually rounded. A1 longer than prosome,
reaching the end of urosome segment 3, or to the end of the furca. P5 large and heavily modified.
Euchirella amoena Giesbrecht, 1888
Female: Prosome 4-5 times longer than urosome. Posterior metasome segments in dorsal view
nearly rectangular. Genital segment symmetrical; A1 reaching half way down the urosome. P5
missing, P4 coxa with around 5 tiny teeth at the base of a plumose seta.
Male: Similar in appearance to female. P5 with uniramous left leg and large biramous right leg.
Right P5 endopodite longer than the exopodite. The first exopodite segment of right P5 with a
large spine- like projection in the mid internal margin, the second exopodite segment is about one
third the length of the first segment
Recorded: CPR.
C
B
A
E
D
G
Euchirella amoena: Female. A, dorsal; B, head lateral; C, urosome ventral; D. urosome lateral;
E, basis of P4. Male. F, dorsal; G, P5. Figures A, E from Sars (1925; as E. brevis); B-D, F, G
from Markhaseva (1996)
174
Euchirella pulchra (Lubbock, 1856)

Female: No crest on anterior cephalosome, but in lateral view this area is rounded or slightly
triangular. Rostrum well developed. Genital segment asymmetrical with a large ear-like
protrusion on the left and a small depression on the right. Anterior metasome segment rounded
and symmetrical. A1 reaches almost to end of genital segment. P5 absent, P4 coxa with 2 spines.
Male: Low crest on anterior cephalosome. Rostrum well developed. P5 left without endopodite,
the first segment of the exopodite not reaching the distal end of the right basis. The P5 right
endopodite with 4 projections along the external margin; the first segment of the exopodite with
3 projections on inner margin
Recorded: Zuza.
B
A
E
H
D
I
Euchirella pulchra: Female. A, head lateral; B, urosome ventral; C, urosome lateral right; D.
urosome lateral left; E, coxa P4. Male. F, head dorsal; G, head lateral; H, P5; I, detail of terminal
segment of left P5 exopod. Figures from Markhaseva (1996).
175
Genus Gaetanus
In the female one pointed rostrum present, which is not large. Frontal spine sometimes present
on anterior cephalosome. Cephalosome and first pedigerous segment fused. Posterior metasome
segment symmetrical, rounded but with spines of different shapes and sized. Genital segment
symmetrical. P5 absent; P4 coxa with group of 11-35 spines in a clump. In males a rostrum is
present, but if they have a frontal spine it is smaller than in the female. Spines on the posterior
metasome segment reduced compared to female. P5 asymmetrical, biramous but endopodites
very reduced.
Gaetanus minor Farran, 1905
Length: females: 1.70-2.40 mm; males:? mm.
Female: Frontal spine present on anterior cephalosome, curving towards the rostrum. Spines on
posterior metasome segment reaching almost to the distal edge of genital segment. Spines
straight and situated towards the dorsal side of the segment. A1 reaches almost to furca. P5
missing; P4 coxa with group of 12-15 small spines beside internal seta.
Male: Undescribed
C
A
Gaetanus minor: Female. A, dorsal; B, lateral; C, urosome dorsal ; D. urosome lateral left; E,
urosome lateral right; F, P4. Figures from Markhaseva (1996).
176
Genus Undeuchaeta
In female crest sometimes present, 1-pointed rostrum present. Cephalosome and first pedigerous
segment indistinctly separated or fused. Last metasome segment produced into obtuse triangular,
or triangular lobes, one of which may be rounded. Genital segment asymmetrical, with small
projection or spine on the right. A1 nearly as long as the prosome, or reaching to around the third
urosome segment. P5 missing; P4 coxa without large spines, rarely with small spinules. In male
the cephalosome and first pedigerous segment fused. Rostrum 1-pointed. Crest on anterior
cephalosome sometimes present. Corners of last metasome segment slightly asymmetrical and
rounded. A1 reaches the end of the prosome, or to around segment 3 of the urosome. P5 large
and robust, biramous, sometimes longer than urosome. The left P5 third exopodite segment
extended into a sharp projection.
Undeuchaeta intermedia A. Scott, 1909
Female: No crest; corners of last metasome segment with triangular lobes, slightly asymmetrical,
sometimes the right corner is rounded in dorsal and lateral view. In dorsal view, spine on right
side of genital segment, in ventral view 2 spines on either side of same segment. In lateral view
large spine like protrusion from genital segment. A1 almost as long as prosome. P5 missing.
Male: P5 biramous, right exopodite segment 1 with sharp spine on the first third of the segment,
with small projection on opposite side; internal segment covered with almost parallel wavy lines.
Left endopodite reaching the distal end of the first exopod segment, second exopodite segment
with spine-like projection on external distal border, third exopodite segment proximal angle
extending into a spine.
Recorded: Zuza.
C
Undeuchaeta intermedia: Female. A, dorsal; B, head lateral; C, urosome lateral left; D. urosome
lateral right; E, urosome ventral; F, genital segment ventral. Figures from Markhaseva (1996).
177
Undeuchaea plumosa (Lubbock, 1856)

Length: 3.00-4.20 females: mm; males: 2.85-3.90 mm.
Female: Posterior cephalosome without dorsal crest, rostrum single pointed, directed ventrally.
Last metasome segment slightly asymmetrical in dorsal view, in lateral view the right corner
rounded, the left more pointed, although this feature is variable. In dorsal view, spine on right
side of genital segment. A1 equal in length to the prosome, or slightly longer. P5 absent, P4 coxa
without internal spines at base of seta.
Male: No crest on cephalosome. A1 around the same length as prosome. Posterior metasome
segment corners rounded. P5 asymmetrical, biramous. Right P5 exopodite and endopodite both
3-segmented. Left P5 exopodite 3-segmented, the apical portion pointed; endopodite 1segmented, reaching around the distal part of the first exopodite segment.
A
E
B
F
Undeuchaea plumosa: Female. A, dorsal; B, head lateral; C, urosome lateral right; D, P4. Male.
E, dorsal; F, P5. Figures A-C, E, F from Sars (1925); D from Markhaseva (1996).
178
Family Arietellidae
Genus Metacalanus
Metacalanus aurivilli Cleve, 1901
Female: Small copepod, in lateral view rounded with high prosome. Posterior of last metasome
segment rounded in dorsal view. A1 short, only reaching to around the third metasome segment,
proximal segments of A1 short, distal segments much longer. Urosome 4-segmented, the second
and anal segment much shorter than the other segments. P5 slightly asymmetrical, uniramous, 2segmented. Distal segment with an internal long plumose seta, a shorter terminal spine and an
outer twisted spine.
Male: Urosome 5-segmented. A1 short, only reaching to around the third metasome segment,
proximal segments short, distal segments much longer. Left A1 geniculate. P5 almost
symmetrical, 5-segmented, uniramous, distal segment in the form of a thick spine. Segments 3
and 4 with thick spines on outer distal ends.
B
A
E
C
Metacalanus aurivilli: Female. A, dorsal; B, lateral; C, P5. Male. D, urosome; E, P5. Figures
from Greenwood (1978).
179
Family Clausocalanidae
Genus Clausocalanus
Small to medium sized copepods which may initially be confused with Paracalanus, but differ in
that the females have a longer urosome in relation to the length of the body and the P5 is more
robust and simple, bifurcated distally with only tiny spikes on the end rather than a long spine.
P5 uniramous on both sides, symmetrical, 3-segmented. The anterior cephalosome and posterior
metasome segments are rounded. Cephalosome fused to first pedigerous segment. Rostrum
bifurcated. Urosome 4-segmented. Male rostrum reduced to a knob. P5 uniramous, legs of
unequal length, longer leg nearly always on the left, 5-segmented with segment 5 short and
attached sub-apically to the previous segment; shorter leg, 3-segmented, less than half the length
of segment 1 of the other leg.
180
Clausocalanus arcuicornis (Dana, 1849)

Female: Rostrum in lateral view short, bifurcated, thick at its base, usually straight or slightly
curved and directed ventrally. Genital segment in lateral view straight or, more often slightly
concave in the region of the seminal receptacle; 1.5 times as long as urosome segment 3. The
third segment of the simple P5 as long as the two preceding segments together. Terminal points
of segment 3 of leg 5 bifurcated, sometimes with tiny spinules on inner and outer margins.
Male: Second urosome segment as long as the two following segments together. Rostrum in
lateral view knoblike and protruding ventrally. P5 uniramous and asymmetrical, right leg short,
usually 3-segmented; left leg robust, longer than urosome, armed distally with long slender
straight setae.
B
A
F
I
H
Clausocalanus arcuicornis: Female. A, dorsal; B, lateral; C, rostrum lateral and dorsal; D.

basipod 2 of P2 (upper), basipod 2 of P3 (lower); E, P5; F genital segment lateral. Male. G,
dorsal; H, lateral; I, urosome lateral; J, urosome dorsal; K, P5; L, basipod 2 of P2 (upper),
basipod 2 of P3 (lower). Figures from Frost and Fleminger (1968).
181
Clausocalanus furcatus (Brady, 1883)

Female: Rostrum in lateral view usually thick, short, and slightly curved, bifurcated at tip.
Genital segment profile in lateral view rather uneve n, generally shorter in length than the second
or third segments. The 2 distal segments of the P5 are of similar length, the coxa very short;
terminal points of bifurcated segment 3 never with tiny spinules.
Male: Rostrum in lateral view not well developed and not protruding ventrally. Longer P5
usually on the right side, occasionally on the left, longer than the urosome; shorter leg usually 2segmented, tiny distal segment spine- like. Recorded: Madagascar.
E
F
G
H
Clausocalanus furcatus: Female. A, lateral; B, genital segment lateral; C, P5. Male. D, lateral; E,
dorsal; F, metasome dorsal; G, urosome; H, P5. Figures A-C from Bradford-Grieve (1994); D-H
from Frost and Fleminger (1968).
182
Clausocalanus paululus Farran, 1926

Female: Rostrum slender and curved posteriorly. In lateral view genital segment with a
depression in mid segment, longer than either of the following 2 segments. Points on distal P5
not divergent and may have tiny spines on inner and outer margins.
Male: Short squat body. In lateral view rostrum knob-like and projecting ventrally. P5 left longer
than urosome; right P5 2-segmented with tiny spine- like distal segment.
B
A
H
D
Clausocalanus paululus: Female. A, lateral; B, rostrum lateral; C, genital segment lateral; D, P5.
Male. E, lateral; F, head lateral; G, urosome dorsal; H, P5. Figures from Frost and Fleminger
(1968).
183
Family Euchaetidae
Most of this group are large copepods. Both the male and female have very large maxillipedes,
associated with their carnivorous diet, and also long urosomes. The innermost (appendicular)
seta on each furca is very well developed and usually much longer than the other caudal setae.
The anterior of the cephalosome in both sexes, in most species, comes to a point when viewed
dorsally. In lateral view a single strong rostrum is obvious and just behind this is a frontal
eminence that varies from a slightly raised ledge with a small sensory bristle to a strong curved
point. The A1 have characteristic very long setae. The female genital segment is very prominent
when viewed laterally and different species have characteristic shapes that are used in
identification. The female often has a single egg sac still attached. The P5 is absent in females.
The male P5 is very large and characteristic, with rudimentary endopodites; left leg with a
complex distal joint.
Genus Euchaeta
The innermost seta on each furca is greatly developed, straight and much thicker and longer than
the other caudal setae. The P5 terminal exopodite segments in both right and left legs of the male
taper into long spines.
Of the three Euchaeta species so far recorded for the area, E. indica and E. rimana belong to the
marina species group while E. longicornis belongs to the concinna species group. These two
groups can be distinguished by the following features.
In both groups the basis of the female maxilliped has both long and short spines along the
proximal half of the medial margin (Fig. A). The serrated lamella of the male left P5 exopodite
reaches or extends beyond a tuft of stiff hairs (Fig. C). In the marina group the two endopodal
setae of the female maxillule 2 are armed with both long and short spines (Fig. B), while in the
concinna group only one seta is armed this way (Fig. D). Males in the marina group have on the
exopod of the left P5, 3 lobes, one which is next to the tuft of hairs (Fig C), not present in the
concinna group (Fig E).
B
D
C
Identification features of limbs mentioned in the text. From Park (1994).
184
Euchaeta indica Wolfenden, 1905

Belongs to the marina species group.
Female: Frontal eminence on forehead well developed. In lateral view, the genital segment is
about twice as long as the broadest part. Genital segment is as long as the combined length of the
next three segments and is irregular in outline when viewed from the dorsal and ventral sides.
Urosome segment 2 is about half the length of the genital segment and is equal to the combined
length of segments 3 and 4. The appendicular seta is much longer than any of the others. The last
metasome segment and the urosome segments are densely covered with short fine hair s. A1
extend to the end of urosome segment 2.
Male: P5 like E. marina in general appearance, distinguished by the structure of the serrated
lamella, which is bilobed distally, with double rows of teeth bordering the lobes.
Recorded: Rodrigues, Zuza, Aldabra, CPR.
A
D
G
Euchaeta indica: Female. A, dorsal; B, head lateral; C, urosome left lateral; D, urosome left
lateral; E, urosome right lateral; F, urosome ventral; G, urosome dorsal; H, maxilla, fifth lobe
and endopod omitted.. Male. I, lateral; J, head lateral; K, exopod of left P5. Figures A, I from
Mori (1937; as E. wolfendeni); B-H, J, K From Park (1994).
185
Euchaeta rimana Bradford, 1974

Also belongs to the marina species group and was originally recorded as E. marina which is now
recognised to be restricted to the Atlantic and Mediterranean.
Female: Can only be separated from E. marina by the structure of the genital segment. In both
dorsal and ventral view, the left side of the genital segment is smoothly curved, without any
prominent projection; the right side with a moderate projection. Posterior metasome segments
are slightly asymmetrical in dorsal view, slightly more extended on right.
Male: The serrated structure of the lamella of the left P5 exopod segment 2 is the only
morphological feature of the appendages found to be different between this species and E.
marina. The distal margin has an unserrated gap in E. marina but not in E. rimana. The thick
spine originating at the base of the lamella is less than half the length of the lamella.
Recorded: Rodrigues, Zuza, Madagascar, CPR.
G
H
Euchaeta rimana: Female. A, head lateral; B, urosome lateral left; C, urosome lateral le ft; D,
urosome left right; E, urosome dorsal; F, urosome ventral. Male. G, head lateral; H, exopod of
left P5; I, distal end of serrated lamella. Figures from Park (1994).
186
Euchaeta longicornis Giesbrecht, 1888

Belongs to the concinna species group
Female: High frontal eminence on forehead and long slender rostrum. In both lateral and dorsal
view, distal end of metasome on each side broadly rounded. Dorsal and ventrally, genital
segment strongly asymmetrical with a broad swelling on proximal half of left side and a large
winglike outgrowth on distal half of right side. Lateral side of winglike outgrowth bearing a
small process.
Male: First exopodite segment of right P5 with 2 patches of small teeth on inner margin. Fingerlike process on left leg exopodite, sharply pointed and far short of reaching the middle of the
serrated lamella. Lamella strongly curved towards exopodite segment 3 and with a conspicuous
notch on the distal margin. Teeth all small along margin.
A
I
F
H
M
N
P
L
Euchaeta longicornis: Female. A, dorsal, lateral; B, head lateral; C, urosome lateral left; D,
urosome dorsal; E, urosome lateral right; F, urosome ventral; G, urosome lateral. Male. H,
dorsal, lateral; I, head lateral; J, metasome lateral; K, right P5; L-O, exopod of left P5 different
angles; P, distal end of serrated lamella. Figures A, H from Grice (1962); B-G, I-P from Park
(1994).
187
Family Phaennidae
Genus Phaenna
Phaenna spinifera Claus, 1863
Female: Prosome is almost round in dorsal view; head and first pedigerous segment separate.
Rostrum bifurcated. Urosome 4-segmented, genital segment slightly longer than the combined
length of the following 3 segments. Anal segment short, furca as wide as long. A1 reaches
around the middle of the urosome. P5 missing.
Male: Prosome broad but more elongated than in the female. Urosome 5-segmented. P5
asymmetrical but legs of almost equal length, left leg slightly longer, uniramous. Left leg, 5segmented, right leg 4-segmented.
Recorded: Seychelles, Zuza.
B
C
A
E
F
Phaenna spinifera: Female. A, dorsal; B, lateral; C, P4. Male. D, dorsal; E, ventral; F, P5.
Figures A, C from Mori (1937); B, D-F from Bradford et al. (1983).
188
Family Scolecitrichidae
Genus Lophothrix
Lophothrix latipes (T. Scott, 1894)
Female: Head in lateral view angular, with small crest. Rostrum bifurcated with fine setae.
Cephalosome and first pedigerous segment fused. Corners of posterior metasome segment
produced into small points. A1 reaches to the genital segment. P5 uniramous, 3-segmented with
3 short spines on the distal segment, the outer the shortest.
Male: Similar to female but anterior cephalosome is squarer in dorsal view and the posterior
metasome segments are more rounded. A1 reaches to around the end of the metasome. P5
reaches the distal end of urosome segment 3. Right P5 exopodite 4-segmented, the last segment
flattened and triangular, endopodite 2-segmented, reaching exopodite segment 2. Left exopodite
3-segmented, last segment with 2 short, broad setae distally and several spines; endopodite 2
segmented, terminating in a flattened seta.
A
B
Lophothrix latipes: Female. A, lateral; B, rostrum; C, P5; D, P5 variant. Figures from Bradford et
al. (1983).
189
Genus Scolecithrix
Scolecithrix bradyi Giesbrecht, 1888
Female: Posterior metasome segments extended, appearing as points in dorsal view but rounded
in lateral view, the right side more prominent than the left, extending beyond the genital
segment. A1 does not reach the end of metasome. Genital segment asymmetrical, left side very
swollen in dorsal view, as long as the combined length of the following 3 segments. The anal
segment is as long as the preceding segment. Furca twice as long as wide. P5 very small and
simple, right leg longer than left, 1-segmented, each leg consisting of an elongated flattened
blade with 2 small terminal spines. This copepod is similar to S. danae, but smaller and has
longer extensions on the posterior metasome segment. S. danae does not have a P5.
Male: Shape similar to female. Anterior cephalosome rounded in lateral view, posterior
metasome segments not produced as in female. P5 left biramous, with 3-segmented exopodite
and 1-segmented endopodite as long as exopodite segment 1. Right leg uniramous, with 2segmented exopodite, distal end of segment 1 with short fingerlike process on inner margin,
terminal segment divided into 2 sub-equal branches. Similar to S. danae, but S. danae does not
have the bifurcated terminal segment on the right P5.
A
E
F
Scolecithrix bradyi: Female. A, dorsal; B, lateral; C, metasome and urosome; D, P5. Male. E,
lateral; F, P5. Figures A, C-F from Bradford et al. (1983); B from Mori (1937; as Scolecithricella
bradyi).
190
Scolecithrix danae (Lubbock, 1856)

