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24. Crustacea Cladocera

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274 Freshwater Invertebrates of the Malaysian Region

Crustacea: Cladocera
Marcelo Silva-Briano1 and Iskandar M. Mirabdullayev2
1
Universidad Autónoma de Aguascalientes. Centro de Ciencias Básicas. Departamento de Biología,
Laboratorio No. 1 de Ecología, Edificio no. 202, Av. Universidad No. 940, Ciudad Universitaria
Verdad. CP. 20100, Aguascalientes, Ags, Mexico
(Email: msilva@correo.uaa.mx)
2
Institute of Zoology, Niyazov st. 1, Tashkent 700095, Uzbekistan
(Email: imirabdullayev@yahoo.com)

INTRODUCTION
Cladocera are commonly known as ‘water fleas’. Most Cladocera are 0.2–3.0 mm
long. The body is not clearly segmented, and consists of two main parts – the head
and the trunk, the latter being covered by a bivalve carapace that ventrally gapes.
The carapace has a dorsal keel and there are often surface reticulations, striations
or other markings. The head bears two pairs of antennae and (usually) a ventral
rostrum, and is normally covered by a head shield (Fig. 1A). The head shield bearing
integumental pores is useful in taxonomy (Figs. 1H, 3E). There is a single compound
eye and sometimes an ocellus. The trunk bears 5–6 pairs of limbs. The abdomen
lacks appendages. The posterior part of abdomen is bent ventrally, forming the
postabdomen. The postabdomen has two strong terminal claws and two long seta
natatoriae on the dorsal side. In the dorsal part of the shell is a brood chamber,
usually containing eggs or embryos and closed posteriorly by a special abdominal
prominence.
The first pair of antennae (antennules) is inserted near the rostrum and is weakly
developed in females (Fig. 1A). The second pair of antennae is well developed and is
the main swimming organ. Each of the antennae consists of a massive basipodite and
two branches: an upper 3–4-segmented exopodite and a lower 3-segmented endopodite
bearing long feathered swimming setae. Externally, there is a hard ridge or fornix,
anterior to the base of each antennae. The mandibles are asymmetrical and they
bear numerous spines and prominences distally. The first pair of maxillae (maxillulae)
has a few short setae, whereas the second pair of maxillae has apparently disappeared
in most groups of Cladocera. Trunk limbs consist of outer (exopodite) and inner
(endopodite) lobes, and one additional outer epipodite. The second pair has a special
maxillary outgrowth near its base, the 3d and 4th pairs are especially large, and their
endopodites bear many long filter setae. The trunk limbs form a rather complex
pump and filter device, which follows a regular rhythmic cycle.
It has been suggested that the Cladocera are an artificial group containing
representatives of rather different phylogenetic origin (Fryer 1987; Negrea et al. 1999).
Crustacea: Cladocera 275

Figure 1. A – Daphnia pulex; B – Simocephalus mesorostris; C – Scapholeberis mucronata; D –

Ceriodaphnia cornuta; E – Moinodaphnia macleayi; F – Moina micrura; G – Bosmina longirostris;
H – Alona quadrangularis; I – Disparalona rostrata; J – Macrothrix spinosa; K – Alonella dadayi.
(Sources: A, C, E–G, I, K, modified after Pennak 1989; B, from Rakhmatullaeva and Mirabdullayev
1998; D, from Smirnov and Timms 1983; H, from Alonso 1996; J, after Sars 1888)
276 Freshwater Invertebrates of the Malaysian Region

GENERAL BIOLOGY
Cladocera occur everywhere from the Arctic region to the Antarctic continent, in
temperate and tropical latitudes, on isolated ocean islands and in high mountain water
bodies, in permanent and temporary waters. They penetrate into ground water and
caves and even inhabit moss-covered trunks and soils in extremely wet cloud forests.
The majority of Cladocera, however, inhabit fresh waters, but a number of them occur
in brackish and saline (up to 100 ‰ and more) habitats or acid environments such as
peat swamps. Some of them live along the shore or in the open water of oceans.
For most of the Cladocera, the antennae are the main organs of locomotion. In
some genera, such as Latona, single vigorous strokes of the antennae produce
powerful leaps. Daphnia and its relatives move by a series of hops, produced by
more rapid and less vigorous strokes. A few genera, including Holopedium and
Scapholeberis, habitually swim upside down. Species occurring on a substrate have
less frequent periods of active swimming and often use the antennae incidentally for
obtaining a purchase on vegetation and bottom debris. The postabdomen is also
useful for locomotion in bottom-inhabiting species. A cladoceran may literally kick
itself along, sometimes by quite respectable hops.
A few species have become remarkably adapted for living in the damp moss and
debris of tropical forest floors; they are unable to swim and only creep about (Frey
1980). A few species (chiefly Chydoridae) inhabit the interstices of stream gravel
deposits.
Complex movements of the highly setose thoracic legs produce a constant current
of water between the valves. These movements further serve to filter food particles
from the water and collect them in a median ventral groove at the base of the legs.
This stream of food is fed forward to the mouthparts where the particles may be
ground between the surfaces of the mandibles before these are taken into the mouth.
Algae and Protozoa have often been assumed to be the main foods, excluding
other materials, but it is now well known that organic detritus of all kinds, as well as
bacteria, are very important and commonly form the great bulk of ingested material.
It has also been successfully demonstrated that the feeding movements of the legs of
plankton species are so efficient that they separate some colloidal organic particles
from the water. It is generally believed that all organic particles of suitable size are
ingested without any selective mechanism. When undesirable material or large
tangles masses are introduced between the mandibles, they may be removed by
spines at the base of the first leg and then kicked out of the shell by the postabdomen.
Cladocera are an important element of freshwater ecosystems, often being the
dominant part of the zooplankton. They are the main food of almost all young
freshwater fishes. Harvested and dried or frozen Cladocera are used in aquaculture
as food for shrimps and fishes. Cladocera are also used in monitoring of water quality.
Crustacea: Cladocera 277

Figure 2. A – Diaphanosoma brachyurum.; B – Pseudosida bidentata; C – Latonopsis australis;

D – Daphnia similis, ephipial female; E – Ilyocryptus sp.; F – Grimaldina brazzai G – Guernella
raphaelis; H – Streblocerus serricaudata; I – Pleuroxus striatus; J – Chydorus sphaericus,
parthenogenetic female; K – Chydorus sphaericus, gamogenetic female; L – Ephemeroporus barroisi.
Scale lines = 0.5 mm. (Sources: A, B, I, from Pennak 1989; D, from Idris 1983; C, E–H, from Silva-
Briano 1998; J, K, from Alonso 1996; L, from Frey 1982)
278 Freshwater Invertebrates of the Malaysian Region

Life cycle
Reproduction is parthenogenetic (asexual) during the greater part of the year in
most habitats, and only female young are produced. The eggs are released via
oviducts into the brood chamber. Depending on the species and environmental
conditions, the number of eggs per clutch varies considerably – usually between 2
and 40. The parthenogenetic eggs undergo further development in the brood chamber
and hatch into young similar to the adult form. The parent then liberates them to the
outside by moving the postabdomen downward.
Sometimes sexual reproduction occurs. At that time males and gamogenic (sexual)
females (Fig. 2D; 2K) capable of fertilisation appear. After copulation females
produce 1–2 (rare more) sexual “resting” eggs. The fertilized eggs pass into the
brood chamber, the walls of which then become thickened and darkened to form an
ephipium. The ephipium can resist freezing and drying, and enable cladocerans to
survive in undesirable conditions and also to disperse.

REGIONAL TAXA
The cladoceran fauna of the Malaysian region is typical of tropical fresh waters.
Sixty-five species have been recorded from the region (Spandl 1925; Johnson 1962; Idris
and Fernando 1981; Idris 1983; Korovchinsky 1992; Dumont and Silva-Briano 2000).