Female: Prosome has a very characteristic shape, being very broad at the mid point. The anterior
metasome segment is produced into points which extend close to the urosome, reaching half way
down the genital segment. The A1 extends beyond the prosome. Urosome of 4 segments. The
genital segment has a downward pointing projection when viewed from the side. Setae on the
swimming legs and furca are typically very feathery and in fresh specimens tinged yellow. The
setae on the furca are of similar length. P5 absent.
Male: Similar to female but posterior metasome segment not produced. Urosome 5-segmented.
P5 asymmetrical, right foot uniramous with 2-segmented exopodite with a short terminal
segment; left foot biramous, the exopodite with 3 segments, the simple endopodite with 1
segment.
D
A
Scolecithrix danae: Female. A, dorsal; B, lateral. Male. C, dorsal; e, P5. Figures from Mori
(1937).
191
Genus Scaphocalanus
Scaphocalanus echinatus (Farran, 1905)
Female: Anterior cephalosome without median crest. Prosome slender and elongated in dorsal
view. Posterior metasome segments slightly extended but not pointed. A1 reach almost to the
end of the genital segment. Genital segment as long as wide. The second seta on the furca is
twice the length of the urosome. P5 symmetrical, uniramous, 2-segmented, second segment with
an outer marginal small spine, a terminal long spine and an inner marginal long spine which is
internally bordered with coarse teeth. Some specimens may have no outer edge spine, but an
additional small spine adjacent to the terminal spine; some specimens may have a rudimentary
endopodite and a shorter terminal spine (Bradford et al., 1983)
Male: Posterior metasome segments not extended. A1 reaches urosome segment 2. P5 reaches
middle of urosome segment 4. Right P5 exopodite 3-segmented, distal segment very small and
slender; endopodite reaches to exopodite segment 1. Left leg endopodite 1-segmented.
A
C
Scaphocalanus echinatus: Female. A, dorsal; B, lateral; C, P5. Male. D, urosome and P5. Figures
A, B from Mori (1937); C, D from Bradford et al. (1983).
192
Genus Scolecithricella
Cephalosome and first pedigerous segment fused. Rostrum bifurcated. No median crest on
cephalosome Female P5 reduced, 2-segmented, uniramous and flattened. Male P5 biramous on
both sides, with very short endopodites.
Scolecithricella minor (Brady, 1883)
Female: Anterior cephalosome rounded in lateral view, posterior metasome segment slightly
produced, rounded in dorsal view and slightly angular in lateral view. A1 reaches just beyond the
prosome. P5 symmetrical. Each foot 2-segmented, oval, twice as long as wide, with a slender
spine on the outer margin, a short spine on the apex and a long spine on the inner margin.
Male: posterior metasome segment rounded in dorsal and lateral view. P5 asymmetrical, longer
than the urosome; right foot uniramous, with a vestigial triangular endopodite; exopodite
segment 3 elongated. Left P5 biramous, endopodite 1-segmented and extends to about the middle
of the second segment of the exopodite. The terminal segment of the exopodite elongated and
sharply pointed.
C
D
A
Scolecithricella minor: Female. A, lateral; B, P5. Male. C, dorsal; D, lateral; E, P5. Figures from
Mori (1937).
193
Scolecithricella ovata (Farran, 1905)

Female: Posterior metasome segment quite rounded in dorsal view. A1 extends to around
urosome segment 3. P5 outer segment flattened and of irregular outline, sometimes fused to
common basal segment, stout spine on internal margin, sometimes a second sub-terminal spine.
Male: A1 reaches urosome segment 3. P5 uniramous on both sides and reaches to the anal
segment. Right P5 short, 3- segmented, the third segment overlapping the beginning of the
second segment of the left leg. Left leg long, 5-segmented
C
D
Scolecithricella ovata: Female. A, lateral; B, urosome lateral; C, P5; D, P5 variant. Male. E,

dorsal; F, P5. Figures A-D, F from Bradford et al. (1983); E from Tanaka (1962).
194
Order Cyclopoida
Cyclopoids include not only free- living species, but also commensal and parasitic species. Of
the free- living cyclopoids, genera such as Oithona can numerically dominate the samples.
Cyclopoids are generally less than 1 mm in length, they range in size from 0.3 to 1.9 mm.
Cyclopoids resemble calanoids in that there is a sharp division between the prosome and
urosome, however, the body articulates between thoracic segments 4 and 5 (rather than
thoracic segment 5 and abdominal segment 1 as in the calanoids). The 5th thoracic segment is
very narrow, it bears the simple P5 and is the first segment of the urosome. The A1 is
elongate, especially in females, although not usually as long as in the Calanoida,. The A2 is
uniramous. Females carry a pair of egg sacs attached to the lateral or subdorsal surface, but
never to the ventral surface as in the Harpacticoida. P1-4 are well developed, except for
parasitic species. P5 are small and similar in both sexes.
Family Oithonidae
The body is usually elo ngate, the prosome wider than the narrow urosome. The Family
includes two genera that are free-swimming in the plankton, Oithona and Paroithona. 25
species have been recorded in the Indian Ocean.
Genus Oithona
Identification of Oithona species is made difficult by the small size of the organisms and the
close relationship they bear to each other, especially so in the males. The males are readily
separated from the females as the A1 are usually shorter and both are geniculate. The males
are usually smaller than the females, and the urosome is 6-segmented in males and 5segmented in females.
195
Oithona attenuata Farran, 1913

Female: Prosome ovoid. Cephalsome truncate anteriorly. Rostrum blunt and not visible
dorsally. Urosome longer than prosome. Genital segment three times longer than wide, with a
prominent lateral processes approximately 1/3 down the genital segment. Anal segment 1.5
times longer than wide. Furca slender and long, four times longer than wide. A1 reaches the
end of the prosome.
Male: Cephalosome truncate anteriorly. Prosome just shorter than urosome. Furca longer than
anal segment.
Oithona attenuata: Female. A, dorsal; B, lateral view of rostrum; C, A2; D, A1; E, P5 and
genital segment; F, furca; G, P4. Male. H, P4; I, dorsal. Figures A-G from Nishida (1985), H
and I from Rosendorn (1927).
196
Oithona fallax Farran, 1913

Female: Cephalosome narrowed anteriorly, but not pointed. Rostrum not visible dorsally.
Anterior 1/3 of genital segment swollen laterally, over twice as long as wide. Anal segment
longer than wide posterior margin serrated. Furca more than twice as long as wide. A1
reaches the beginning of genital segment. Very similar to O. similis, however terminal spines
on the 3rd joint of the P2 exopod are shorter, and there is a small inner edge seta on the 1st
joints of P2-4.
Male: Anterior of cephalosome rounded. Prosome longer than urosome. Genital segment
longer than anal segment and furca together. Furca longer than anal segment. A1 geniculate
once.
Recorded: Western Indian Ocean.
Oithona fallax: Female. A, dorsal; B, lateral view of rostrum; C, A2; D, A1; E, P5 and genital
segment; F, furca; G, P4. Figures from Nishida (1985).
Oithona fallax: Male. A, dorsal; B, urosome; C, P4; D, A1. Figures from Rosendorn (1927).
197
Oithona nana Giesbrecht, 1892

Female : Cephalosome truncate anteriorly. Rostrum blunt and not visible dorsally. Prosome
short and stout, approximately the same length as the urosome. Genital segment
approximately twice as long as wide and very rounded in the anterior quarter. Anal segment
as long as wide. Furca twice as long as wide. A1 as long as the prosome.
Male: Prosome shorter and stouter than female. Genital segment longer than wide. A1
geniculate. Anal segment as long as wide. Furca twice as long as wide.
Oithona nana: Female. A, dorsal; B, lateral view of rostrum; C, A2; D, A1; E, P5 and genital
segment; F, furca; G, P3. Male. H, dorsal. Figures A-G from Nishida (1985), H from Mori
(1937).
198
Oithona oculata Farran, 1913

Female: Pair of dark eyes visible through the anterior section of the cephalosome. Anterior
section of cephalosome truncate. Rostrum present and blunt, not visible dorsally. Prosome
stocky, approximately 1.5 times longer than the urosome. Genital segment 1.5 times longer
than wide, the first quarter swollen laterally. Anal segment just shorter than wide, posterior
ventral margin serrated. A1 shorter than prosome.
Male: Cephalosome similar to female. Prosome oblong in dorsal view, more than twice as
long as wide. Genital segment just longer than wide. Anal segment shorter than wide,
posterior ventral margin serrated. Furca twice as long as wide. P1-4, the terminal spine on the
exopods is very long.
Oithona oculata: Female. A, dorsal; B, anal segment and furcal; C, A2; D, A1; E, genital
segment and P5; F, P4. Figures from Nishida (1985).
Oithona oculata: Male. A, dorsal; B, lateral view of rostrum; C, cephalosome dorsal view; D,
A2; E, A1; F, furca; G, lateral view of P5 and genital segment; H, P4. Figures A, B, & D-H
from Nishida (1985), C from Rosendorn (1927).
199
Oithona plumifera Baird, 1843

Female: Anterior section of cephalosome narrowed into the pointed rostrum. Rostrum visible
in dorsal view. Genital segment twice as long as wide. Genital segment swollen laterally on
the anterior third, with a characteristic tuft of hairs on the ventral side. Furca shorter than anal
segment and three times longer than wide. A1 long, reaching the anal segment. Segments of
A1 fringed with small spinules. Plumose setae arise from swimming legs of some specimens,
not always present.
Male: Anterior of cephalosome very different to female, it is not pointed and the rostrum is
blunt. A1 is twice geniculate, with a sheath just beyond the proximal elbow and a semicircular
process on the 1st segment beyond the distal elbow. Genital segment large, about as wide as it
is long. The anal segment is similar in length to the two preceding segments.
Oithona plumifera: Female. A, dorsal; B, dorsal view with appendages; C, A2; D, A1; E.
furca; F, P5 and genital segment; G, P4; H, lateral view of rostrum. Male. I, dorsal; J,
endopod of P4. Figures A and C-H from Nishida (1985), B, I and J from Giesbrecht (1892).
200
Oithona pseudofrigida Rosendorn, 1917

Length: females: 1.22-1.25 mm.
Female: Anterior section of cephalosome narrowed into a pointed rostrum. Rostrum visible in
dorsal view. Anterior third of genital segment swollen laterally, genital segment without tuft
of hairs ventrally. Genital segment 2.5 times longer than wide. Anal segment longer than
wide. Anterior ventral section of anal segment with long setae. Furca over 3 times longer than
wide. Segments of A1 fringed with small spinules.
Oithona pseudofrigida: Female. A, dorsal; B, lateral view of rostrum; C, A2; D, A1; E, furca,
ventral view; F, furca, dorsal view; G, lateral view of P5 and genital segment; H, P4. Figures
from Nishida (1985).
201
Oithona robusta Giesbrecht, 1891

Female: Cephalosome pointed anteriorly. Rostrum just visible dorsally. Anterior 1/3 of
genital segment slightly swollen laterally. Genital segmement twice as long as wide. Anal
segment 1.5 times longer than wide, only the lateral margins of anal segment lined with
spinules. Furca three times longer than wide. A1 shorter than prosome length.
Male: Cephalosome truncated anterioirly. Prosome longer than urosome. Genital segment
relatively short. Furca longer than anal segment.
Oithona robusta: Female. A, dorsal; B, lateral view of rostrum; C, A2; D, A1; E, lateral view
of P5 and genital segment; F, furca; G, P4. Male. H, dorsal; I, P4. Figures A-G from Nishida
(1985), H and I from Rosendorn (1927).
202
Oithona setigera (Dana, 1849)

Female: Allied to O. plumifera, but the body is larger and A2 is shorter. Cephalosome
narrows anteriorly to sharp pointed rostrum that is visible dorsally. Anterior third of genital
segment swollen laterally. Genital segment twice as long as wide. Anal segment
approximately twice as long as wide. posterior margin of anal segment lined with spinules,
Anterioir ventral section of anal segment with long setae. Furca approximately three times
longer than wide. A1 longer than prosome.
Male: Rostrum not pointed. Prosome longer than urosome. Furca just longer than anal
segment.
Oithona setigera: Female. A, dorsal; B, lateral view of rostrum; C, A2; D, A1; E, ventral view
of furca; F dorsal view of furca; G, P4. Figures from Nishida (1985).
Oithona setigera: Male. A, dorsal; B, urosome; C, lateral view of anterior cephalosome; D,

P1; E, P4. Figures from Rosendorn (1927).
203
Oithona similis Claus, 1866

Female: Cephalosome narrowed anterioirly but not to a point. Rostrum present but not visible
dorsally. Urosome approximately the length of the prosome. Anterior third of genital segment
swollen laterally. Genital segment more than twice as long as wide. Anal segment just longer
than width. Furca shorter than anal segment and divergent, more than twice as long as wide.
Second furcal seta is longer than the urosome. Segments of A1 fringed with small spinules.
A1 doesnt reach the genital segment. Ovisacs when present usually with a single row of very
large eggs.
Male: Rostrum short and blunt. A1 twice geniculate, with a sheath just beyond the proximal
elbow and a semicircular process on the 1st segment beyond the distal elbow, as in O.
plumifera. Furca about as long as wide.
Oithona similis: Female. A, dorsal; B, lateral view of rostrum; C, A2; D, A1; E, P5; F, furca;
G, P5 and genital segment, lateral view; H, P5 and genital segment, dorsal view. Figures from
Nishida (1985).
Oithona similis: Male. A, dorsal; B, A1. Figures from Wilson (1932).
204
Oithona simplex Farran, 1913

Length: females: 0.36-0.42 mm; males: 0.39-0.41
Female: Prosome oval. Cephalosome narrows anteriorly, narrow region truncate at tip.
Rostrum blunt and not visible dorsally. Anterior third of genital segment swollen laterally.
Genital segment 1.5 times longer than wide. Anal segment short, half as long as wide,
posterior margin lined with spinules. Furca more than 1.5 times longer than wide. Furca
longer than the anal segment. A1 short, reaches to the beginning of the 2nd metasome
segment. The segments of A1 without a fringe of spinules.
Male: Prosome oval. Rostrum blunt and not visible dorsally. Anterior of cephalosome
truncate. Genital segment just longer than wide. Anal segment short, twice as wide as long,
posterior margin lined with spinules. Furca 1.5 times longer than wide.
Oithona simplex: Female. A, dorsal; B, lateral view of rostrum; C, A2; D, A1; E, lateral view
of P5 and genital segment; F, dorsal view of P5 and genital segment; G, P6; H, furca; I, P4.
Figures from Nishida (1985).
Oithona simplex: Male. A, dorsal; B, lateral view of rostrum; C, A2; D, A1; E, furca; G,
dorsal view of P5 and genital segment; G, lateral view of P5 and genital segment; H, P4.
Figures from Nishida (1985).
205
Oithona tenuis Rosendorn, 1917

Female: Body slender and elongate. Cephalosome narrowed anteriorly into pointed rostrum.
Rostrum visible dorsally. Genital segment only slightly swollen laterally in anterior third
section. Genital segment approximately three times longer than wide. Anal segment
approximately 1.5 times longer than wide. Posterior margin of anal segment lined with
spinules. Furca approximately 2.5 times longer than wide.
Oithona tenuis: Female. A, dorsal; B, lateral view of rostrum; C, A2; D, A1; E, lateral view of
P5 and genital segment; F, furca; G, P4. Figures from Nishida (1985).
206
Oithona vivida Farran, 1813

Female: Cephalosome very pointed anteriorly, nearly triangular at the anterior- most section.
Prosome approximately the same length as urosome. Anterior third of genital segment
swollen laterally. Genital segment more than twice as long as wide. Anal segment more than
twice as long as wide. Postero-ventral margin of anal segment lined with spicules. Furca
approximately three times longer than wide. A1 longer than prosome. P6 with a very long seta
distinctive to O. vivida.
Oithona vivida: Female. A, dorsal; B, lateral view of rostrum; C, A2; D, A1; E, lateral view of
P5 and genital segment; F, furca; G, lateral view of anterior prosome; H, P5. Figures from
Nishida (1985).
207
Order Harpacticoida
Although a large group (approximately 3372), the Harpacticoida are numerically one of the
least common sampled. Only 17 species of harpacticoids can be considered truly pelagic, most
are either benthic or epibenthic. Benthic and epibenthic harpacticoids are a specialist group and
are not included in this guide, although at times they are caught in plankton samples. Female
harpacticoid copepods typ ically carry single egg sacs on the ventral surface, however, female
Miraciidae carry paired egg sacs. In general there is no demarcation between the prosome and
the urosome. The A1 is short and, in males both are geniculate. Males and females have 6 pairs
of swimming legs, the P6 is usually rudimentary in females. The familes with pelagic species
are Miraciidae, Ectinosomatidae, Clytemnestridae and Euterpinidae.
Family Miraciidae
All Miraciidae are planktonic and essentially tropical and subtropical. The family
characteristically possesses a pair of cuticular lenses (absent in Macrosetella) on the front of the
head. The prosome comprises the cephalosome and three pedigerous segments. The urosome is
5-segmented in females and 6-segmented in males. The furca are usually parallel, they are as
long or longer than the last 2 urosome segments combined. Each furca is furnished with 7 setae,
setae I-III are closely set together and setae IV-V are spinulose. P5 are fused medially in the
male but free in the female. Male P6 are symmetrical with 1-3 setae, female P6 are rudimentary.
Females carry paired egg sacs, the eggs are typically large and low in number (4 in O. gracilis
and D. minor, 4-6 in M. efferata and 2-11 in M. gracilis, the number varying with size of the
female). The family comprises four genera Miracia, Distioculus, Oculosetella and
Macrosetella.
208
Miracia efferata Dana, 1849

Female: The front of the head is rounded. The large pair of cuticular lenses touch middorsally. the cephalosome is wider than the following segment and there are distinct
constrictions between each of the prosome and urosome segments. Rostrum is very small. A1
is 8-segmented. Furca 3 times longer than wide, the outer margin is stepped 2/3 down the
furcal length. Seta V is shorter than the urosome. P5 is large and P6 is rudimentary.
Male: Body similar to female. A1 is 10-segmented with the geniculation between segments 7
and 8. P5 fused medially. P6 symmetrical.
A
Miracia efferata: Female. A, dorsal; B, lateral. Male. C, lateral. Figures A-C reproduced from
Taxonomy, biology and phylogeny of Miraciidae (Copepoda: Harpacticoida) by Huys &
Bttger-Schnack from Sarsia, www.tandf.no/sarsia, 1994, volume 79, pages 212 and 218, by
permission of Taylor and Francis AS.
209
Oculosetella gracilis (Dana, 1852)

Length: females: 1.20-1.30 mm; males: 0.80 mm
Female: Body slender and elongate, without a distinct boundary between the prosome and
urosome. Cephalosome is only slightly wider than first prosome segment. The front of the
cephalosome bears a pair of large cuticular lenses that touch in the dorsal midline. The rostrum
is conspicuous and distinctively elongate. A1 is 7-segmented. No distinct constrictions
between each prosome and urosome segments. Row of spinnules on the posterior margin of
the ventral surface of the last four urosome segments. Furca about 3 times longer than wide,
the outer margin is almost straight (not stepped as in Miracia). All the setae are located in the
posterior 1/6 of the furca, seta V is the longest and about 75% of total body length. P5 large.
P6 vestigial. Females carry paired egg sacs, each with 4 large eggs.
Male: Body shape similar to female. A1 is 10-segmented with the geniculation between
segments 7 and 8. P5 is fused medially and P6 is symmetrical.
Recorded: Seychelles, Rodrigues.
A
B
Oculosetella gracilis: Female. A, dorsal; B, lateral. Male. C, lateral. Figures A-C reproduced
from Taxonomy, biology and phylogeny of Miraciidae (Copepoda: Harpacticoida) by Huys
& Bttger-Schnack from Sarsia, www.tandf.no/sarsia, 1994, volume 79, pages 222 and 227,
by permission of Taylor and Francis AS.
210
Macrosetella gracilis (Dana, 1848)

Female: Body is slender, elongate and without paired cuticular lenses. Cephalosome relatively
large, about 1/3 of body length (excluding furca), and as wide as the following segment.
Prosome and urosome without distinct constrictions between segments. Ventral margin of
penultimate segment, and ventral and lateral margins of anal segment with spinules. Anal
segment narrow. Furca about 11 times longer than wide, narrowest halfway its length. Setae V
very long, distinctly longe r than total body length. The rostrum is relatively large and
elongate. A1 is 8-segmented. P5 is large, and P6 is rudimentary.
Male: Body shape similar to female. A1 is 10-segmented with the geniculation between
segments 7 and 8. P6 are weakly developed and symmetrical. Post-genital segments with rows
of spinules at the ventral posterior margin.
Recorded: Seychelles, Rodrigues, Zuza, Aldabra, Mauritius, Madagascar
B
A
Macrosetella gracilis: Female. A, dorsal; B, lateral. Male. C, lateral. Figures A-C reproduced
211
Distioculus minor (T. Scott, 1894)

Female: Cephalosome slightly narrower than the following prosome segment. The pair of large
cuticular lenses (often deep red in colour) do not touch mid-dorsally. The rostrum is minute.
A1 is 8-segmented. Prosome and urosome without distinct constrictions between segments.
Furca about 3 times longer than wide and the outer margin is stepped at about halfway down.
Setae V is the longest, but shorter than the urosome. P6 is rudimentary.
Male: Body similar to the female. A1 is 10-segmented with the geniculation between segments
7 and 8. P6 is symmetrical. Post- genital segments with rows of spinules at the ventral posterior
margin. Recorded: Rodrigues, Zuza.
A
Distioculus minor: Female. A, dorsal; B, lateral. Male. C, lateral. Figures A-C reproduced
212
Family Ectinosomatidae
The only truly pelagic genus in the family Ectinosomatidae is Microsetella.
Genus Microsetella
The body is slender and laterally compressed. The urosome is as wide as the metasome. The
urosome is 4-segmented in the female and 6-segmented in the male. The A1 is slender,
elongate, 5-segmented, and, in males is geniculate. The female P5 is 2-segmented and
symmetrical, the male P5 is rudimentary and symmetrical. The two species of Microsetella
are radily separated by the relative lengt hs of their furcal setae. The furcal setae are shorter
than the body in Microsetella norvegica and nearly twice as long as the body in Microsetella
rosea.
Microsetella norvegica (Boeck, 1864)
Length: females: 0.35-0.53 mm; males 0.33-0.42 mm.
Female: Body slender and laterally compressed. Rostrum short and turned downwards.
Transverse rows of minute spicules on the urosome segments. Furca about as long as wide,
and divergent. The longest furcal seta is nearly as long as the body. The second furcal seta is
about the length of the body. P5 with the two inner setae of very different lengths, the
outermost seta is twice as long as the neighbouring seta.
Male: Smaller than the female, the body is similar in shape. The furca are a little wider than
long. A1 is geniculate.
A
B
C
D
Microsetella norvegica: Female. A, dorsal; B, lateral; C, P5. Male. D, P5. Figures A from
Wilson (1932), B from Sars (1904), C & D from Giesbrecht (1892).
213
Microsetella rosea (Dana, 1848)

Length: females 0.64-0.85 mm.
Female: Body sometimes has a rosy or reddish tinge. Body similar in shape to Microsetella
norvegica, but almost twice the length. The 2nd, 3rd and 4th metasome segments, and the three
urosome segments with transverse rows of spinules near the anterior margins. The longest seta
of the furca is nearly twice as long as the body. P5 with the two inner setae approximately
equal in length.
B
A
Microsetella rosea: Female. A, dorsal; B, lateral; C, P5. Male. D, P5. Figure A from Wilson
(1932), B & C from Giesbrecht (1892), D from Johnson (1942).
214
Family Clytemnestridae
Body considerably flattened. Cephalosome and first three metasome segments with prominent
angular projections at the posterior corners. Prosome comprises cephalosome and three free
pedigerous segments. Urosome 5-segmented in female, 6-segmented in male. Anal segment as
long as the penultimate segment. Furca short and wide. A1 7- or 8-segmented, geniculate both
sides between the last two segments in males. P5 rudimentary and symmetrical, 2-segmented,
segments narrow and elongate. Females carry single egg sacs.
Clytemnestra scutellata Dana, 1848
Female: Body shape as for the family. A1 is 8-segmented; 4th , 5th and 8th segments with long
sensory hairs. The last segment is twice as long as the preceding one. The furca is
approximately twice as long as it is wide, and the 2 longest setae are plumose.
Male: Body similar to female. Both A1 are geniculate. 6th segment of A1 is short and has a
spine. The longest furcal setae are longer than those of the female.
Clytemnestra scutellata: Female. A, dorsal; B, lateral; C, furca. Male. D, furca; E, P5. Figures
A-E from Giesbrecht (1892).
215
Clytemnestra rostrata (Brady, 1883)

Female: Similar body shape to Clytemnestra scutellata, but generally smaller in size. A1 is 7segmented, the last segment is nearly five times as long as the preceding one. The furca is
about as long as wide, none of the furcal setae are plumose.
Male: Body similar to female, but is usually smaller. A1 geniculate between the last two
segments, both elongate. P5 resembles that of the female, but is a little shorter and narrower.
Clytemnestra rostrata: Female. A, furca, dorsal; B, P5. Male. C, A1. Figures A-C from
Giesbrecht (1892).
216
Family Euterpinidae
Prosome comprises cephalosome and three free pedigerous segments. Urosome 5-segmented
in females and 6-segmented in males. Females carry single egg sacs. Only one species within
the family, Euterpina acutifrons.
Euterpina acutifrons (Dana, 1848)
Female: The rostrum is stout. The anterior of cephalosome is sharply pointed. Body widens to
the posterior of the cephalosome then narrows posteriorly. A1 is 7-segmented. P5 is
symmetrical and 1-segmented. Body articulation is distinct.
Male: Body similar in shape to female. Both sides of A1 are geniculate; the 4th and 5th
segments are fused and much thickened; 6th and 7th segments are fused and make up the last
segment. P5 is symmetrical and rudimentary.
Recorded: Rodrigues, Seyc helles, Zuza, Aldabra, Madagascar.
C
D
Euterpina acutifrons: Female. A, dorsal; B, P5. Male. C, lateral; D, Posterior prosome and
urosome segments. Figures A-D from Giesbrecht (1892).
217
Order Poicilostomatatoida
Family Lubbockiidae
The genus Lubbockia has recently been reassigned from the family Oncaeidae to the family
Lubbockiidae (Huys & Bttger-Schnack, 1997). Lubbockiids are small oceanic copepods. The
prosome comprises the cephalosome and four free pedigerous segments. There is a distinct
boundary between the prosome and urosome. The urosome is 5-segmented in females and 6segmented in males. The genital apertures (on the genital segment) are paired and situated
dorsolaterally or laterally in females and ventrally in males. A2 is 3 or 4-segmented. The
maxillipeds end in a hook or claw and are 4-segmented in females and 5-segmented in males.
Females carry paired egg sacs.
Genus Lubbockia
Lubbockia squillimana Claus, 1863
Female: The body is slender and elongate. The anterior of the cephalosome is pointed. The
greatest width is at the posterior margin of the cephalosome; the remaining prosome segments
taper posteriorly. The prosome is 5-segmented and is longer than the urosome. The last prosome
segment is not pronounced at the posterior corners. The genital segment is longer than the
segment following. The furca is slightly longer than the anal segment, and is nearly 5 times as
long as wide. A1 is 5-segmented. The maxilliped is characteristic, the 1st segments has no
spines, the 2nd segment has 5 large denticles on the inner surface of the distal two thirds, the 3rd
segment has no spines, the 4th has smooth margins and ends in a claw.
Male: Similar body shape to the female, but larger in size. Prosome is longer than the urosome.
The last prosome segment is rounded posteriorly. The maxilliped has no spines on the first
segment, the 2nd segment has a row of fine spinules and a distal tubercle on the inner margin, 3rd
segment very short with no spines, 4th segment terminates in a claw.
F
C
Lubbockia squillimana: Female. A, dorsal; B, urosome ventral view; C, A2; D, maxilliped. Male.
E, urosome ventral view; F, maxilliped. Figures from G.A.Boxshall, The planktonic copepods of
the northeastern Atlantic Ocean: some taxonomic observations on the Oncaeidae (Cyclopoida),
Bulletins of the Natural History Museum, Zoology, 1977, Published by Cambridge University
Press, reprinted with permission.
218
Family Oncaeidae
Oncaeids are small copepods, half the known species are less than 0.6 mm in length, that can be
abundant in tropical coastal waters. The prosome comprises the cephalosome and four free
pedigerous segments. There is a distinct boundary between the prosome and urosome. The
urosome is 5-segmented in females and 6-segmented in males. The genital apertures (on the
genital segment) are paired and situated dorsally in females and ventrally in males. A1 is 6segmented in females and 4-segmented in males. A2 is 3-segmented. The maxillipeds are 3 or 4
segmented and end in a hook or claw, adapted as a grasping tool in males. Females carry paired
egg sacs.
The family Oncaeidae has recently been under review (Huys & Bttger-Schnack, 1997), with
the genera Lubbockia and Pachos, previously considered to be within the family Oncaeidae,
reassigned to separate families. The genera Conaea and Oncaea are included in the family
Oncaeidae.
Genus Oncaea
The genital segment is characteristically large and elongated, particularly so in the males. The
genus Oncaea shows a variation in size within species. The front margin of the cephalosome is
rounded. Characteristics used in separating the species of Oncaea are the proportional lengths
of the body segments, the furca, the 2nd and 3rd segments of A2, the maxilliped and the structure
of P4.
219
Oncaea conifera Giesbrecht, 1891

Female: In lateral view, the 2nd free pedigerous segment of the prosome has a prominent
projection. The genital segment is nearly twice as long as the rest of the urosome. The 3rd
usosome segment is longer than the 4th segment. The furca is as long as the anal segment.
Maxilliped, the 2nd segment has two spines, the distal spine is thicker and longer than the
proximal spine, and the inner margin of the terminal claw is lined with spinules. P5 is
cylindrical, the inner seta is twice as long as the outer one.
Male: Considerably smaller than the female. There is no dorsal hump on the 2nd free pedigerous
segment of the prosome. The genital segment is 3 times longer than the rest of the urosome. The
lappets at the posterior corners of the genital segment are large, triangular and pointed
outwards. The apical setae on P5 are unequal in length.
A
F
E
I
J
Oncaea conifera: Female. A, dorsal; B, lateral; C, A1; D, urosome dorsal; E, maxilliped; F, P5.
Male. G, lateral; H, dorsal; I, genital segment and P5; J, maxilliped. Figures from Heron &
220
Oncaea media Giesbrecht, 1891

Length: females: 0.50-0.92; males: 0.60-0.80 mm.
Female: The widest part of the prosome is at the base of the cephalosome. The genital segment
is twice the length of the rest of the urosome. The furca is nearly twice as long as wide. The
distal spine on the 2nd segment of the maxilliped is thicker but of a similar length to the
proximal spine. Approximately half of the inne r margin of the terminal claw is lined with
spinules. P5 is short, apical setae unequal in length.
Male: Genital segment is 3 times longer than the rest of the urosome. The lappets are large,
rounded with a pointed projection directed outwards. The terminal claw on the maxilliped is not
as long as the preceding segment. The apical setae on P5 are unequal in length. The furca are
approximately twice as long as wide.
Oncaea media: Female. A, A1; B, dorsal; C, lateral; D, urosome; E, maxilliped; F, P4. Male. G,
dorsal; H, lateral; I, maxilliped; J, P4. Figures A,B,C, G & H from Heron & Bradford-Grieve
(1995), D-F, I & J from Razouls (1974).
221
Oncaea minuta Giesbrecht, 1892

Length: female: 0.45-0.58 mm; male: unknown
Female: A very small copepod with a slender body. The prosome is more than twice as long as
wide. The urosome is less than half the length of the prosome. The furca is shorter than the anal
segment, and is approximately twice as long as wide. The outer seta on the furca arises midway up the outer margin. P5 is as wide as long, the inner apical seta is twice as long as the outer
seta.
Oncaea minuta: Female. A, dorsal; B, urosome; C, maxilliped; D, A2. Figure A from Wilson
(1932), B-D from Razouls (1974).
222
Oncaea mediterranea (Claus, 1863)

Length: females: 1.00-1.30 mm; males: 0.70- 1.05 mm.
Female: Posterior of last prosome segment not smoothly rounded, tips directed slightly
outwards. Genital segment elongate and more than twice the length of the rest of the urosome.
Furca is longer than anal segment and approximately three times longer than wide. The outer
seta on the furca arises in the first third of the outer margin.
Male: Smaller than female. Genital segment nearly 3 times as long as the rest of the urosome.
Lappets at the distal corners of the genital segment have prominent backward directed
projections.
G
I
J
K
Oncaea mediterranea: Female. A, lateral; B, P5; C, urosome; D, maxilliped; E, A2. Male. F,

dorsal; G, lateral; H, A2; I, A1; J, urosome; K, maxilliped. Figures A, B & F-H from Heron &
Bradford-Grieve (1995), C-E & I-K from Razouls (1974).
223
Oncaea venusta Philippi, 1843

Female: Prosome widest at the base of the cephalosome. Last prosome segment rounded at the
distal corners. The genital segment is longer than the rest of the urosome and is more rounded
than in O. mediterranea. The furca is longer than the anal segment, and nearly 4 times longer
than wide. P5 is very small, same width as length, the two apical setae are not equal in length.
Male: The body is narrower than the female. The three segments between the genital segment
and the anal segment are very short. The genital segment is more than 3 times longer than the
rest of the urosome. The lappets at the base of the genital segment have small lateral points
turned outward. The furca is shorter than 3 times longer than wide.
Recorded: Rodrigues, Seychelles, Madagascar
K
J
Oncaea venusta: Female. A, dorsal; B, lateral; C, urosome; D, maxilliped; E, A2; F, A1. Male.
G, dorsal; H, lateral; I, urosome; J, maxilliped; K, A2. Figures A, B, G and H Heron &
Bradford-Grieve (1995), C-F and I-K from Razouls (1974).
224
Family Sapphirinidae
Saphiriniids may have a typical cyclopoid body, or the body may be characteristically flattened
dorsoventrally. Cuticular lenses are present at the front of the cephalosome in all females and
some males. The prosome comprises the cephalosome and three or four free pedigerous
segments. Urosome typically 4-segmented in females and typically 6-segmented in males.
Genital apertures are paired and situated dorsolaterally in females and ventrally in males. A1 3to 6-segmented. Male A1 not geniculate. A2 is 4-segmented. Females carry paired egg sacs.
The genera Copilia, Sapphirina and Vettoria (=Corissa) are included in the family
Sapphirinidae.
Genus Copilia
Relatively large transparent copepods with very distinctive features that readily separate them
(especially the females) from the rest of the family. The males and females have very different
body shapes. The female cephalosome is usually rectangular in shape, with two prominent
anterior cuticular lenses. A1 is 6-segmented. A2 is 4-segmented and uniramous. The metasome
segments taper backwards. The posterior margin of the 3rd metasome segment has a stout
median spine mid-dorsally that is pointed backwards. P5 is degenerate, 1-segmented and
uniramous. There is a definited division between the prosome and urosome. The urosome
consists of four segments and the furca are long and slender, usually longer than the urosome.
The male body shape resembles that of Sapphirina, but the furca are long and slender in
Copilia but leaf- like in Sapphirina, and the cephalosome of the male Copilia bears no cuticular
lenses. A2 is 4-segmented and more slender than that of the female. There is no obvious
division between the prosome and urosome. The prosome segments and the first four urosome
segments are wide whereas the anal segment is very small. The furca are rod- like and generally
shorter than those in the female.
225
Copilia mirabilis Dana 1849

Female: The cephalosome is longer than broad; it is as long as the rest of the body (not
including the furca) with the widest part at the posterior margin. The space between the two
cuticular lenses is nearly twice the diameter of the lens. 1st segment of A2 with many short
spines, and a large spine at the distal end covered in spinules. A large characteristic spine with
several large spinules on the mid-section of the 2nd segment. The posterior margins of the
urosome segments have rows of spinules. The anal segment is longer than the rest of the
urosome, and wider posteriorly. The furca is longer than the urosome, with short setae near tip.
Male: The cephalosome is very rounded, and shorter than wide. The first two metasome
segment s are wider than the cephalosome, and the remainder of the segments taper backwards.
A2 is slender, the 1st segment is longer than the 2nd and the 2nd segment is longer than the 3rd.
The 4th segment is almost twice as long as the 3rd. There are two varieties Copilia mirabilis var.
typical and C. mirabilis var. platonyx.
A
B
E
Copilia mirabilis: Female. A, dorsal; B, urosome; C, A2; D, close up of A2; E, P4. Male. F,
dorsal. Figures A-G from Lehnhofer (1926), F from Wilson (1932).
D
C
F
A
B
Copilia mirabilis var. typical: Male. A, A2; B, close up of segments 3 and 4 of A2; C,
maxilliped. Copilia miarabilis var. platonyx: Male. D, A2; E, close up of segments 3 and 4 of
A2; F, maxilliped. Figures from Lehnhofer (1926).
226
Copilia quadrata Dana, 1849

Length: females: 2.20-4.40 mm, males: 3.50-5.70 mm.
Female: The cephalosome is quadrate in dorsal view, and is as long as wide. The space
between the two cuticular lenses is more than three times the diameter of the lens. The 1st
segment of A2 is longer than the 2nd, with a spine at the distal end. The spine on the middle
section of the 2nd segment with no spinules off it. The anal segment is as long as the rest of the
urosome. The furca is more than twice as long as the urosome.
Male: The front margin of the head is straight, or slightly concave. The cephalosome is shorter
than wide. The 1st segment of A2 is nearly twice as long as the 2nd segment, and the 3rd segment
is as long as the 4th .
A
Copilia quadrata: Female. A, dorsal; B, ventral urosome; C, P4; D, segments 3 and 4 of A2; E,
tip of A2. Figures from Lehnhofer (1926).
Copilia quadrata: Male: A, close up of segment 2 of A2; B, A2; C, close up of segments 3 and
4 of A2; D, tip of A2; E, maxilliped. Figures from Lehnhofer (1926).
227
Copilia vitrea (Haeckel, 1864)

Female: The cephalosome is about 1.5 times longer than wide. The anterior of the cephalosome
is narrow; the space between the cuticular lenses is less than the diameter of the lens. The anal
segment is short, less than half the length of the rest of the urosome. The furca is less than twice
as long as the rest of the urosome.
Male: The 1st segment of A2 is shorter than the 2nd segment; the 3rd segment is shorter than the
4th .
A
D
B
F
C
Copilia vitrea: Female. A, dorsal; B, ventral view of urosome; C, segments 3 and 4 of A2.
Male. D, A2; E, segments 3 and 4 of A2; F, tip of A2; G, maxilliped. Figures from Lehnhofer
(1926).
228
Copilia hendorffi Dahl, 1892

Female: The cephalosome is quadrangular in shape, longer than broad, front margin is straight.
The space between the cuticular lenses is just over twice the diameter of the lens. The 1st
segment of A2 has a large spine at the distal end that is about twice as long as the spine on the
2nd segment which is located about one third along its length. The anal segment is as long as the
rest of the urosome. Rows of spinules on posterior margins of 2nd and 3rd urosome segments.
Male: The cephalosome is shorter than wide, and rounded anteriorly. The first two metasome
segments are wider than the cephalosome, and the rema inder of the segments taper backwards.
The 1st segment of A2 has a large spine projecting from its distal end, the spine reaches
approximately half way along the 2nd segment. 1st segment of A2 is longer than the 2nd segment,
the 2nd segment is longer than the 3rd segment, and 4th segment is only just longer than the 3rd.
Recorded: Rodrigues
A
D
Copilia hendorffi: Female. A, dorsal; B, ventral view of urosome; C, distal margin of anal
segment; D, A2; E, segments 3 and 4 of A2; F, P5. Figures from Lehnhofer (1926).
A
B
E
H
Copilia hendorffi: Male. A, dorsal; B, dorsal view of articulation between metasome and
urosome, urosome flipped upwards; C, first 2 urosome segments; D, tip of furca; E, maxilliped;
F, A2; G, tip of A2; H, segments 3 and 4 of A2. Figures from Lehnhofer (1926).
229
Genus Sapphirina
Sapphirina species are large copepods that are flattened dorso-ventrally The majority of species
are transparent and some are iridescent. A pair of large cuticular lenses are situated on the
anterior of the cephalosome. The males are typically larger than the females. The prosome
comprises the cephalosome and up to 4 pedigerous segments. The urosome is 5-segmented in
females and 6-segmented in males. The 1st segment of the urosome (the last thoracic segment)
is usually half the width of the preceding segme nt. In females the remaining urosome segments
are a similar width whereas in males the following segment (genital segment) is nearly as wide
as the last prosome segment. In males the anal segment is small. The furca is leaf- like, with a
superficial seta rising from the blade and 4 seta rising from the outer margin, the exact shape of
the furca varies with species. A1 is between 3- and 6-segmented. A2 is 4-segmented and ends
in a claw. P5 is uniramous, 1 segmented and ends in two minute setae.
The characteristics for separating species include the overall body shape, the length of
cephalosome in relation to the rest of the body, whether the cuticular lenses are visible dorsally,
the shape of the furca, respective lengths of the 4 segments of A2, the number of segments in
A1, the type of spines on the endopod of P2 and the relationship between exopod and endopod
of P4.
230
Sapphirina angusta Dana, 1849