KEY TO FAMILIES OF MALAYSIAN CLACOCERA

1. Dorsal ramus of second antenna bearing 8 or more setae; six pairs of trunk legs of similar
structure.. ................................................................................................................ SIDIDAE
- Dorsal ramus of second antenna bearing 5 or less setae; five or six pairs of trunk legs of different
structure .................................................................................................................................. 2
2. Dorsal ramus of antennae 4-segmented, ventral ramus 3-segmented ...................................... 3
- Dorsal and ventral rami 3-segmented ...................................................................................... 6

3. First antennae not mobile (Fig. 1A) ................................................................. DAPHNIIDAE

- First antennae mobile (Fig. 1E) ............................................................................................... 4

4. First antennae 2-segmented ................................................. ILYOCRYPTIDAE: Ilyocryptus

(Ilyocryptus is the only genus – Fig. 2E)
- First antennae 1-segmented .................................................................................................... 5
5. First antennae situated on antero-ventral end of head (Fig. 1J) ............. MACROTHRICIDAE
- First antennae situated on postero-ventral margin of the head, but not at the anterior end (Fig.
1E,F) ..................................................................................................................... MOINIDAE
6. First antennae fused with the rostrum, forming long proboscis-like structure (Fig. 1G) .............
........................................................................................................................ BOSMINIDAE
- First antennae not fused with the rostrum, and covered by head shield or rostrum ....................
...................................................................................................................... CHYDORIDAE
Crustacea: Cladocera 279

KEYS TO GENERA OF MALAYSIAN CLADOCERA

FAMILY DAPHNIIDAE
1. Valves with long, sharply pointed postero-dorsal spine; proximal to the postabdomen; 2 well
developed abdominal processes .............................................................................................. 2
- Valves without postero-dorsal spine; proximal to the postabdomen with only 1 abdominal process.. 3
2. Body size large (more than 2 mm); head broadly rounded without horn, cervical depression
absent; postero-dorsal corner of valves with a long spine (Figs. 1A, 2D) ................... Daphnia
- Body size small (less than 1 mm); head with or without horn-like process, cervical depression
present; postero-dorsal corner of valves with 2 short sharply pointed spines (Fig. 1D) .......
.......................................................................................................................... Ceriodaphnia

Figure 3. A – Dunhevedia crassa; B – Dadaya macrops; C – Leydigia acanthocercoides; D –

Graptoleberis testudinaria; E – Karualona karua, Up, detail of head pores; F – Kurzia longirostris; G
– Camptocercus australis; H – Acroperus harpae; I – Oxyurella singalensis; J – Euryalona orientalis;
K – Indialona globulosa. Scale lines = 0.1 mm. (Sources: A, from Alonso 1996; B–D, F from Thorp and
Covich 1991; E–K, from Smirnov 1974)
280 Freshwater Invertebrates of the Malaysian Region

3. Posterior margin of valves almost straight vertically, ventral margin straight horizontally with
spine on postero-ventral corner; preanal corner and anal groove of postabdomen evenly rounded
with 4 or 5 denticles (Fig. 1C) .......................................................................... Scapholeberis
- Posterio-dorsal and posterio-ventral margin of valves broadly rounded, posterior corner
projecting forming an acute angle; postabdomen with distinct and projecting preanal corner;
anal groove slightly concave with 5–10 long and strong denticles decreasing in size proximally
(Fig. 1B) ............................................................................................................. Simocephalus

FAMILY MOINIDAE
1. Body compressed, valves elliptical and completely covering the body; ocellus present; first
antennae long with one long lateral seta (Fig. 1E) ............................................. Moinodaphnia
- Body thick and heavy, valves somewhat rhomboid, not wholly covering the body; ocellus
absent; first antennae covered by short spines and long fine setules laterally (Fig. 1F) ... Moina