Both males and females are large copepods, they have very elongated bodies, and possess a
wide blunt projection on the inner distal margin of the furca.
Female: Body elongate. Cephalosome approximately 1.5 times longer than wide. Cuticular
lenses are adjacent, visible dorsally, and make up the anterior section of the cephalosome.
Urosome elongate, approximately 1/3 the length of the prosome. Furca is twice as long as wide
with a broad and bluntly rounded process on the distal end of the inner margin. A1 is 5segmented, the 2nd segment is at least as long as the next three segments together. A2 has a long
spine at the distal end of the 1st segment; the 2nd segment is longer than the last two segments
(3rd and 4th ) together. The endopod of P4 is shorter than the exopod, with two foliacious spines
of equal length at the tip.
Male: Body elongate, more than twice as long as wide. Cuticular lenses not visible dorsally.
First four metasome segments wider than the cephalosome. Furca similar to that of the female.
Last endopod segment of P2 with three inner setae, three fo liacious spines and two claw- like
spines at the tip.
Recorded: Western Indian Ocean
A
D
C
F
Sapphirina angusta: Female. A, dorsal; B, furca; C, A2. Male. D, dorsal; E, A1; F, A2; G, P2.
Figures B, C, E, F, G from Lehnhofer (1929), figures A and D from Giesbrecht (1892).
231
Sapphirina auronitens Claus, 1863

Female: Cephalosome wider than long. Cuticular lenses visible dorsally. 1st metasome segment
not as wide as cephalosome, 2nd metasome segment wider than 1st , 3rd metasome segment
approximately 2/3 the width of the 2nd, 5th metasome segment half the width of the 4th . Furca
are ovate in shape and appear inclined inwards with a small spine at the distal end of the inner
margin. The superficial seta on the furca is behind the first marginal seta. 4th segment of A2 is
twice as long as the 3rd, the 2nd segment is as long as the 3rd and 4th together. P4 with two spines
of unequal length at the tip.
Male: Cephalosome wider than long. Cuticular lenses visible dorsally. Endopod of P4 is shorter
than exopod. Furca similar in shape to female, and 1.5 times longer than wide. A1 is nearly half
the length of A2. The 2nd segment of A1 is longer than the 3 following segments together. The
last endopod segment of P2 has a hook- like process on the inner distal corner, a foliacious,
coarsely denticulate spine at the tip, and a long foliacious spine on the outer distal corner. The
endopod of P4 is longer than the exopod, there are two foliacious spines of unequal length at
the tip.
Sapphirina auronitens: Female. A, dorsal; B, A2; C, P4. Male. D, dorsal; E, A1; F, A2; G, P2;
H, P4. Figures A-C from Wilson (1932), D-H from Mori (1937)
232
Sapphirina bicuspidata Giesbrecht, 1891

The furca is distinctive, there are two pointed projections on the inner distal end.
Female: Body short. Cephalosome wider than long, and approximately half the length of the
prosome. Cuticular lenses visible dorsally. Urosome short, approximately a quarter of the
length of the prosome. Furca longer than broad with two pointed projections at the inner distal
margin, the superficial seta is situated in front of the 1st marginal seta. 2nd segment of A2 is as
long as the following two segments. Two foliacious spines of equal length at tip of endopod of
P4.
Male: Body short and rounded. Cephalosome wider than broad. Cuticular lenses not visible
dorsally. 1st metasome segment as wide as posterior section of cephalosome. Remainder of
segments decrease in width giving a smoothly rounded edge (apart from the last thoracic
segment, usually hidden from view dorsally). Furca longer than broad with characteristic two
projections at the inner distal margin.
B
A
Sapphirina bicuspidata: Female. A, dorsal; B, A2; C, furca; D, endopod of P4. Male. E, dorsal;
F & G, left and right endopod of P2. Figures C & F from Lehnhofer (1929), A, B, D, E & G
233
Sapphirina darwini Haeckel, 1864

Female: Cephalosome wider than long. Cuticular lenses visible dorsally and set apart. 1st
metasome segment narrower than the cephalosome. Urosome 6-segmented. Anal segment
small. Furca longer than wide with a small projection on the distal inner margin. The superficial
seta is behind the 1st marginal seta. A1 is 3-segmented and nearly as long as A2. A2 with 1 long
spine on the distal end of the first segment. The last segment on A2 is twice the length of the
terminal claw. Endopod of P4 is less than half the length of the exopod, tip of endopod with
two equal length foliacious spines.
Male: Cephalosome wider than long and rounded anteriorly. Cuticular lenses visible dorsally
and set apart. Furca similar to female.
C
Sapphirina darwini: Female: A, dorsal; B, A1; C, A2; D, P2; E, P4. Male: F, dorsal; G, furca;
H, A2; I, endopod of P4. Figures A-E from Mori (1937), F-I from Giesbrecht (1892).
234
Sapphirina gastrica Giesbrecht, 1891

Similar body shape to S. nigromaculata. In S. nigromaculata, the superficial seta on the furca is
situated in front of the 1st marginal seta, whereas in S. gastrica it is situated behind the 1st
marginal seta, nearly level to the 2nd marginal seta.
Female: Cephalosome wider than long. Cuticular lenses just visible dorsally. Furca longer than
wide with a small pointed process on the distal end of the inner margin, superficial seta situated
behind 1st marginal seta. A1 is 5-segmented and just over half the length of A2. The 2nd
segment of A1 is shorter than the combined length of the following 3 segments. One long spine
at the distal end of the 1st segment of A2, the terminal claw is the length of the last segment.
Endopod of P4 is longer than exopod and ends in two foliacious spines.
Male: Cephalosome wider than long. Cuticular lenses only just visible dorsally. Furca more
than twice as long as wide. A1 is 6-segmented. A2 and P4 resemble female. Endopod of P2 has
3 foliacious spines off the 3rd segment.
E
A
C
D
F
H
B
Sapphirina gastrica: Female. A, dorsal; B, furca; C, P2; D, P4. Male. E, dorsal; F, A2; G, furca;
H, P2. Figures E, F. G & H from Lehnhofer (1929), A, B, C & D from Giesbrecht (1892).
235
Sapphirina intestinata Giesbrecht, 1891

Female: Cephalosome wider than long. Cephalosome makes up over half the length of the
prosome. Cuticular lenses adjacent and visible dorsally. Urosome 6-segmented. Furca more
than twice as long as wide, small pointed projection at the distal inner margin, superficial seta
in front of the 1st marginal seta. A1 is 5-segmented, 2nd segment is longer than the three
following segments. Segment 1 of A2 with a long spine at the distal end. Segment 2 of A2 as
long as the two following segments, the last segment is nearly three times longer than the
terminal claw. Segment 3 of exopod of P2 with three foliacious spines. Endopod of P4 is less
than half the length of the exopod.
Male: Cephalosome wider than long. Cuticular lenses visible dorsally. Furca similar to fe male.
The 3rd segment of the exopod of P2 with 2 foliacious spines.
B
C
D
E
H
Sapphirina intestinata: Female. A, dorsal; B, A2; C, A1; D, endopod of P2; E, P4; F, furca.
Male. G, dorsal; H, endopod of P2. Figures A-E from Mori (1937), F & H from Lehnhofer
(1929), G from Giesbrecht (1892).
236
Sapphirina maculosa Giesbrecht, 1892

Length: females: 1.81-2.13; males: 2.65-2.71 mm.
Female: 2nd segment of A2 longer than the following two segments. Furca longer than wide;
one blunt projection at the distal inner margin, a second blunt projection at the distal tip;
superficial seta approximately the same level as the 1 st marginal seta.
Male: Cephalosome wider than long. Cuticular lenses not visible dorsally. Furca similar to
female. 2nd segment of A2 longer than the following two segments. Exopod of P4
approximately half the length of the endopod.
D
E
F
Sapphirina maculosa: Female. A, A2. Male. B, dorsal; C, A2; D, furca; E, endopod of P2; F,
P4. Figures A, C, D, E & F from Lehnhofer (1929), B from Giesbrecht (1892).
237
Sapphirina metallina Dana, 1849

Furca is a characteristic shape with two long foliacious spines off the distal margin.
Female: Body elongate. Cephalosome approximately as wide as long. Cuticlar lenses adjacent
and visible dorsally. Length of cephalosome approximately half total prosome length. Furca is
characteristic, rectangular, twice as long as wide, superficial seta behind the 1st marginal seta. 2
long spines at the distal end of the 1st segment of A2. Endopod of P4 is 2/3 the length of the
exopod, tip has two spines, one is foliacious.
Male: Body elongate. Cephalosome approximately as long as wide. Cuticular lenses adjacent
and visible dorsally. Furca similar to female. A2 similar to female. Endopod of P2 ends in 3
foliacious spines.
C
F
E
B
D
K
I
Sapphirina metallina: Female. A, dorsal; B, A2; C, P2; D, P4. Male. E, dorsal; F, close up of
urosome; G, A1; H, A2; I, endopod of P2; J, P4; K, furca. Figures A-E from Mori (1937), F-L
from
Giesbrecht
(1892).
238
Sapphirina nigromaculata Claus, 1863

Recognised by the very reduced endopod on P4.
Female: Cephalosome rounded anteriorly and wider than long. Cuticular lenses adjacent and
visible dorsally. Urosome 5-segmented. Furca is elongate ovate and twice as long as wide, the
superficial seta is situated anterior to the 1st marginal seta, the inner margin ends in a small
tooth. A1 is 5-segmented. 2nd segment of A2 as long as the last two segments together. Endopod
of P4 less than half the length of the exopod, two equal length foliacious spines at the tip of
endopod.
Male: Cephalosome wider than long. Cuticular lenses adjacent. Furca similar to female. 2nd
segment of A2 just shorter than the last 2 segments together. Tip of 3rd segment of endopod of
P2 has two foliacious spines and a third spine with one margin denticulate. P4 similar to female.
F
A
C
B
H
G
I
E
J
D
Sapphirina nigromaculata: Female. A, dorsal; B, A1; C, A2; D, endopod of P2; E, furca. Male.
F, dorsal; G, A1; H, A2; I, endopod of P2; J, P4. Figures A & F from Giesbrecht (1892), C, H,
& I from Lehnhoffer (1929), B, D, E, G & J from Mori (1937).
239
Sapphirina opalina Dana, 1849

Female: Cephalosome wider than long, anterior rounded. Cuticular lenses visible dorsally and
just separate. Urosome 5-segmented. Furca characteristic shape, same length as width, pointed
projection at the distal inner margin, superficial seta behind 1st marginal seta. A1 is 3segmented, tip of 3rd segment with two setae fused at the base. A2 longer than A1. 4th segment
of A2 longer than the terminal claw. 3rd segment of endopod of P2 with 3 foliacious spines at
the tip.
Male: Cephalosome wider than long. Cuticular lenses separate and visible dorsally. Furca as
wide as long with projection similar to female. Swimming legs similar to female.
Recorded: Seychelles, Madagascar
G
C
E
D
Sapphirina opalina: Female. A, dorsal; B, A1; C, A2; D, urosome; E, P4. Male. F, dorsal; G,
endopod of P2. Figures A-G from Giesbrecht (1892).
240
Sapphirina ovatolanceolata Dana, 1849

Female: Cephalosome wider than long. Cuticular lenses adjacent and visible dorsally. 2nd
metasome segment longer than 1st segment. A1 is 5-segmented. 3rd and 4th segments of A2 of
similar length, and together shorter than the 2nd segment. Furca twice as long as wide,
superficial seta just behind the 1st marginal seta.
Male: Cephalosome wider than long and pointed on the frontal margin. Cuticular lenses not
visible dorsally, positioned on the anterior ventral surface. A2 very different to female, longer
and more slender. Furca similar to female.
A
E
C
Sapphirina ovatolanceolata: Female. A, dorsal; B, A2; C, P4; D, furca. Male. E, dorsal; F, A2.
Figures A-F from Giesbrecht (1892).
241
Sapphirina scarlata Giesbrecht, 1891

Male and female are moderately wide forms.
Female: Cephalosome wider than long. Cephalic lenses adjacent and just visible dorsally.
Lateral margins of prosome not smooth due to the varying widths of the metasome segments. 1st
metasome segment narrower than the cephalosome, 2nd metasome segment wider than the 1st ,
3rd metasome segment 2/3 as wide as the 2nd, 4th metasome segment same width as 3rd. Furca
twice as long as wide, superficial seta situated in front of the 1st marginal seta, pointed
projection on the distal inner margin. A1 is 5-segmented. Endopod of P4 approximately half the
length of exopod ending in two foliacious spines.
Males: Cephalosome nearly twice as wide as long. Cephalic lenses not visible dorsally. Body
tapers posteriorly, lateral margins smooth. Furca similar to female. Last segment of endopod of
P2 with 2 foliacious spines and a denticulate spine.
Recorded: Madgascar
Sapphirina scarlata: Female. A, dorsal. Male. B, dorsal; C, A2; D, endopod of P4; E, endopod
of P2; E, furca. Figures A-F from Giesbrecht (1892).
242
Sapphirina stellata Giesbrecht, 1891

Female: Cephalosome wider than long. Cuticular lenses are adjacent and visible dorsally.
Urosome is 6-segmented. Furca is longer than wide, small projection on the inner distal margin,
superficial seta is anterior to the 1st marginal seta. A1 is 5-segmented, the 3rd and 4th segments
with a stout spine. The 2nd segment of A2 is longer than the 3rd and 4th segments together, 4th
segment is more than twice the length of the terminal claw. Endopod of P4 is less than half the
length of the exopod.
Male: Cephalosome wider than long. Body oval in shape. Cuticular lenses adjacent and visible
dorsally. Furca similar to female. Last segment of endopod of P2 with 3 foliacious spines at the
tip, also a claw- like process on the inner distal margin of the segment.
Sapphirina stellata: Female. A, dorsal; B, A1; C, P4; D, A2. Male. E, dorsal; F, A2; G,
endopod of P2. Figures A-C and G from Mori (1937), D, E and F from Giesbrecht (1892).
243
Sapphirina sinuicauda Brady, 1883

Female: Cephalosome wider than long. Cuticular lenses adjacent and visible dorsally. Furca
longer than wide, sharply pointed projection on distal inner margin, superficial seta behind 1st
marginal seta. 2nd segment of A2 as long as the following two segments together.
Male: Cephalosome wider than long. Cuticular lenses adjacent and visible dorsally. 1st urosome
segment is narrow and visible dorsally. Furca similar to female. Last segment of endopod of P2
with one foliacious spine and two denticulate spines.
A
Sapphirina sinuicauda: Female. A, dorsal; B, A2; C, furca. Male. D, dorsal; E, endopod of P2.
Figures A, C & D from Giesbrecht (1892), B & E from Lehnhofer (1929).
244
Family Corycaeidae
A large pair of cuticular lenses at the anterior of the cephalosome. The cephalosome is long,
typically longer than the rest of the prosome. Prosome comprises cephalosome and two or three
free pedigerous segments, The third and fourth segments sometimes fused. The 3rd segment is
produced at each posterolateral corner into sharply pointed lateral points. The fourth segment,
sometimes fused with the previous one, bears posteriorly directed lateral points that sit inside
the epimeral plates. The urosome is 2- or 3-segmented in females, and 3-segmented in males.
Furca is narrow and elongate. A1 is short and 6-segmented. A2 is 4-segmented and prehensile
in males. Females carry paired egg sacs. The family Corycaeidae comprises the genera
Corycaeus, and Farranula.
Corycaeus (Agetus) flaccus Giesbrecht, 1891

Female: Prosome 4-segmented. Prosome more than twice the length of the urosome. The outer
lateral points extend to about halfway down the urosome. The inner lateral points are not sharp.
Urosome 1-segmented. Urosome with a small projection mid-dorsally, visible from lateral view.
Lateral margins of the urosome with short hairs. Furca greater than half the length of the
urosome. The inner marginal furcal seta is the longest.
Male: Prosome 3-segmented. Prosome less than twice the le ngth of the urosome. Urosome 2segmented. Genital segment 1.5 times longer than wide, and not as long as the anal segment and
furca together. Furca about 7 times longer than wide.
A
B
E
C
Corycaeus flaccus: Female A, lateral; B, dorsal; C, A2; D, P4. Male. E, dorsal; F, lateral view
of urosome; G, A2. Figures A-G from Tanaka (1957).
245
Corycaeus (Agetus) typicus (Kryer, 1849)

Length: females: 1.45-1.65 mm; males:1.30-1.54 mm.
Female: Prosome 4-segmented. Urosome one-segmented, the shape of the urosome and the
form of the inner lateral points at the end of the prosome are characteristic. The outer lateral
points do not extend to the genital openings. The inner lateral points rise up at right angles to
the body and extend posteriorly to almost the same length as the outer lateral points. The furca
is longer than half the length of the urosome. The inner marginal furcal seta is longer than the
furca itself.
Male: Prosome 3-segmented. Urosome two-segmented. The anterior section of the genital
segment rounded, the posterior section forming prominent rounded corners, each with a seta off
the ventral surface. Genital segment 1.5 times as long as wide. Urosome twice as long as furca.
Furca more than 6 times longer than wide.
Corycaeus typicus: Female. A, dorsal; B, lateral; C, A2; D, P4. Male. E, dorsal; F, lateral view
of urosome; G, A2. Figures A-G from Tanaka (1957).
246
Corycaeus (Corycaeus) crassiusculus Dana, 1849

Female: Prosome 4-segmented. The lateral points at the end of the prosome do not extend as far
as the genital segment. Urosome 2-segmented. The genital segment is large, oval, and about 1.3
times longer than wide. The posterior dorsal section of the genital segment overlaps the anal
segment. The posterior margin of the ventral side of the genital segment with a row of minute
spinules. The anal segment narrows slightly. Posterior margins of anal segment fringed with
small spinules. Furca is divergent, about 6 times longer than wide, and about 3/5 the length of
the urosome.
Male: Prosome narrower than in the female and 3-segmented. The outer lateral points at the end
of the prosome extend to the centre of the genital segment. Urosome 2-segmented. Genital
segment about 1.5 times longer than wide, oval, the lateral margins are smooth and each bear a
plumose seta near the posterior end. The ventral posterior margin of the genital segment with a
row of minute spinules. The posterior margin of the anal segment with a row of spinules along
the join with the furca. The furca are parallel, about 8x longer than wide, and about the length
of the urosome.
Corycaeus crassiusculus: Female. A, dorsal; B, ventral; C, A2. Male. D, dorsal; E, A2. Figures
A, B, C and E from Sewell (1947), D from Tanaka (1957).
247
Corycaeus (Corycaeus) speciosus Dana, 1849