FAMILY SIDIDAE
1. Posterior and/or postero-ventral valve margins with long feathered and especially long naked
setae (Fig. 2C) ........................................................................................................ Latonopsis
- Posterior and postero-ventral valve margins without long setae; relatively short feathered setae
may be present only on antero-ventral margins ..................................................................... 2
2. Antennules short; postabdomen without anal teeth or with small solitary anal teeth (Fig. 2A)
........................................................................................................................ Diaphanosoma
- Antennules long; postabdomen with cluster of anal teeth (Fig. 2B) ...................... Pseudosida

FAMILY MACROTHRICIDAE
1. Head relatively small (less than 0.2 of the body size); ventral margin of valves not serrated;
first antennae slender and long (ratio of width-length less than 0.2); postabdominal claw long
with 2 basal spines, dorsal-distal margin with long and strong spines .................................. 2
- Head relatively large (more than 0.25 of the body size); ventral margin of valves serrated; first
antennae relatively shorter (ratio of width-length more than 0.2), postabdominal claw short
without basal spines, dorsal-distal margin without long spines ............................................ 3

2. Ocellus situated far from the eye, closer to the apex of rostrum; first antennae with lateral setae
near the base; dorsal-distal margin of postabdomen with 11–12 long and strong spines in
groups, dorsal margin with a series of fine setules (Fig. 2F) ................................. Grimaldina
- First antennae short and broad (ratio of width-length more than 0.4); ventral margin of valves
serrated, without setae; postabdomen with transverse rows of spinules, without anal spines or
anal denticles (Fig. 2G) ............................................................................................ Guernella
3. Body size less than 0.25 mm (the smallest macrothricid), head about 0.5 of the body size;
labral plate pointed postero-ventrally and with setules on the apex; ventral setae not arranged
in a group; first antennae armed with 4 long spine–like setae on the anterior surface (Fig. 2H)
............................................................................................................................. Streblocerus
- Body size more than 0.25 mm, head about 0.25 of the body size; labral plate straight or
concave and somethings with protuberance in the middle; ventral setae arranged in groups of
three; first antennae sometimes with spines on the surface (Fig. 1J) .................... Macrothrix
Crustacea: Cladocera 281

FAMILY CHYDORIDAE
1. Exopodite of 4th trunk legs bearing less than 7 setae; small head pores between main pores (Fig.
3E) ................................................................................................ Subfamily Chydorinae ... 2
- Exopodite of 4th trunk legs bearing 7 setae; small head pores laterally to main pores (Fig. 1H)
.................................................................................................................................................
Subfamily Aloninae .. 8
2. Ventral margin of valves with a series of setae attached marginally on entire length, head with
long and pointed rostrum (2.0–2.5 of length of first antennae) ............................................... 3
- Ventral margin of valves with a series of setae attached submarginally on the anterior and posterior
part of the margin, head with shorter rostrum (less than 1.5 of length of first antennae) ............ 5

3. Head shield with very long and pointed rostrum, postpore distance greater than interpore
distance, postabdomen slightly long, narrow and tapering distally (Fig. 2I) ................ Pleuroxus
- Head shield with moderately long rostrum, postpore distance less than interpore distance,
postabdomen short and slightly tapering distally ...................................................................... 4

4. Head shield fan-shaped with reticulations and longitudinal striations, postpore distance slightly
less than interpore distance; posterior margin convex, postero-ventral margin rounded, without
denticle; postabdomen slightly broad and rounded distally (Fig. 1I) ...................... Disparalona
- Head shield with fine striations, postpore distance much less then interpore distance; posterior
margin straight, postero-ventral corner with denticle or serration (except Alonella hamulatus);
postabdomen tapering slightly distally (Fig. 1K) .......................................................... Alonella
5. Body spherical in shape, dorsal margin shorter than maximum height ..................................... 6
- Body oval in shape, dorsal margin longer than maximum height, postero-ventral corner with
denticles .................................................................................................................................. 7
6. Body spherical in shape, dorsal margin shorter than maximum height, postero-ventral corner
without denticles, dorsal and ventral margin of postabdomen parallel or almost parallel (Fig.
2J,K) ......................................................................................................................... Chydorus
- Body spherical in shape, dorsal margin shorter than maximum height, postero-ventral corner
with denticles, dorsal and ventral margin of postabdomen parallel or almost parallel (Fig. 2L) ..
....................................................................................................................... Ephemeroporus
7. Eye rather small, ocellus smaller than the eye; first antennae not reaching the apex of rostrum;
head shield with 2 main pores and 2 lateral pores (Fig. 3A) ................................. Dunhevedia
- Eye rather large, ocellus elongated and about the same size as the eye; first antennae project far
beyond the apex of rostrum; head shied with only one head pore situated far from the posterior
margin (Fig. 3B) .......................................................................................................... Dadaya
8. Postabdomen short or broadly rounded ................................................................................. 9
- Postabdomen narrow and long ............................................................................................. 12
9. Postabdomen broadly rounded or almost semi-circle; anal denticles short and small, lateral
setae long and longer than anal denticles and projecting far beyond the dorsal margin; claw with
very short basal spine (less than 0.25 the basal diameter of the claw). (Fig. 3C) ........ Leydigia
- Postabdomen rather short, lateral setae fine and usually smaller than anal denticles and some-
times slightly projections beyond the dorsal margin; claw with relatively longer basal spine
(more than 0.5 the basal diameter of claw) .......................................................................... 10
282 Freshwater Invertebrates of the Malaysian Region

10. Head, dorsal and posterior margin evenly rounded (forming a semicircle shape) with indistinct
postero-dorsal corner; ventral margin straight; valves and head shield heavily reticulated (Fig.
3D) .................................................................................................................... Graptoleberis
- Not like this ............................................................................................................................ 11
11. Head and dorsal margin evenly rounded with distinct postero-dorsal corner; ventral margin corner
slightly convex; valves not heavily reticulated; with only faith longitudinal striations or plain (Fig.
1H) ................................................................................................................................. Alona
- Two main head pores, with two small lateral head pores and distal pseudopore, valve densely
reticulated; postero-ventral corner mostly with 2–5 spines, labrum helmet-shaped, with strong
keel and lateral indentation (Fig. 3E) ....................................................................... Karualona
12. Shell with keel, head shield with 3 connected main pores ...................................................... 13
- Shell without keel, head shield with 4 main pores (2 middle pores connected) or with only single
pore ....................................................................................................................................... 15
13. Only shell with keel; rostrum long and attenuated; postero-ventral margin of valves without
denticles (Fig. 3F) .......................................................................................................... Kurzia
- Both head and shell with keel; rostrum blunt, postero-ventral corner of valves usually with denticles
............................................................................................................................................. 14
14. Postabdomen very long, narrow and tapering distally, anal denticles and lateral setae numerous
and conspicuous (Fig. 3G) .................................................................................. Camptocercus
- Postabdomen slightly shorter with almost parallel dorsal and ventral margin, lateral setae
conspicuous, anal denticles very small (Fig. 3H) ...................................................... Acroperus

15. Head shield with 4 median pores (anterior and posterior pores isolated, 2 middle pores connected)
and 2 lateral pores; postabdominal claw with a long basal spine and 3 accessory spines at the
base (Fig. 3I) ............................................................................................................ Oxyurella
- Head shield with 1 head pore; postabdominal claw with a short basal spine without accessory
spines .................................................................................................................................... 16
16. Head shield broadly rounded posteriorly and with one head pore closer to the posterior edge;
labral plate rounded with nipple-like structure on the apex; postabdomen long with sharply pointed
anal denticles; lateral setae not projecting beyond the dorsal margin (Fig. 3J) ......... Euryalona
- Head shield slightly pointed posteriorly, head pores formed by 2 semicircular slits situated slightly
to the middle; labral plate serrated antero-ventrally with slightly pointed apex; postabdomen
slightly broader with small anal denticles; lateral setae projecting beyond the dorsal margin (Fig.
3K) ........................................................................................................................... Indialona

FAMILY BOSMINIDAE (see the figure in Idris, 1983)

In Malaysia and Singapore the only genus is Bosminopsis.

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