Length: females: 1.90-2.18 mm; males: 1.481.90 mm.
Female: The female is easily recognised by its large size and the moderately long and divergent
furca. The prosome is 3-segmented. The outer lateral points at the end of the prosome extend
beyond the end of the genital segment. The urosome is 2-segmented. The furca are divergent
and as long as the rest of the urosome. The spine at the distal end of the 1st segment of A2 is
more than twice the length of the spine at the proximal end of the 2nd segment.
Male: Prosome narrower than the female and 3-segmented. The outer lateral points at the end
of the prosome reach the centre of the genital segment. Urosome 2-segmented. Genital segment
with a seta on each postero-lateral margin. Anal segment more than twice as long as wide.
Furca as long as the urosome, slender, and about 12 times longer than wide.
Recorded: Rodrigues, Seychelles, Madagascar, CPR.
A
H
C
I
D
Corycaeus speciosus: A, dorsal view; B, urosome; C, P4; D, A2; E, tip of furca. Male. F,
dorsal; G, urosome; H, P4; I, A2; J, tip of furca. Figures A-J from Cervignon (1964).
248
Corycaeus (Monocorycaeus) robustus Giesbrecht, 1891

Female: A large and very robust copepod. Prosome 4-segmented. The urosome is 2-segmented.
The genital segment is large and oval in shape with a distinct tubercle present on the dorsolateral margins. The posterior dorsal section of the genital segment characteristically overlaps
the anal segme nt. The anal segment is very short. The furca are about 1.5 times longer than the
anal segment and are divergent.
Male: Prosome 4-segmented. Urosome 2-segmented. Genital segment large. Anal segment
short. Furca divergent and more than twice as long as anal segment.
A
C
D
Corycaeus robustus: Female. A, dorsal; B, lateral; C, A2; D, P4. Male. E, dorsal; F, lateral; G,
A2; H, P4. Figures A-G from Motoda (1963).
249
Corycaeus (Urocorycaeus) lautus Dana, 1849

Female: Prosome 3-segmented. Urosome 2-segmented. The female is similar to both C. furcifer
and C. longistylis, but is larger in size and differs in the form of the urosome. The genital
segment is relatively narrow. The anal segment is widest at the anterior end. The spine at the
distal end of the 1st segment of A2 is more than three times longer than the length of the spine at
the proximal end of the 2nd segment (in the females of C. furcifer and C. longistylis, the two
spines are of a similar length). Furca are not divergent and are slightly longer than the rest of the
urosome.
Male: Prosome 4-segmented. Urosome 2-segmented. Furca not divergent and about 1.2 times
longer than the rest of the urosome (in C. furcifer the furca is almost 1.5 times longer than the
urosome). The male is similar in general appearance to C. furcifer but is much larger. On P4
there are two setae on the endopod of C. lautus whereas in C. furcifer there is only one.
Recorded: Rodrigues.
A
D
H
G
Corycaeus lautus: Female. A, dorsal; B, lateral; C, A2; D, P4. Male. A, dorsal; F, lateral; G,
A2; H, P4. Figures A-H from Motoda (1963).
250
Corycaeus (Urocorycaeus) longistylis Dana, 1849

Female: Prosome 4-segmented. Prosome a little longer than the urosome and furca together.
The outer lateral points at the posterior of the prosome extend 2/3 down the genital segment.
Urosome 2-segmented. Furca slender and approximately 16 times longer than wide. The spine
at the distal end of the 1st segment of A2 is about the same length as the spine at the proximal
end of the 2nd segment.
Male: Prosome 4-segmented. Prosome only just longer than urosome. Urosome 1-segmented,
the urosomes of C. lautus and C. furcifer are 2-segmented. Furca greater than 1.2 times the
length of the urosome, and not divergent.
Recorded: Rodrigues, Seychelles, Madagascar
A
D
I
E
J
Corycaeus longistylis: A, dorsal; B, lateral; C, A2; D, P4; E, endopod of P4. Male. F, dorsal; G,
lateral; H, A2; I, P4; J, endopod of P4. Figures A-J from Motoda (1963).
251
Corycaeus (Ditrichocorycaeus) andrewsi Farran, 1911

Female: Prosome 4-segmented, and twice as long as the urosome. The outer lateral points at the
end of the prosome are short (not reaching the genital openings) and are notched on the inner
margin. The inner lateral points are short and pointed at the tip. Urosome 2-segmented. The
genital segment has a characteristic shape in lateral view, the antero- ventral angle is almost a
right angle, and at the apex there is a small patch of spinules, there is no ventral hook. The
genital segment is about 1.5 times longer than the anal segment. Furca about 5 times longer
than wide and less than 1/3 the length of the rest of the urosome. The spine at the distal end of
the 1st segment of A2 is about 1.3 times longer than the length of the spine at the proximal end
of the 2nd segment.
Male: Prosome 4-segmented, and is about 1.5 times longer than the urosome. Segment
immediately following the cephalosome has a slight swelling at the central lateral margins. The
outer lateral points at the end of the prosome extend 1/3 the way down the genital segment.
Urosome is 2-segmented. Genital segment broadly oval with a small ventral hook. Anal
segment about 1.5 times longer than wide.
A
B
C
E
Corycaeus andrewsi: Female. A, dorsal; B, lateral view of urosome; C, A2; D, P4; E, outer and
inner lateral points. Male. F, dorsal; G, lateral; H, A2; I, P4. Figures F-I from Mori (1937) and
A-E from G.P. Farran Plankton from the Christmas Island, Indian Ocean I. On the Copepoda
of the family Corycaediae from Proceedings of the Zoological Society London, 1911, , The
Zoological Society of London published by Cambridge University Press, reprinted with
permission.
252
Corycaeus (Ditrichocorycaeus) asiaticus F. Dahl, 1894

Female: Prosome 4-segmented and nearly twice as long as the urosome. The lateral points at the
end of the prosome are long and wide, almost as wide as the widest part of the cephalosome,
and they extend to the genital openings. The urosome is 2-segmented. The genital segment has
a small blunt process behind the genital opening, the processes are visible dorsally as small
lateral projections. There is no ventral hook on the genital segment. There is a small seta from
each genital opening that can be difficult to see. Furca is short and about 5 times longer than
wide.
Male: Prosome 4-segmented, and about 1.3 times longer tha n the urosome. The lateral points at
the end of the prosome extend to the middle of the genital segment. Urosome 2-segmented.
Genital segment is about 1.3 times longer than wide with a prominent ventral hook. Anal
segment is twice as long as wide. Furca about 6 times longer than wide.
A
C
Corycaeus asiaticus: Female. A, dorsal; B, lateral view of urosome; C, A2; D, P4. Male. E,
dorsal; F, lateral view of urosome; G, A2. Figures A-F from Tanaka (1957), G from Motoda
(1963).
253
Corycaeus (Ditrichocorycaeus) erythraeus Cleve, 1904 (= C. dubius Farran, 1911)

Lemgth: females: 0.88-1.09 mm; males: 0.83-0.84 mm.
Female: Prosome slender and 4-segmented. Prosome about 2.4 times longer than wide. The
outer lateral points at the end of the prosome are acute and extend beyond the centre of the
genital segment. The inner lateral points are sharp and acute. Genital segment with a ventral
hook and setae on the genital openings. Genital segment is the same length as the anal segment.
Furca just longer than anal segment. The spine at the distal end of the 1st segment of A2 is
nearly 3 times longer than the length of spine at the proximal end of the 2nd segment.
Male: Prosome 3-segmented. The outer lateral points at the end of the prosome do not extend to
the middle of genital segment. Urosome 2-segmented. Genital segment with a ventral hook,
ventral posterior margin lined with spinules. Anal segment shorter than genital segment. Furca
longer than anal segment.
C
B
Corycaeus erythraeus: Female. A, dorsal; B, lateral view of urosome; C, A2; D, P4. Male. E,
dorsal; F, lateral view of urosome; G, A2. Figures A-G from Tanaka (1957).
254
Corycaeus (Onychocorycaeus) agilis Dana, 1849

Length: females:0.88-1.00; males: 0.71-0.84 mm.
Female: Prosome robust and 4-segmented. Prosome is about 1.5 times longer than the
urosome. Urosome is slender and 2-segmented. The lateral points at the end of the prosome
do not extend to the genital apertures. The genital segment is rounded centrally and the genital
aperture furnished with one seta. Anal segment slightly shorter than the genital segment. Anal
segment more than twice as long as wide. Furca about the same length as the genital segment.
The spine at the distal end of the 1st segment of A2 is about three times longer than the spine
at the proximal end of the 2nd segment. The 2nd segment of A2 is broad and has two teeth on
the distal inner margin.
Male: Prosome 4-segmented. Urosome is 2-segmented. Genital segment is oval, and the
widest section equals its length. Some specimens may have a small ventral hook on the
genital segment. Anal segment shorter than genital segment. Furca longer than anal segment.
Recorded: Rodrigues, Seyc helles, Madagascar.
E
F
B
Corycaeus agilis: Female A, dorsal; B, lateral view of urosome; C, A2; D, P5. Male. H, dorsal;
F and G, lateral view urosome of two individuals; H, A2. Figures A-H from Tanaka (1957).
255
Corycaeus (Onychocorycaeus) catus F. Dahl, 1894

Female: Prosome robust and 4-segmented. Prosome more than twice the length of the urosome.
The outer lateral points at the end of the prosome extend to the genital openings. The inner
lateral points are characteristically long and pointed. Urosome 2-segmented. Genital segment is
longer than the anal segment and furca together. Anal segment just wider than long. Furca
comparatively short, slightly longer than anal segment.
Male: Prosome 4-segmented. Prosome about 1.5 times longer than the urosome. Urosome 2segmented. Genital segment longer than the combined length of the anal segment and furca.
Genital segment with a ventral median hook.
E
B
F
Corycaeus catus: Female. A, urosome dorsal view; B, lateral; C, A2; D, P4. Male. E, dorsal; F,
lateral view of urosome. Figures A-F from Tanaka (1957).
256
Corycaeus (Onychocorycaeus) ovalis Claus, 1863

Female: The large size and the width of the last two prosome segments are characteristic. The
prosome is 4-segmented. The outer lateral points at the end of the prosome extend beyond the
genital opening. The inner lateral points are very broad with acute points. The urosome is 2segmented. The genital segment is three times longer than the anal segment. There is a seta on
each genital opening. Furca is longer than the anal segment. The spine at the distal end of the 1st
segment of A2 is longer than the length of the spine at the proximal end of the 2nd segment.
Male: Prosome narrower than in female and is 4-segmented. The outer lateral points at the end
of the prosome do not reach the centre of the genital segme nt. The genital segment is twice as
long as anal segment. Widest part of genital segment behind the centre, anterior end rounded.
The furca is 1.5 times longer than the anal segment.
G
C
Corycaeus ovalis: Female. A, dorsal; B, A2; C, P4. Male. D, dorsal; E, urosome; F, A2; G, P4.
Figures A-G from Mori (1937).
257
Genus Farranula
Female Farranula are readily separated from Corycaeus by the presence of a beak shaped
posteriorly directed ventral process, visible laterally. The last two prosome segments are fused.
Inner lateral points at the posterior of the prosome are absent (present in Corycaeus species), the
outer lateral points are present. The urosome is 1-segmented in both males and females.
Farranula carinata (Giesbrecht, 1891)
Female: Prosome 2-segmented. Cuticular lenses in profile when viewed dorsally. Lateral points
at the posterior of prosome are long and slender, they extend beyond the centre of the urosome.
Furca half as long as urosome, and fo ur times as long as wide. In lateral view the prosome
appears hump-backed. The urosome has a wide projection dorsally, and a setose pad midventral anteriorly.
Male: Prosome 2-segmented. Urosome widest at the centre, tapered both anteriorly and
posteriorly. Urosome 2.5 times longer than the furca. Furca about ten times longer than wide.
Posterior section of genital projection approximately 2/3 down the urosome.
D
H
Farranula carinata: Female. A, dorsal; B, lateral; C, A2; D, P4. Male. E, dorsal; F, lateral; G,
A2; H, P4. Figures A-G from Motoda (1963).
258
Farranula concinna (Dana, 1849)

Female: Prosome 2-segmented. Prosome is twice the length of the urosome and furca together.
Urosome 1-segmented, slightly swollen towards the anterior end. Furca approximately 1/3 the
length of the urosome and 3 times longer than wide. There is a small patch of setae on the
antero-ventral part of the abdomen. The lateral points at the posterior of the prosome do not
reach the centre of the urosome.
Male: The male is very similar to F. gibbula, only separated by size. F. concinna is shorter by
0.1 mm.
Farranula concinna: Female A, dorsal; B, lateral; C, A2. Male. D, dorsal; F, lateral view of
urosome. Figures A-E from Tanaka (1957).
259
Farranula curta (Farran, 1911)

Female: Similar to female C. carinata but the body is more slender, the lateral spines on the
prosome are shorter and do not reach the genital openings on the urosome. There is no patch of
setae on the dorso-ventral surface of the urosome. Furca 3 times longer than wide.
Male: Urosome 3 times longer than furca. Furca three times longer than wide. Distinctive shape
and features of urosome shown in figure.
Farranula curta: Female. A, dorsal; B, lateral; C, urosome dorsal view; D, A2. Male E,
urosome. Figures A- E from from G.P. Farran Plankton from the Christmas Island, Indian
Ocean I. On the Copepoda of the family Corycaediae from Proceedings of the Zoological
Society London, 1911, , The Zoological Society of London published by Cambridge
University Press, reprinted with permission.
260
Farranula gibbula (Giesbrecht, 1891)

Female: Prosome 2-segmented. Prosome approximately 2.5 times longer than the urosome. A
dorsal hump is conspicuous on the 2nd prosome segment. Lateral points at the posterior of
prosome do not reach the centre of the urosome. The urosome has a characteristic shape. There
is a small setose section on the antero-ventral part of the urosome. The dorsal side of the
urosome is irregular in shape. Furca more than 3 times longer than wide. Urosome about 1.7
times longer than the furca.
Male: Males of F. gibbula and F. concinna are very similar, but are of different lengths, F.
gibbula is longer by approximately 0.1 mm.
B
D
E
Farranula gibbula: Female. A, dorsal; B, ventral view of urosome; C, A2. Male. D, dorsal; E,
lateral; F, A2. Figures A-F from Tanaka (1957).
261
Farranula gracilis (Dana, 1849)

Female: Prosome 3-segmented. No mid-dorsal hump as in F. gibbula. Urosome about twice as
long as furca. In lateral view of the urosome, anterior section the dorsal margin is almost
parallel to the ventral margin.
Male: Prosome 3-segmented. Urosome less than three times longer than the furca. Furca longer
than twice the width. The posterior margin of the genital enlargements approximately 2/3 down
the length of the urosome.
D
I
E
Farranula gracilis: Female. A, dorsal view of prosome; B, lateral; C, A2; D, close up of anteroventral urosome; E, P4. Male. F, dorsal; G, lateral; H, A2; I, P4. Figures A-I from Cervignon
(1964).
262
Order Monstrilloida
Monstrilloid copepods are rare in the plankton. The adults and first nauplius are free-swimming
while all other stages are parasitic in polychaetes or gastropods. Adult males and females are
immediately recognised by their elongate cylindrical shape and their lack of feeding
appendages and second antennae. The first antennae are parallel and directed forwards, in line
with the body. The adult copepod cannot feed and must subsist on food accumulated as a
juvenile, the adults primary function is to reproduce.
The female genital segment is equipped with a pair of slender spines from the ventral surface on
which the eggs are carried. The male genital segment has a variable shaped appendage where
the spermatophores are carried. In this guide we have not attempted to identify the monstrilloid
copepods to species level.
Cymbasoma longisponosum: A, female dorsal; B, female lateral; C, male dorsal. Monstrilla

helgolandica: D, female dorsal; E, female lateral. From Sars (1916).
Order Siphonostomatoida
Family Caligidae
The family is made up entirely of parasites, largely ectoparasitic on fish. The nauplii larvae are
typical and free-living. Adults mainly remain attached to their host, although most are able to
swim freely and are occasionally found in the plankton.
Example of a female parasitic copepod Alebion sp. from the Indian

Ocean. Figure from Wilson (1932).
263
CLASS OSTRACODA
Ostracods are small crustaceans which have a carapace of two valves, hinged across the
top. They are sometimes confused with the cyprid stage of barnacles which they
superficially resemble. They have 7 pairs of appendages, specialized for different tasks.
Their feeding and swimming limbs stick out below the carapace. Some species have a
lateral eye-spot. They shed the carapace seven times during life, growing a new larger
one each time. They are sexually dimorphic, males and females are slightly different
shapes.
Examples of ostracods (From Wickstead, 1965).
264
CLASS MALACOSTRACA
Order Stomatopoda
The stomatopods are more commonly known as mantis shrimps. The adults are benthic,
and typically live in burrows or rock or coral crevices. They range in size from 5 cm to
greater than 35 cm. The second pair of legs are characteristic, very large and powerful.
They can be modified for smashing hard-shelled prey (bivalves, shelled gastropods, crabs
etc) or for spearing fish and other soft-bodied organisms. For those modified for smashing,
some species can harness enough power to shatter a pain of aquarium glass in a single high
speed punch.
Adult females brood their eggs and tend them until they hatch. The larvae are transparent
and pass through up to nine planktonic stages, remaining in the plankton for up to three
months. All stages resemble the adult, with more adult features taken on as development
progresses. In all stages the large 2nd pair of legs and large carapace are distinctive features.
Larvae are usually less than 10 mm, but may reach up to 5 cm in length in some species.
A
Stomatopoda: A, adult Lysiosquilla maculata; B, early larval stage of a species of Squilla.

Figure A from Richmond (1997), B from Calman (1911).
A
Squilla sp. larval stages: A, stage I; B, stage IV; C, stage VII. Figures A-C from Diaz
(1998).
265
Order Mysida
These are commonly known as opossum shrimps and can be especially numerous in areas
such as estuaries. They are shrimp- like, but can be recognized by the presence of very
obvious circular organs called statocysts in the tail fan. The eyes are usually black but
sometimes red. Adult females often have a brood pouch with eggs or developing young
inside.
Examples of mysids and structure of the tail fan showing the statocysts (From Wickstead,
1965).
Order Amphipoda
Amphipods are large crustaceans, sometimes common in the plankton samples. The eyes
are usually well developed, large with many facets (compound eye), but are sometimes
reduced or lacking. Their bodies are elongate and more or less flattened from side to side.
They brood their young and plankton samples often contain large numbers of juveniles
released from the female during the sampling process. They have two pairs of antennae,
with one pair usually very small.
Examples of amphipods (From Wickstead, 1965).
266
Order Isopoda
Isopods are crustaceans with a flattened (dorso-ventrally) body. The body is divided into a
series of segments, which generally decrease in width towards the tail, although they are
very variable in shape. They usually have a pair of simple, small eyes widely separated on
each side of the head and uniramous (single branch) walking legs. Some are associated
with the very surface film and are only collected when this is sampled. An unusual species
which can occur in plankton samples is a parasite of fish called Gnathia. This has a tubular
body with a yellow tinge, probably because of stored lipids. The larvae of another parasitic
isopod group, the epicarids, are often found in plankton samples in low numbers. An
epicarid larvae, which resembles a small isopod hatches from the egg and seeks out and
attaches to a copepod. It moults several times, becoming a microniscid and eventually a
cryptoniscid larvae. It then leaves the copepod and finds its next host, usually a crab.
C
D
G
F
Selection of isopods. A-E, Representatives of family Idoteidae; F, Gnathia sp. male dorsal;
G, female lateral; H, juvenile dorsal; I, Cirolanidae isopod; J, Cryptoniscid larva (A-E
From Naylor, 1957a, F-I from Naylor 1957b, J from from Wickstead, 1965).
267
Order Cumacea
Cumaceans are bottom dwelling, shrimp- like organisms, which can occur in plankton
samples when they emerge at night, or if the bottom is stirred up by wave action in shallow
waters. They generally have an enlarged carapace, a slender abdomen, no tail fan and most
do not have obvious eyes.
Example of a cumacean (from Wickstead, 1965).
268
Order Euphausiacea
Euphausiids, or krill, are relatively large shrimp- like organisms, of which there are 86 known
species. There are only two families of euphausiids, Bentheuphausiidae and Euphausiidae, of
which only the latter have been sampled during the Darwin Programme. Euphausiids often
swarm in large numbers and are generally found in greatest numbers away from the coast.
Sexes are separate and eggs are either released directly into the water column (57 species) or
held in an egg sacs or sacs, carried between the thoracic legs. Eggs are large, typically 0.30.9 mm in diameter, with, at least in free spawned eggs, a perivitelline space of varying
diameter between the embryo and the outer membrane. The larva which emerges from free
spawned eggs is the typical nauplius larva of crustaceans and has only 3 pairs of limbs. The
nauplius develops through a second nauplius stage and then a metanauplius stage, which has
only 2 pairs of limbs. Following the metanauplius are three calyptopis stages, the final of
which moults into the first of several furcilia stages, the number varying with species and
also determined by environmental conditions. Eyes emerge from below the carapace and start
to pigment during the furcilia stages. Light organs (photophores), which are usually red,
develop during this stage and are characteristic of all but 2 species of this order. Generally
there are one at the base of each eye, a pair on the coxae of the second and seventh thoracic
limbs and single ones on the mid line between each of the first to fourth pleopods. In the
Genus Stylocheiron these numbers are reduced. There is a reduction in the number of spines
on the telson as euphausiids approach adulthood, till there are only three spines. However,
they cannot be considered adult until they are capable of reproduction, and till this time are
called postlarval. In the later developmental stages their eyes are generally black or brown,
sometimes red, and can be circular, sub-circular or elongated.
Developmental stages of euphausiids: A, egg; B, nauplius; C, metanauplius; D, calyptopis; E

various successive furcilia stages. From Einarsson (1945).
269
Changes in the telson of the furcilia stage during development, showing the gradual reduction
in the number of spines. From Einarsson (1945).
Euphausiid morphology
The body is divided into the cephalothorax and the abdomen. The cephalothorax includes the
carapace, exposed gills, 5 paired mouthparts, 8 paired thoracic limbs, 2 pairs of antennae and
a pair of eyes. There are 6 segments in the abdomen, which ventrally carry the paired
pleopods, which are used in locomotion. The telson and uropods are found posteriorly.
The paired first antennae, or antennules, each consists of a 3-segmented antennular peduncle
and 2, multi-segmented, antennal flagellae. The anterior-dorsal edge of the first segment of
the antennular peduncle is often extended, forming a plate called the antennular lappet, which
varies in shape between species. The paired second antennae each consist of a basal segment,
which has a 2-segmented antennal peduncle and an antennal scale. On the outer distal margin
of the basal segment there is a spine, which varies in length. The antennal scale lies to the
outside of the peduncle, which terminates in a single long flagellum.
Each of the abdominal pleopods carries a pair of setose plates, an inner endopodite and an
outer exopodite. In Family Euphausiidae, the endopodites of the first and second pleopods
are modified as sexual organs in males. The various hooks and processes, which develop on
270
the pleopods, are termed the petasma. The structure can be used in the identification of
mature individuals. The external reproductive organ of the female is known as the thelycum
and is found on the ventral surface of the sixth and seventh thoracic segments.
Generalised euphausiid, illustrating terminology. 1, Cephalothorax; 2, abdomen; 3, carapace;

4, frontal plate; 5, rostrum; 6, eye; 7, first antennal peduncle; 8, flagella of first antenna; 9,
second antennal peduncle; 10, flagellum of second antennae; 11, antennal scale; 12, thoracic
limbs; 13, exopodite of thoracic limb; 14, endopodite of thoracic limb; 15, gills; 16,
pleopods; 17, male copulatory organ; 18, dorsal keel; 19, dorsal tooth; 20, pleuron; 21,
telson; 22, uropods. From Boden (1954).
Typical euphausiid sex organs. A, female thelycum with two spermatophores attached; b,
male petasma. From Einarsson (1945).
It should be noted that keys are based on adult feature and these same features may not be
present in juveniles.
271
An excellent key and descriptions to the euphausiids of the world are given in Baker et al.
(1990).
Family Euphausiidae
The eyes are well developed. The first and second pairs of pleopods in the male are
transformed into copulatory organs. Photophores are present. The lower posterior margin of
the carapace is not serrated but smooth, or carries one or two small denticles
Genus Euphausia
Denticle are generally present on the sides of the carapace, either one or two pairs. Eyes are
always circular, or slightly sub -circular. Shape of the rostrum is variable, either pointed or
gently rounded. The basal segment of the antennular peduncle often has a small lappet at the
distal end. The seventh thoracic leg consists of a minute process with the exopodite missing,
while the eighth thoracic leg is tiny.
Euphausia diomedeae Ortmann 1894
Length: 10-18 mm.
The anterior extension of the carapace is short, sometimes extended over the eyestalks and
terminates in a slender rostrum. Two denticles are present on the carapace, anteriorly and
posteriorly, towards the lower edge. The basal segment of the first antennal peduncle bears a
bifurcated lobe, the ends of which point forwards and outwards. The second segment has 2
processes distally, the outer one blunt and the other sharp. There is a short low ridge on the
third segment.
Euphausia diomedeae: Adult male from left side and right first antennal peduncle from
above. From Boden(1954).
272
Euphausia gibboides Ortmann 1893

Length: 22-27 mm.
Body quite robust. The anterior projection of the carapace is short, with a broad based but
terminally sharp rostrum. There is a single spine on the carapace, on the lo wer margin,
around half way along. There is a pronounced keel on the dorsal carapace. The eyes are large
and circular. The basal segment of the first antennular peduncle is extended distally into a
long lobe, which points forwards and upwards for the first half of its length and then tapers
abruptly and bends sharply outwards. The upper distal end of the second segment is concave
but projects as a lobe over the proximal end of the third segment. The third segment has a
high dorsal keel with an acute tooth at its dorsal, distal edge. The scale of the second antennal
segment reaches to around the middle of the third segment of the first antennal peduncle. The
spiny process on the outer edge of the scale is just under half as long as the scale. There is a
short tooth on the dorsal surface of the third abdominal segment.
Euphausia gibboides: Adult male from left side. From Boden (1954).
273
Euphausia tenera Hansen 1905

Length: 7-9 mm.
The rostrum is quite short but pointed and there is a slight keel on the dorsal carapace. There
is a single denticle on the edge of the carapace, just behind its mid point. The eyes are small
and slightly oval. The basal segment of the first antennular peduncle of the male has several
strong setae, but no processes. The second segment has a distal, which extends over the third
segment. There are no strong setae on the basal segment of the first antennular peduncle of
the female, but the upper terminal margin has a small flat process, pointing upward and
visible from the side. The second segment has a much smaller lobe than in the male, which
terminates in an angle. There are no processes on the third segment of the peduncle in either
sex.
Euphausia tenera: Adult male from left side and right male first antennal peduncle from
above. From Boden (1954).
274
Genus Pseudeuphausia
Pseudeuphausia latifrons G.O. Sars, 1883
Length: 7-16 mm.
This is the only species of the genera recorded in the region. Clo sely related to Euphausia.
The carapace is produced into a frontal plate, which is transversely cut off so there is no
rostrum. The plate is concave anteriorly and longitudinally. There is a prominent median keel
on the carapace. A small lateral denticle si present on the lower posterior border of the
carapace. The first segment of the antennular peduncle has a dorsal keel bearing 9-10 spines
and cuts across at an angle to the outer corner, with a conspicuous spine on the outer edge.
Before the distal edge of the second segment there is a group of stiff curved setae, which
converge at their tip near the middle of segment 3.
Recorded: Zuza, Madagascar
Pseudeuphausia latifrons: Front part of carapace and first antennal peduncle from above,
and right first antennal peduncle from above. From Boden (1954).
275
Genus Stylocheiron
Mostly small slender species, tending to have elongated eyes with large crystal cones at the
top. They never have denticles on the carapace. The second and third segments of the
antennular peduncle in the female are long and slender while in the male they are short and
thick. The upper flagellum is shorter than the lower in the male and the segments are
flattened and broad, while in the female they are slender and cylindrical. The peduncle of the
second antennal endopodite is extremely elongated, reaching beyond the end of the scale.
The first 2 thoracic limbs are short, while the third is extremely elongated, terminating in a
claw-like segment called a false chelae. However, this limb is fragile and is often broken off
during sampling. The remaining thoracic limbs gradually reduce in size and the last is
rudimentary. The number of photophores is reduced in this genera. They are found at the
base of the eyes, on the coxae of the seventh thoracic limbs and in the ventral midline of the
first abdominal segment.
Stylocheiron affine Hansen, 1910
Rostrum long and slender in the female, reaching to, or beyond the anterior eye, shorter in
the male. The carapace is keeled dor sally. The eyes are less than twice as high as broad, with
the lower end less than twice as wide as the upper end. At the upper narrow end are 4-7
crystalline cones in a transverse row. The first antennal peduncle in both sexes is about as
long as the carapace, relatively shorter in the male. In the female the upper flagellum is
cylindrical and slender and around as long as the peduncle while in the male the distal parts
of both flagella are flattened and wider. The elongated third thoracic limb terminates in the
typical chelae of the genera. The second last segment has a long, strong curved spine and 2
shorter spines.
Length: 5.4-8.5 mm.
Stylocheiron affine: Adult female from left side and terminal segment of third thoracic limb.
From Boden (1954).
276
Stylocheiron carinatum G.O. Sars, 1883

Length: 6-12 mm.
The anterior carapace is short, with a sharp rostrum. The eyes are sub-circular, quite broad at
the base, upper lobe short and narrow with 6-8 slightly enlarged crystalline cones at the top,
in a transverse row. There is a high dorsal keel on the carapace behind the rostrum. The basal
segment of the first antennular peduncle is similar in both sexes and has no lobes or
processes. The flagellae are shorter than the peduncle, the lower slightly shorter than the
upper, and in the male thick at the base. The female flagellae are much thinner than in the
male. At the distal end of the third last segment of the elongated third thoracic limb, there is a
raised portion armed with a spine. The second last segment has three strong setae on its lower
edge.
Recorded: Zuza, Madagascar
Stylocheiron carinatum: Adult female from left side and terminal segments of third thoracic
limb. From Boden (1954).
277
Stylocheiron longicorne G.O. Sars, 1883

Length: 6.2-11.3 mm.
The rostrum is short but pointed. There is a low dorsal keel without a prominent crest. The
eyes are twice as high as the width across the base, the upper part almost as wide as the lower
part. The upper eye has 7-19 enlarged crystalline cones in a transverse row. The false chelae
bear a number of strong bristles on the last 2 segments. The sixth abdominal segment is
slightly longer than the fifth segment.
Stylocheiron longicorne: Adult male from left side and false chelae of the third thoracic limb.
From Boden (1954).
278
Stylocheiron microphthalma Hansen, 1910

Length: 6.1-6.9 mm.
Frontal plate triangular in the male but produced to a slender rostrum in female. Antennal
scale long and narrow, about 12 to 20 times longer than wide at its widest point. Eye small,
upper lobe with 2 enlarged crystalline cones in a transverse row. The false chelae of the third
thoracic leg have a long distally curved spine and 2 shorter spines on the second last
segment. The terminal segment has several strong spiniform bristles. The eyes of this species
are small compared to S. affine and S. suhmii.
Stylocheiron microphthalma: Adult female and false chelae of third thoracic leg. From
Boden et al. (1955)
279
Stylocheiron suhmii G.O. Sars, 1883

Length: 5.5-7.8 mm.
The rostrum is short and pointed. Antennal scale long and narrow, about 12 to 20 times
longer than wide at its widest point. No dorsal crest on carapace. Eyes narrower than in S.
affine and with only 3 well developed crystal cones in a transverse row across the top.
Stylocheiron suhmii: Adult female from left side and false chelae of third thoracic limb.
From Boden (1954).
280
Order Decapoda
The decapods are a very diverse group that comprise the familiar crabs, prawns and shrimps. The
group is split into two suborders, Dendrobranchiata and Pleocymata, separated by the gill
structure. The Dendrobranchiata comprises the pelagic sergestoid and penaeoid prawns. The
suborder Pleocyemata comprises the shrimps, crabs, hermit crabs and lobsters.
Suborder Dendrobranchiata
Two superfamiles Penaeoidea and Sergestoidea, the commercial prawns are penaeids, the truly
planktonic genus Lucifer is a sergestid
Superfamily Penaeoidea
Mature females do not carry or brood their eggs. Eggs are released directly into the surrounding
water. On hatching the larva passes through four distinct larval types, a nauplius, a protozoa, a
mysid, and a postlarva. The nauplius possesses a median eye and three pairs of appendages. There
are five naupliar stages and three protozoeal stages. The protozoea have paired stalked eyes, they
have five pairs of thoracic appendages and two pairs of thoracic maxillipeds. The mysis stage has
eight pairs of thoracic appendages and uropods. In penaeids the uropods appear before the
pleopods. The pleopods develop by the postlarval stage, over several moults the telson and
carapace slowly change to the adult form.
E
F
Penaeus sp. A, adult; B, nauplius; C, protozoa dorsal; D; mysis dorsal; E, mysis lateral; F,
postlarva lateral. Figures A from Richmond (1997), B-F from Cook and Murphy (1971).
281
Superfamily Sergestoidea
Sergestoidea are represented by two families, Luciferidae and Sergestidae. The rostrum is shorter
than the eye-stalk in adults. In members of the Luciferidae, the body is strongly compressed and
gills are absent. For the Sergestidae, gills are present and the body is moderately compressed.
Family Luciferidae
The family is represented by a single genus, Lucifer. Lucifer typically have a long neck, a short
pointed rostrum and very distinctive stalked-eyes. Lucifer species lack gills. Only the first three
pairs of legs (pereopods) are present in the adult, and these lack chelae. The length of eye stalks
are significant in Lucifer species, some species have long eye stalks, where the eye and stalk are
about as long as the neck; others have short eye stalks, where they are about half the length of the
neck. The females loosely carry eggs on the third pleopods, in preserved specimens the eggs have
usually become detached. A nauplius hatches out of the egg and passes through protozoeal and
mysis stages before taking on the adult features. Mature males have a characteristic petasma (male
genital structure) on the first pleopod, two distinctive ventral processes on the sixth abdominal
segment, and a swollen section on the ventral surface of the telson. The females and immature
males are not so easy to separate. Species are identified by the length of the eye-stalk, the spines
on the telson, and in males, the petasma, and the shape of the swollen section on the ventral surface
of the telson. Five species of Lucifer have been recorded in the south and western Indian Ocean,
Lucifer chacei, L. hanseni (recorded from Madagascar), L. orientalis (recorded from the east coast
of South Africa), L. penicillifer and L. typus.
A
Lucifer: A, female carrying eggs on 3rd pleopods; B, nauplius; C, protozoea; D, juvenile. Figures
A-D from Brooks (1882)
282
Lucifer penicillifer Hansen, 1919

Length: females: 10-11 mm; males: 9.5-10 mm
The length of the neck is just greater than twice the length of the eye and stalk.
Male: There are two large hook- like processes on the ventral surface of the 6th abdominal segment,
the second one is larger than the first and tapers to a point. The swollen section on the ventral
surface on the telson is much smaller than that of L. typus and is situated further away from the tip
of the telson. The last pair of lateral spines on the telson are close to the tip, they are large and have
spinules off them on the first half of their length. The centre pair of spines at the tip of the telson
are very short. The outer pair of spines are as long as the width of the tip of the telson, or slightly
longer, and have between 5 and 8 spinules on either side.
Recorded: Seychelles, east African coast.
A
C
B
D
F
Lucifer penicillifer: Male. A, lateral; B, close up of telson; C, telson, dorsal; D, telson, lateral; E,
close up of swollen section. Female. F, thorax and 1st abdominal segment, sp spermatophore case,
pl 1 st pair of pleopods. Figures A from Barnard (1950), B-F from Gordon (1956).
283
Lucifer typus H. Milne-Edwards, 1837

The length of the animals neck is just greater than the length of the eye and stalk.
Male: The swollen section on the telson of the male is large. The end of the telson narrows
abruptly after the last pair of lateral spines on the telson. There are three pairs of spines on the tip
of the telson. The outer pair of spines are the longest, they are shorter than the width of the tip of
the telson, and each of the spines has 4 spinules on the first half of the inside edge. The two pairs
of inner spines do not have any spinules.
Recorded: Seychelles, Rodrigues, east coast of South Africa.
A
D
Lucifer typus: Male. A, lateral; B, posterior part of thorax and first abdominal somite of the male
showing a fully formed spermatophore and two developing spermatophores, p the petasma, III,
base of 3rd pereiopod; C, close up of telson; D, telson, dorsal; E, telson, lateral; F, close up of
swollen section. Figures A from Prez Farfante & Kensley (1997), B-F from Gordon (1956).
284
Lucifer chacei Bowman, 1967

Length of the neck is greater than twice the length of the eye and stalk.
Male: Two large processes on the ventral surface of the 6th abdominal segment, the second is
larger than the first, not as pointed as in L. penicillifer. The swollen section on the ventral surface
on the telson is situated closer to the tip of the telson than that of L. penicillifer. Three pairs of
spines at the tip of the telson. The centre pair of spines are approximately half the length of the pair
of spines alongside. The outer pair of spines have several spinules on either side, they are not as
long as the width of the tip of the telson.
A
F
D
Lucifer chacei: Female. A, lateral; B, telson, lateral. Male. C, 6th abdominal segment and telson,
lateral; D, telson, lateral; E, telson, dorsal; F, apex of telson, dorsal. Figures A-F from Bowman
(1967).
285
Family Sergestidae
The family comprises six genera of free-swimming prawns, Acetes, Sicyonella, Peisos, Petalidium,
Sergia and Sergestes. In this guide we give an example of Sergestes nudus. Adult Sergestes are
difficult to separate, but the larvae are usually very distinct.
Sergestes nudus Illig, 1914
Length: females: 12-31 mm; males: 14-25 mm.
Carapace moderately compressed. Rostrum is short, only just extending beyond the anterior
margin of the carapace, pointed at the tip. Ventral antennular flagellum modified in male to form
clasping organ. Carapace with supraorbital and hepatic spines. Telson lacking movable spines,
tapering sharply towards the tip.
A
Sergestes nudus: A, adult lateral; B, mysis larva at 3.5 mm. Figures A and B from Illig (1927).
B
A
Sergestid larval types: A, protozoa; B, mastigopus; C, Acanthosoma. Figures A-C from Gurney
(1960).
286
Suborder Pleocyemata
The Pleocyemata comprise the shrimps, lobsters, crabs and hermit carbs. Females carry eggs
attached to pleopods. Eggs hatch as the larvae pass through a varying numbers of zoeal stages
before the megalopa stage that settles on the seabed and metamorphoses into a juvenile. The group
is separated into a number of infraorders: Caridea (shrimps), Palinura (spiny lobsters), Brachyura
(true crabs), Anomura (porcelain crabs), Thalassinidea (burrowing lobsters).
Infraorder Caridea
Carideans comprise most of the shallow water shrimps. A characteristic feature of adults is the 2nd
pleura on the abdomen which overlaps the 1st and 3rd. The prezoeal telson has six spines on either
side, the rostrum is cylindrical or laterally compressed, never horizontally flattened throughout.
Abdominal somite 3 sometimes with dorsal process. Abdominal somite 5 often with lateral spines.
Palaemon pacificus: A, stage I; B, adult. Figure A from Gurney (1960); B from Richmond (1997).
Periclimenes sp.: A, late stage; B, adult. Figures A from Gurney (1960), B from Richmond (1997).
287
Infraorder Palinura
The typical larval form of the palinuran families Palinuridae and Scyllaridae is the phyllosoma.
The size of phyllosoma larvae varies with species and development, but can range from 1.5 mm
through to 85 mm. We have not attempted to identify the phyllosoma larvae to species, or genus.
Females carry their eggs until the larvae hatch. The larva passes through a short- lived naupliosoma
stage which moults to the 1st phyllosoma stage. The phyllosoma is very distinctive, it is
transparent, very broad and extremely dorsoventrally flattened. Other unusual features include the
eyes, which are stalked from the first phyllosoma stage. The thin oval carapace does not cover the
whole of the disc-shaped thoracic region. There are four pairs of slender legs, and a relatively
small abdomen. The larvae pass through several phyllosoma stages, the uropods and pleopods
making their appearance from stage V. The last phyllosoma stage passes into an unusual form of
megalopa, the puerulus which resembles a lobster, it has large setose pleopods. The puerulus stage
may remain in the plankton for several weeks before settling and metamorphosing into a juvenile.
Development from the puerulus to the adult is gradual. The larval life ranges from 1 month for
some Scyllarus species up to 23 months for Jasus edwardsii. The families of warm-water species
generally have a shorter larval life than the colder water forms.
Scyllarides sp. slipper lobster: A, late stage phyllosoma; B, adult. Palinurus sp.: C, phyllosoma; D,
adult.
Figures
A
reproduced
by
permission
of
CSIRO
Publishing,
http://www.publish.csiro.au/journals/mfr, from the Australian Journal of Marine and Freshwater
Research, vol. 32 (BF Phillips, PA Brown, DW Rimmer & SJ Braine, 1981), B and D from
Richmond (1997), C from Calman (1911).
288
Infraorder Brachyura
Brachyurans are the true crabs. The complete larval development isnt known for all species,
however the majority of species pass through a number of planktonic zoeal stages and a megalopa
stage. The typical zoea has four carapace spines: a rostral, a dorsal and two lateral. The abdomen
is narrow and ends in a characteristically forked telson. The compound eyes are not on stalks.
Immediately after hatching the zoea is without the characteristic spines, but soon moults to the first
zoeal stage. The megalopa stage resembles the adult, however the abdomen hasnt yet folded under
the body.
C
D
Portunus pelagicus. A, zoea I, lateral; B, zoea I posterior view; C, megalopa; D, adult. Figures AC from Wear & Fielder (1985), D from Richmond (1997).
The zoeal larva of the spider crab, Elamena, are unusual among the brachyurans as the carapace is
only equipped with a rostral spine, the lateral and dorsal spines are absent.
Elamena sp. A, zoea I, lateral; B, 1st juvenile crab; C, adult. Figures A and B from Wear & Fielder
(1985), C from Richmond (1997).
289
Infraorder Anomura
The hermit crabs, squat lobsters and porcelain crabs. Anomurans hatch out as a zoea and pass
through several zoeal stages before metamorphosing to a megalopa, or, in the hermit crabs, a
glaucothoe larva. In zoeal larvae the posterior end of the carapace is produced into two spines,
these may be short as in the hermit crabs and squat lobsters, or long and conspicuous as in some
porcelain crabs. The rostrum is typically robust and not long, with the exception of the porcelain
crabs whose rostrum may be several times longer than the body.
A
Porcelain crab. A, unknown porcellanid larva. Petrolisthes sp. B, zoea I; C, zoea II; D, adult.
Figures A from Wickstead (1965), B & C from Shenoy & Sankolli (1967), D from Richmond
(1997).
B
Hermit crab, Pagurus sp. A, zoea I; B, glaucothoe (equivalent to a megalopa); C, adult. Figures A
and B from Shenoy (1967), C from Richmond (1997).
Squat lobster. A, unknown galatheid zoea; B, Allogalathea elegans adult. Figure A from
Wickstead (1965), B from Richmond (1997).
290
Infraorder Thalassinidea
Thalassinids are burrowing lobsters, they are filter- feeding lobster-shaped crustaceans that live in
complex tunnel systems within mud or muddy sand. Females carry the fertilised eggs. On
hatching, the larva passes through several zoeal stages and a post-larval settlement stage.
Thalasinid larvae can be separated into two types, the homarine group (families Axiidae and
Callianassidae) and the anomuran group (families Upogebiidae and Laomediidae).
Homarine group. The typical zoea has a long flattened rostrum with serrated edges. There is
usually at least one dorsal spine on the abdominal segments. All zoeal stages have a median spine
on the telson.
B
C
A
Homarine group, Callianassa sp., last zoeal stage. A, rostrum and eye, dorsal; B, lateral; C, telson
and uropods, dorsal. Figures A-C from Williamson (1967).
Anomuran group. The typical zoea has a small round rostrum. There are no dorsal spines on the
abdomen. The median spine on the telson is small or absent, always absent on zoea stage I. In
Upogea sp. there are four planktonic zoeal stages and two postlarval stages.
A
E
B
G
Burrowing lobster Upogea sp. A, zoea I; B, telson zoea I; C, zoea II; D, zoea III; E, telson zoea III;
F, zoea IV; G, postlarva I; H, adult. Figures A-G from Shenoy (1967), H from Richmond (1997).
291
PHYLUM PHORONIDA
Phoronids are wormlike organisms, which live in tubes on the seabed. Their actinotroch larvae
can be up to 2 mm in length and may be found in plankton samples. As the larvae grow they
develop a crown of long tentacles.
Examples of actinotroch larvae. A: Early larva; B: Late larva (From Wickstead, 1965).
PHYLUM BRYOZOA
Phylum Bryozoa is sometimes called Ectoprocta. The adults of this group are colonial, forming
gelatinous or firm encrusting mats, or raised coral- like structures. They have larvae with a
chitinous, flattened shell composed of two triangular plates, called a cyphonautes larvae. The
larvae are 0.2-0.7 mm in length and can be common in the plankton at particular times of year.
Bryozoan cyphonautes larva (From Wickstead, 1965).
292
PHYLUM BRACHIOPODA
Commonly known as lamp shells, the adults of these live on the seabed and superficially
resemble bivalve molluscs. They have an external calcareous bivalve shell and attach to rocks
with a fleshy stalk. Their larvae occur in the plankton and also have a bivalve shell which is
usually transparent. Internally they have a series of ciliated tentacles that can emerge from the
shell.
Examples of brachiopod larvae (From Wickstead, 1965).
PHYLUM ECHINODERMATA
Echinoderms include starfish, brittle stars, feather stars, sea urchins and sea cucumbers (which
includes the Bche-de-mer). The larvae of all of these can be common in the plankton and are
all slightly different. Starfish have a typical bipinnaria larvae which has ciliated bands (which
are difficult to see in preserved specimens). As they develop they become more complex, till a
small star-shaped larvae develops at one end. This then settles on the sea bottom and develops
into an adult. The bipinnaria larva is similar to the auricularia larvae of the sea cucumber and
the doliolaria larva of the feather star. Brittle stars and sea urchins have similar larvae, which
resemble arrowheads.
Examples of echinoderm larvae. A: Ophiopluteus larvae of brittle star; B: Late ophiopluteus

larva; C: Echinopluteus larvae of sea urchin; D: Auricularia larva of sea cucumber; E:
bipinnaria larva of starfish; F: Doliolaria larva of feather star. (From Wickstead, 1965).
293
PHYLUM CHAETOGNATHA
Chaetognaths, commonly called arrow worms, because they are slender and dart like arrows,
are a very individual group, so are in a phyla of their own. Most are pelagic, but there are a few
benthic species which may be taken in inshore samples. They are important, common
predators, with a set of long, generally brown, grasping hooks on each side of the head for
capturing prey. The mouth is behind the hooks and has one or two rows of external teeth on
either side. Chaetognaths have a tubular elongated body which is usually transparent when they
are alive, but in many species becomes opaque when preserved. The animal is divided by
internal partitions into three sections head, trunk and tail. The head is usually separated from
the trunk by a slight constriction. Most have a pair of small eyes. Posterior to the eyes, on the
head and/or anterior part of the trunk, is an oval band of ciliated cells known as the corona
ciliata. This feature is important in the separation of the various genera, but is difficult to see in
most preserved specimens, so has not been included in the descriptions.
Chaetognaths are hermaphrodite, having organs of both sexes. The ovaries are situated in the
trunk, are usually elongate, extending from the trunk/tail septum (partition) at the posterior end
of the trunk, forwards to different degrees, depending on the species and stage of maturity.
Eggs are generally visible inside the ovaries in mature specimens. Because chaetognaths go
through cycles of egg release and egg maturation, large mature individuals can appear
immature if they have just released all their mature eggs.
General chaetognath structure: Ventral view of complete chaetognath and dorsal head view.
(From Pierrot-Bults and Chidgey, 1988).
294
Seminal vesicles, in which sperm is stored after being produced in the testes, are found on
either side of the tail section, the degree of development depending on maturity. Inside the
trunk, extending along its whole length, is the intestine, which terminates in the anus on the
ventral surface, at the tail/trunk septum. Around the neck region, the intestine in some species
may be extended into a short lateral extension or diverticulum. There are paired fins on either
side of the body, either a single short pair, two pairs or a pair of long continuous fins. The
shape and arrangement of the fins is particularly useful in identification, as is also the position
at which the anterior fin starts in relation to a nervous ganglion situated on the ventral anterior
surface. This ganglion resembles an opaque patch. There is a further single fin on the tail.
There are supportive fin rays in the fins, which may be continuous or partial, depending on
species.
Some chaetognaths have a thickening of the epidermis in the neck region forming a collarette
of tissue between the head and the anterior fin, particularly developed in Pterosagitta draco.
Some species are rigid when preserved, while others are flaccid. There is gradual development
of adult characteristics from hatching of the egg, so it can be difficult to positively identify
immature specimens.
Chaetognaths from the Darwin samples have not been examined in detail, so mainly
descriptions of the species previously recorded from the western Indian Ocean have been
included here.
Further useful information on identification can be found in Alvario (1967), Bieri (1991),
Lutschinger (1993), Michel (1984) and Pierrot-Bults and Chidgey (1988).
295
Family Pterosagittidae
Genus Pterosagitta
Only one pair of lateral fins, which are completely rayed. Two rows of teeth. Massive
collarette, present even in young specimens. Only one widely distributed species.
Pterosagitta draco (Krohn, 1853)
Opaque, firm and stout body. Large head with obvious neck. On each side of head 8-10 hooks,
6-10 anterior teeth, 8-18 posterior teeth. No intestinal diverticula. The ventral ganglion is
situated about midway along the trunk, slightly closer to the neck. Massive collarette, present
even in young specimens, extending from the neck over the paired fins, thinning towards the tip
of the tail. Collarette can be damaged during sampling. A single pair of lateral fins,
semicircular in shape and with no rayless zones, run between the trunk/tail septum and the
seminal vesicles. Mature ovaries can reach to the neck region, ova in 2 rows arranged
dorsoventrally. Ova large and round, but can be compressed into flat cylinders in mature
specimens. The seminal vesicles touch the lateral fins and are close to the tail fin. Length up to
16 mm.
Pterosagitta draco: A, maturity stage 1; B, stage 2; C, stage 3; D, stage 4 (From Alvario,

1967).
296
Family Krohnittidae
Genus Krohnitta
This genus has one pair of lateral fins, extending from the tail section to less than half way
along the trunk. There is no collarette or gut diverticula and they have a single row of teeth on
either side at the top of the head. The teeth are long and meet at the tips to give a funnel shape.
Krohnitta pacifica (Aida, 1897)
The head is small with a conspicuous neck. On either side of the head there are 8-11 hooks and
up to 16 teeth in a single row. The body is slender and transparent, with a single pair of lateral
fins extending from just beyond half way down the trunk, to the base of the tail fin. The fin rays
cover the outer half of the fin surface, leaving the internal and anterior part without rays. There
is no collarette or gut diverticula. The ovaries in mature specimens can reach the region of the
ventral ganglion, extending sometimes to the neck. The ova are in one line. The seminal
vesicles touch both the lateral and tail fins. Total length 6-8mm
Krohnitta pacifica: A, maturity stage 1; B, stage 2; C, stage 3; D, stage 4; E, specimen from the
Atlantic, stage 2; F detail of head (From Alvario, 1967).
297
Krohnitta subtilis (Grassi, 1881)

The body is long, thin, transparent and floppy. On either side of the small head there are 6-9
strong hooks and 10-13 very prominent teeth in a single row covering the anterior of the head.
No collarette or gut diverticula present. The single pair of broad lateral fins begin at a point
between two thirds and half the distance between the ventral ganglion and the caudal septum.
One third of their length is on the trunk and they are wide and semicircular in shape, almost
totally without rays. The rays are short and wide apart. Lack of supportive rays means these
fins are often badly damaged. The seminal vesicle touch both the lateral and tail fins. The
ovaries are short, reaching the anterior end of the lateral fins. The large ova are in two dorsoventral rows. Total length 12-16 mm.
Krohnitta subtilis: A, maturity stage 1; B, stage 2; C, stage 3; D, stage 4; E detail of head; F right
teeth (From Alvario, 1967).
298
Familly Sagittidae
Genus Aidanosagitta
Lateral fins are completely rayed, with fin rays almost at right angles to the body wall.
Intestinal diverticula present. Seminal vesicles generally situated just behind the posterior fins,
more or less apart from the tail fin. Tail segment relatively long.
Aidanosagitta bedfordii (Doncaster, 1903)
A small chaetognath. Prominent collarette, especially from neck to ventral ganglion.
Conspicuous gut diverticula. On each side of the head 8-12 hooks, 1-3 anterior teeth and 2-3
posterior teeth. The ventral ganglion is wide and long for the size of the chaetognath. Fins are
completely rayed. In mature individuals the ovaries reach the ventral ganglion. The ova are
small in number, large, oval and arranged in a single row, separated unequally. The seminal
vesicles are close to the posterior lateral fins, but separated from the tail fin. Length up to 4
mm.
Recorded: Central Indian Ocean (Tokioka, 1956), but usually a coastal species??
Aidanosagitta bedfordii: A, maturity stage 3; B, stage 4; C, detail of head (From Alvario,

1967).
299
Aidanosagitta neglecta (Aida, 1897)

Body firm rigid and opaque, the same width from the head to the tail septa. Small head with an
inconspicuous neck. A gut diverticula is present. Anterior fins begin a little behind the ventral
ganglion, the posterior ones are broader and mostly situated on the tail section. Both pairs of
fins are completely rayed. On each side of head 6-8 hooks, 5-7 anterior teeth and 13-18
posterior teeth. A collarette extends from the head to the front of the anterior fin and also from
the seminal vesicles to the tail fin. Mature ovaries may reach beyond the ventral ganglion. The
ova are large, round and arranged in a single row. The seminal vesicles almost touch the
anterior fin, but are well separated from the tail fin. Length up to 10 mm.
Aidanosagitta neglecta: A, maturity stage 1; B, stage 2; C, stage 3; D, stage 4; E detail of head

(From Alvario, 1967).
300
Aidanosagitta regularis (Aida, 1897)

The body is opaque, rigid and brittle, the same width from the head to the tail septum. The head
is small and pointed at the top, and the neck is not conspicuous. Anterior fins begin just behind
the ventral ganglion and are shorter and narrower than the posterior fins. Both pairs of fins are
completely rayed. On each side of the head up to 11 hooks, 2-4 anterior teeth, 4-7 posterior
teeth. Strongly developed collarette on the neck and also on mid-body and posterior to the
seminal vesicles. A gut diverticula is present. Mature ovaries extend to the posterior end of the
ventral ganglion. The ova are large and arranged in a single row. The seminal vesicles touch the
posterior fin, but are well separated from the tail fin. Length up to 9.5 mm.
D
Aidanosagitta regularis: A, maturity stage 1; B, stage 2; C, stage 3; D, stage 4; E detail of head

(From Alvario, 1967).
301
Genus Decipisagitta
Intestinal diverticula present, Lateral fin rays diagonal to body wall rather than at right angles,
missing in some areas.
Decipisagitta decipiens (Fowler, 1905)
Slender, variable transparency, tending to be opaque. Small square head with a conspicuous
neck. On either side of the head 5-7 long strong hooks (usually 6), 4-13 anterior teeth and 6-20
posterior teeth. Well developed gut diverticula. No collarette. Anterior fins narrow, reaching
the posterior edge of the ventral ganglion and have a rayless zone at their anterior edge. The
posterior fins are equally situated over the trunk and tail sections, triangular and also have an
anterior rayless zone. Seminal vesicles equidistant between posterior and tail fins. In mature
specimens the ovaries can reach the posterior end of the anterior fins. The ova are quite large,
oval-cylindrical in shape and arranged in a single row. Length up to 20 mm.
Decipsagitta decipiens: (From Pierrot-Bults and Chidgey, 1988).
302
Genus Ferosagitta
Intestinal diverticula present. The 2 species recorded from the region in this genera F. ferox and
F. robusta are very similar and there has been great confusion in their taxonomy by various
authors, which may still not have been adequately resolved. Because of this, some of the details
given for them may not be correct. F. hispida was recorded as common in Madagascar by
Furnestin and Radiguet (1964). However F. hispida is a neritic American Atlantic species, so
this record is questionable.
Ferosagitta ferox (Doncaster, 1903)
Body opaque, rigid and firm, the same width from the neck to tail section. Large head with
conspicuous neck. On either side of head normally 5-6 large, strong hooks, 7-10 anterior teeth
and 12-14 posterior teeth. Thick collarette, extending from neck to ventral ganglion and
covering the rest of the body as a thin layer. Conspicuous gut diverticula. Anterior fins start at
middle of ventral ganglion and are longer than the posterior fins. The anterior fins are
completely rayed while the posterior fins have a small internal rayless zone. The ovaries are
wide, in mature specimens, filling the body cavity and reaching the neck region. The seminal
vesicles touch both the posterior and tail fins. Length up to 13 mm.
Ferosagitta ferox: A, maturity stage 1; B, stage 2; C, stage 3; D, stage 4 (From Alvario, 1967).
303
Ferosagitta robusta (Doncaster, 1903)

Strong, firm, opaque body, the same width from the head to the tail septum. On either side of
the head 5-7 hooks, 6-11 anterior teeth and 8-15 posterior teeth. Anterior fins start at the
posterior edge of the ventral ganglion. Anterior fins completely rayed, posterior fins with a
narrow rayless zone, mostly on the tail section. Very broad head and large eyes. Conspicuous
collarette extending from the head to the edge of the anterior fins. Gut diverticula present.
Mature ovaries can extend to the neck region. Seminal vesicles close to both lateral and tail
fins. Length up to 22 mm.
Ferrosagitta robusta: A, maturity stage 1; B, stage 2; C, stage 3; D, stage 4 (From Alvario,

1967).
304
Genus Flaccisagitta
Very limp, flaccid body. The lateral fins show distinctive rayless zones. No intestinal
diverticula. Anterior fins separate from posterior fins and start some distance behind ventral
ganglion.
Flaccisagitta enflata (Grassi, 1881)
Body wide, floppy, transparent and widest around its mid- length. On either side of the head 710 hooks, 6-11 anterior teeth and up to 16 posterior teeth. Head is wider than long with a
conspicuous neck. Anterior and posterior lateral fins short, approximately the same length, well
separated from each other. Two thirds of the posterior fins are on the trunk section. Both lateral
fins are only rayed on their outer posterior edges. The ventral ganglion is situated midway
between the head and the anterior fins. No collarette or intestinal diverticula. Ovaries short,
broad, oval, extending when mature no further than the posterior edge of the anterior fin, but
are usually shorter. Ova are quite large, arranged in 3 parallel lines. Small spherical, seminal
vesicles touch the caudal fin, but are well separated from the posterior fins. Length up to 30
mm.
Flaccisagitta enflata: A, maturity stage; B, stage 2; C, stage 3; D, stage 4 (From Alvario, 1967).
305
Flaccisagitta hexaptera (dOrbigny, 1843)

Body transparent and floppy, widest around its mid- length. On either side of the head 6-10
hooks, 2-6 long, sharp anterior teeth which distinctively usually project forwards, 2-8 posterior
teeth. Anterior fins are small and rounded, starting well behind the ventral ganglion, rays only
on outer edge. The greater part of posterior fin located on the trunk section, fin rays only on
outer edge, missing on the anterior portion. Broad head is small for the size of the body and the
neck is conspicuous. No gut diverticula or collarette. Ovaries long and narrow and do not fill
the trunk cavity, in mature specimens sometimes extending to the level of the ventral ganglion.
Ova small, arranged in 3 rows. Small, spherical seminal vesicles, lying nearer to the caudal
than lateral fins, but separate from both. Length up to 70 mm.
Flaccisagitta hexaptera: A, maturity stage 1; B, stage 2; C, stage 3; D, stage 4 (From Alvario,

1967).
306
Genus Mesosagitta
Body musculature weak. Collarette inconspicuous or missing. Intestinal diverticula present.
Posterior lateral fins more on the trunk than tail section. Seminal vesicles well behind the
posterior fins. Fin rays are sparse or missing in some parts of the lateral fins.
Mesosagitta minima (Grassi, 1881)
Slender, opaque, limp body, wider at the level of the ovaries. Lateral fins rounded, both with
widely spaced fin rays along the edges. The fins are delicate and are usually damaged. Anterior
fins start slightly behind ventral ganglion. Head small and neck well defined. On either side of
head 7-9 short, strongly curved hooks, 2-7 anterior teeth and 6-16 posterior teeth. No collarette.
Small intestinal diverticula. The mature ovaries are short and broad, reaching half way up the
posterior fins. They contain a few very large eggs arranged in a single row. Seminal vesicles
are close to the caudal fin, but well separated from the posterior fins. Length up to 11 mm.
Mesosagitta minima: A, maturity stage 1; B, stage 2; C, stage 3; D, stage 4 (From Alvario,

1967).
307
Genus Pseudosagitta
Body limp, but not as much as in Flaccisagitta. Distinct rayless zones on lateral fins. No
intestinal diverticula. Anterior and posterior fins joined by inflated tissue. Anterior fins start at
or close to the ventral ganglion.
Pseudosagitta lyra (Krohn, 1853)
Bulky, floppy, transparent body. Head wider than long. On either side of the head up to 10
hooks, 2-8 anterior teeth and up to 12 posterior teeth. Anterior and posterior fins joined by a fin
bridge of tissue. In smaller specimens this fin bridge is wide, giving the impression of
continuous fins. Anterior fins long and rounded, starting near the posterior end of the ventral
ganglion, increasing in width towards the posterior. Internal region of both lateral fins rayless
and anterior section of anterior fins completely rayless. The tail fin is split into two paddles. No
gut diverticula or collarette. Mature ovaries long and broad and may reach the middle of the
anterior fins. Ova are in 4 dorsoventral rows. The small, spherical seminal vesicles are separate
from both the posterior and caudal fins, but are closer to the posterior fins. Length up to 42
mm.
Pseudosagitta lyra: A, maturity stage 1; B, stage 2; C, stage 3; D, stage 4 (From Alvario, 1967).
308
Pseudosagitta maxima (Conant, 1896)

Body large and transparent. A broad fin bridge connects the lateral fins. The anterior fins start
at the anterior edge of the ventral ganglion, compared to P. lyra where they start at the posterior
edge. Fin rays are found only on the outer posterior edges of the lateral fins. On each side of
the head 5-11 hooks, 4-6 anterior teeth and 5-8 posterior teeth. Collarette and gut diverticula
absent. Mature ovaries can reach around the ventral ganglion. Seminal vesicles are closer to the
lateral than the caudal fins. Length up to 90 mm.
Distribution: A deep mesopelagic species, bathpelagic in warm waters, unlikely to occur near
the surface.
Pseudosagitta maxima: Maturity stage 0 (From Lutschinger, 1993).
309
Genus Sagitta
Intestinal diverticula absent, fins completely rayed.
Sagitta bipunctata Quoy and Gaimard, 1827
Body rigid and opaque. Trunk is about the same width along its length. No gut diverticula.
Anterior fins wide and roundish, beginning just behind the ventral ganglion. The posterior fins
are wider and longer than the anterior fins and are equally distributed over the body and tail
sections. Fins completely rayed. On either side of head 5-10 hooks, 4-8 anterior teeth and 8-16
posterior teeth. There is a prominent collarette from the neck to the anterior fins, between the
lateral fins and also between the posterior fins and the seminal vesicles. Mature ovaries are long
and can reach the ventral ganglion. Ova are quite large, irregularly arranges in 1 or 2 rows. The
seminal vesicles touch the caudal fins, situated well behind the posterior fins. Length up to 19
mm.
Sagitta bipunctata: A, maturity stage 1; B, stage 2; C, stage 3; D, stage 4 (From Alvario, 1967).
310
Genus Serratosagitta
Intestinal diverticula absent. Hooks serrated. Seminal vesicles situated just behind the posterior
fins and well separated from the tail fin. This group requires sorting out with genetical
techniques as misidentifictions are probably occurring (A. Pierrot-Bults, pers. com.)
Serratosagitta pacifica (Tokioka, 1940)
Body translucent, slender, firm and rigid, with the trunk region of the same width along its
whole length. Head small with a conspicuous neck. Lateral fins completely rayed, apart
sometimes for a small portion at the front of the anterior fin. Anterior fins start slightly behind
posterior edge of ventral ganglion. On either side of the head 5-9 serrated hooks, 3-13 anterior
teeth and 8-24 posterior teeth. No intestinal diverticula. A thin collarette can be present on the
neck and between the seminal vesicles and caudal fin. Mature ovaries can reach the anterior
edge of ventral ganglion. Ova are in a single row. Seminal vesicles touch the posterior fins, but
are separated from the caudal fin. When they are in good condition the vesicles have a series of
chitinous teeth on the side. Length up to 14 mm.
Serratosagitta pacifica: A, maturity stage 1; B, stage 2; C, stage 3; D, stage 4 (From Alvario,

1967).
311
Serratosagitta pseudoserratodentata (Tokioka, 1939)

Body quite transparent and slender, slightly wider in middle. Neck not obvious, sometimes with
a slight collarette. No gut diverticula. On either side of head 6-7 serrated hooks, 4-6 anterior
teeth and 10-12 posterior teeth. Anterior fin starts just behind ventral ganglion, without any
rayless zone. Posterior fins longer than anterior fins and rounded in shape with anterior portion
and inner zone in front of tail septum rayless. In mature specimens the ovary can extend nearly
to the ventral ganglion. Seminal vesicles are situated close to the end of the posterior fin, but
separated from the caudal fin. This is a smaller species than S. serratodentata, with fewer teeth
and different shape of seminal vesicles. Length up to 9 mm.
Distribution: Recorded along the east coast of Africa (Andreu et al. 1989), but questionable as
is a north Pacific species.
A
C
Serratosagitta pseudoserratodentata: A, stage 4 of maturity; B, hook; C, detail of hook end; D,

detail of head (From Tokioka, 1939).
312
Serratosagitta serratodentata (Krohn, 1853)

Body needle- like, firm and opaque. Anterior fins start just at the posterior edge of the ventral
ganglion. Lateral fins with small anterior internal rayless zones. On either side of the small
head 5-9 serrated hooks, 6-11 anterior teeth and 12-20 posterior teeth. Small collarette present
in the neck region and no gut diverticula. Ovaries, when mature, can extend to the ventral
ganglion. Large ova. The seminal vesicles almost touch the posterior fins, but are separate from
the tail fin, almost touching both fins when mature. Seminal vesicles in mature and perfect
specimens with two anterolateral papillae. Even in immature specimens in good condition, a
membranous supporting structure can extend from the lateral fins to the anterior part of each
vesicle. Length up to 15 mm.
Distribution: Madagascar, east Africa.
Serratosagitta serratodentata: Maturity stage 2; B, detail of hook (From Lutschinger, 1993).
313
Serratosagitta tasmanica (Thompson, 1947)

Rigid, firm, opaque body. Anterior fins begin at posterior end of ventral ganglion. Anterior fins
rayless in their anterior part, posterior fins rayless in their anterior part and inner part. Lateral
fins close together. On both side of the head 6-9 serrated hooks, 6-9 anterior teeth and 9-15
posterior teeth. No gut diverticula. A tiny collarette may occur at the neck and between the
posterior and caudal fins. Mature ovaries, with quite large ova, extend to between the anterior
and posterior fins. Seminal vesicles with conspicuous anterior knob with protuberances,
separated from posterior and tail fins, closer to posterior fins. Length up to 30 mm.
Distribution: Really an Atlantic species, but may just penetrate round South Africa. S. pacifica
is more likely in the Indian ocean.
Serratosagitta tasmanica: Maturity stage 2 (From Lutschinger, 1993).
314
Genus Solidosagitta
Body stout and stiff with very strong muscles and prominent collarette. Intestinal diverticula
obvious. Posterior lateral fins more on body than tail section. Small pigmented area in eye.
Distinct rayless zones on lateral fins.
Solidosagitta planctonis (Steinhaus, 1896) and Solidosagitta zetesios (Fowler, 1905)
These two species are very similar and are possibly only different forms of the same species - S.
planctonis, forma planctonis and forma zetesios.
General description - Body strong, firm and opaque. Collarette is prominent, extending to the
anterior fins and even to the tail in mature specimens, forming a connecting bridge between
lateral fins. Intestinal diverticula present. Anterior fins may reach the posterior end or middle of
the ventral ganglion and are rayless in the anterior and inner parts. Posterior fins are triangular,
with an anterior and inne r rayless zone. In mature specimens the ovaries can reach beyond the
ventral ganglion. Ova are small and arranged dorsoventrally in 3 rows. The seminal vesicles are
located midway between the posterior and tail fins, touching neither.
Forma planctonis has a maximum body length of 37 mm, 8-11 hooks, 6-9 anterior teeth, 10-14
posterior teeth and the anterior lateral fin starts mid-way up the ventral ganglion.
Forma zetesios has a maximum body length of 45 mm, up to 11 hooks (usually 8-10), up to 12
anterior teeth (usually 8-10), up to 22 posterior teeth (usually 15-19) and the anterior fin starts
at the posterior end of the ventral ganglion.
Distribution: Deep water species, unlikely to be sampled above 100 m.
A: Sagitta planctonis forma planctonis. B: Sagitta planctonis forma zetesios (From PierrotBults and Chidgey, 1988).
315
Genus Zonosagitta
No gut diverticula. Both pairs of fins have an anterior and long internal rayless zone.
Zonosagitta bedoti (Beraneck, 1895)
Body opaque, firm and wider at the middle. Small head with conspicuous neck. On each side of
the head 6-8 thick, short, not strongly curved hooks, 8-10 anterior teeth and 18-22 thin, closely
packed posterior teeth. A short thin collarette can be present. The anterior fin starts in the
middle of the ventral ganglion. The anterior and inner part of the anterior fins is rayless. The
innermost part of the posterior fins is rayless. The seminal vesicles touch both the posterior and
caudal fins. When mature the ovaries reach the ventral ganglion. The ova are round, but can be
compressed in mature specimens, arranged in 3 rows. Length up to 15 mm.
Zonosagitta bedoti: A, maturity stage 1; B, stage 2; C, stage 3; D, stage 4 (From Alvario, 1967).
316
Zonosagitta pulchra (Doncaster, 1902)

Slender body, widest at its midpoint, firm and transparent. Small head. On either side of the
head 6-7 hooks, 10 anterior teeth and 15 posterior teeth. Well-developed collarette in the neck
region, extending one third of the distance to the ventral ganglion, also appearing in the seminal
vesicle region. Anterior fins start in the middle of the ventral ganglion, broadest at their
posterior part. The anterior and inner part of the anterior fin is rayless, or has a few rays, while
the posterior edge is rayed. The posterior fins are roundish, with the anterior and inner parts
rayless. In mature specimens the ovaries reach the ventral ganglion. The ova are roundish and
arranged in 2, sometimes 3 rows. The seminal vesicles are close to both the posterior and
caudal fins but separate from them. Length up to 24 mm.
Zonosagitta pulchra: A, maturity stage 1; B, stage 2; C, stage 3; D, stage 4 (From Alvario,

1967).
317
Family Eukrohniidae
Genus Eukrohnia
Characterised by having only one set of paired fins, which extend from the tail to the level of
the ventral ganglion, tail segment comparatively short, no gut diverticula and only one row of
teeth. In juveniles, the hooks on the head may have wide serrations.
Eukrohnia hamata (Mbius, 1875)
The body is firm, rigid and opaque. The head is small with a distinct neck, making the head
appear broader. The eyes are large with no pigment and there are 8-10 hooks with curved tips
on each side of the head. There may be as many as 26 teeth on each side of the head, in a single
row. There is no collarette or gut diverticula. The intestine characteristically often contains oil
droplets, which range in colour from bright orange to olive green. The single lateral fins are
broader at the level of the tail septum. Most of the fins are without rays, but they are present at
the posterior outer margins. The ovaries can reach up to the middle of the trunk. The ova are
small, round and arranged in four rows. The seminal vesicles are elongated and oval, situated
close to the posterior end of the paired fins and separate from the tail fin. Length up to 43 mm.
Eukrohnia hamata: A, maturity stage 1; B, stage 2; C, stage 3; D, stage 4 (From Alvario,

1967).
318
PHYLUM HEMICHORDATA
Hemichordates are wormlike animals living in tubes in the bottom mud or among rocks or
masses of plant material. At one time they were considered to belong to the same phylum as we
belong (Phylum Chordata), but now have a phylum of their own. Their casts, left exposed at
low tide, are a familiar sight in the tropics. Their common name is acorn worms and their
tornaria larvae can occur in plankton samples. The larvae are usually between 0.5 and 1 mm
long and have obvious bands of cilia.
Example of tornaria larvae (From Wickstead, 1965)
PHYLUM UROCHORDATA
Urochordates do not have a backbone but share our phylum, because during a stage in their life
cycle they have a primitive notochord. The y are represented in the plankton by three main
classes.
CLASS ASCIDIACEA
A common ascidacean is the sea squirt, which is typically found on the seabed or attached to
the bottom of ships. It has a larval stage called an ascidian tadpole (as it superficially resembles
a frog tadpole) which often occurs in plankton samples.
Ascidian tadpole larvae (from Wickstead, 1965).
319
CLASS LARVACEA
Larvaceans, also known as appendicularia, have an elongated, flat paper- like body of varying
width. They have a head which usually appears to be very loosely attached to the body and is
often broken off in samples. They secrete a jelly- like house inside which they live, but the
house and animal are generally sampled separately when they are caught in the plankton net.
Examples of appendicularia (From Wickstead, 1965).
CLASS THALIACEA
There are two main groups of thaliaceans. The doliolids are barrel-shaped and have complete
and separate circles of muscle-bands around the body e.g. Doliolum. They have a complex life
history involving three stages, apart from the larvae. One of the stages is sometimes found in a
disintegrating form (old nurse), where the individual becomes very flabby and the muscle band
widen till they can become continuo us in some species.
The second group, the salps, are generally larger than doliolids and have more jelly in their
bodies. In salps, some of the muscle bands join together at certain points and do not go
completely round the body e.g. Thalia. They have a less complex life cycle than dolioids. The
asexual individuals live singly and are called solitary. They bud off sexual individuals in a
chain, which stick together for some time, so are called the aggregate phase.
D
Examples of thaliaceans (A-C From Wickstead, 1965). A: Thalia democratica, aggregate form;
B: Thalia democratica, solitary form; C: Doliolum sp. D: Doliolum sp. old nurse stage (From
Fraser, 1947).
320
SUBPHYLUM CEPHALOCHORDATA
The only members of this group are the lancets, which were once called Amphioxus. A
common genus is Branchiostoma. They are rigid slender organisms, flattened side to side, with
segmented muscle bands and gill slits. They have a nerve cord along the whole body. They
generally live in sandy bottoms but often emerge into the plankton, especially the juveniles.
Examples of cephalochordate larvae A: Branchiostoma sp. larva (1.5 mm); B: Branchiostoma

sp. larva (2.8 mm); C: Branchiostoma sp. young adult (5 mm); D: Amphioxides sp. adult (9
mm). (From Wickstead, 1967).
321
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GUIDE TO THE

DEFRA Darwin Initiative Zooplankton Programme

Marine Biological Association of the United Kingdom

GUIDE TO THE COASTAL AND SURFACE ZOOPLANKTON

UK-DEFRA Darwin Initiative Project 162/09/004

Marine Biological Association of the United Kingdom

Phytoplankton and cyanobacteria

Examples of phytoplankton and cyanobacteria. A: Dinoflagellates; B: Diatoms; C:

Representative radiolaria (From Haeckel, 1862, 1867).

Representative Acantharians (From Trgouboff & Rose, 1957).

Examples of foraminferans (From Wickstead, 1965)

Examples of tintinnid shapes (From Daday, 1886)

A, planula larva; B, aracnactis larva (From Wickstead, 1965).

Ephyra larvae (From Wickstead, 1965)

Chrysaora hysoscella (Linnaeus, 1766)

Chrysaora hysoscella (Figure by J. Corbera, From Pags et al., 1992)

A, Structure of medusae; B, Details of umbrella margin (From Russell, 1953).

Actinula larvae (From Wickstead, 1965).

Euphysa aurata (Figure by J. Corbera, From Pags et al., 1992)

Zanclea costata. Different development stages (From Russell, 1953).

Cladonema radiatum. Different developmental stages (From Russell, 1953).

Turritopsis nutricula. (From Mayer, 1910).

Podocoryne spp. A, (From Mayer, 1910); B, (From Russell, 1953).

Bougainvillia macloviana Different development stages. A, (From Russell, 1953); B, (Figure by J.

Leukartiara octona (Figure by J. Corbera, From Pags et al., 1992).

Velella velella Colony. (Figure by J. Corbera, From Pags et al., 1992)

Clytia sp. (Figure by J. Corbera, From Pags et al., 1992).

Obelia sp. (Figure by J. Corbera, From Pags et al., 1992)

Phialella quadrata. A, B, different development stages; C, gonad and marginal tentacle;

Proboscidactyla stellata. (Figure by J. Corbera, From Pags et al., 1992)

Liriope tetraphylla (Figure by J. Corbera, From Pags et al., 1992)

Aglaura hemistoma (Figure by J. Corbera, From Pags et al., 1992)

Solmundella bitentaculata. (Figure by J. Corbera, From Pags et al., 1992)

Physalia physalis (Figure by J. Corbera,

Rosacea cymbiformis. (Figure by J. Corbera, From Pags et al., 1992)

General Diphydae structure, polygastric, stage, showing nectophores separated (Figures by J.

Sulculeolaria turgida A anterior nectophore; B, posterior nectophore (Figures by J. Corbera, From

Sulculeolaria chuni (Lens and van Riemsdijk, 1908)

Sulculeolaria chuni. A, anterior nectophore; B, posterior nectophore (Figures by J. Corbera, From

Diphyes chamissonis. Anterior nectophore

Diphyes dispar Chamisso and Eysenhardt, 1821

Diphyes dispar. A, anterior nectophore; B, posterior nectophore (Figure by J. Corbera,

Lensia subtilis (Chun, 1886).

Lensia subtiloides (Lens and van Riemsdijk, 1908).

Lensia hotspur Totton, 1941

Chelophyes appendiculata (Eschsoltz, 1829).

Chelophyes appendiculata. A, Anterior nectophore; B, Posterior nectophore (Figure by J. Corbera,

Chelophyes contorta. A, anterior nectophore; B, anterior nectophore side view; C, Posterior

Eudoxoides spiralis (Bigelow, 1911).

Eudoxoides mitra. A, anterior nectophore; B, posterior nectophore; C, eudoxid stage (Figures by J.

Abylopsis eschscholtzi. A, complete polygastric stage; B, eudoxid stage (Figures by J. Corbera,

Bassia bassensis (Quoy and Gaimard, (1833) 1834).

Examples of ctenophores (From Wickstead, 1965).

Typical Turbellarian shapes.

Pilidium larvae (From Wickstead, 1965).

Examples of rotifers (From Berzins, 1960a,b).

Chiton development stages. A, trochophore; B, later trochophore; C, juvenile; D, adult. Figures

Examples of Sepioidea larvae (From Muus 1963a)

Examples of the Genus Octopus (From Muus 1963d).