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Rotifera: Monogononta

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106 Freshwater Invertebrates of the Malaysian Region

Rotifera: Monogononta
Hendrik Segers
Royal Belgian Institute of Natural Sciences, Freshwater Biology,
Vautierstraat 29, B-1000 Brussels, Belgium
(Email: hendrik.segers@naturalsciences.be)

INTRODUCTION
The rotifers (Phylum Rotifera) are a diverse assemblage of pseudocoelomate,
primary bilaterally symmetrical worms. The taxon classically includes three groups,
the freshwater Monogononta and Bdelloidea, and the marine epizoic Seisonacea,
and is closely related to, amongst others, the parasitic Acanthocephala. Monogononta,
the largest of the three, comprises about 1450 species, distributed over 29 families
and 106 genera, worldwide (Segers 2002).
Rotifers are diagnosed by the presence of an anterior ciliary organ, the corona,
used in locomotion and collection of food, a specialized muscular masticatory
apparatus or mastax containing a complex set of jaw-like structures, the trophi, and
a tegumental skeletal lorica which, in many species, forms a stiff defensive cover. In
addition, all post-embryonic tissues are syncytial, without cell membranes separating
individual cells. They are eutelic, having a constant, and species-specific number of
nuclei ranging from 900 to 1000, for those taxa that have been studied (Nogrady et
al. 1993). Characteristic features for monogononts are the presence of typical trophi
types, unpaired gonads, and a reproductive cycle consisting of parthenogenetic phases
intermitted by short periods of sexual reproduction (heterogonic reproduction).
The body of monogonont rotifers (Fig. 1) typically consists of three main regions:
head, trunk and foot. Although these regions may be marked or subdivided by
transverse folds in the body wall, giving the appearance of joints, no true segmentation
is present. The head bears the corona and several types of sensory organs: tactile
cirri, chemoreceptive pores, eye(s), and a single or (rarely) double dorsal antenna.
The corona is a ciliary organ, which, in its simplest form, consists of a circumapical
ring of cilia surrounding an apical field without cilia, and a ciliated mouth field. In the
corona two bands of cilia, the anterior trochus and posterior cingulum can be
discerned. This basic corona pattern is commonly modified into diverse functional
types in conjunction with specific adaptations: in many creeping forms, the ventral
part of the corona is enlarged to facilitate gliding over surfaces, whereas in sessile
taxa the circumapical ring can be prolonged, eventually by the formation of lobes,
increasing the filtering potential. The head contains the brain, frontal or cervical
eyespots with associated pigment spots, and the retrocerebral organ (a gland of
Rotifera: Monogononta 107

unknown function). Most monogononts can retract their head and foot into the trunk
by the contraction of strong retractor muscles.
The trunk contains a spacious pseudocoelom in which are suspended most organs
of the digestive, reproductive, and excretory system, muscles and nerves. There are
no respiratory or circulatory systems. The excretory system consists of a paired

Figure 1. Scheme of the anatomy of a monogonont rotifer, Brachionus rubens Ehrenberg in dorsal
view. Abbreviations: b, bladder; co, corona; cr, corona retractor; da, dorsal antenna; e, egg; ey, eye; fg,
foot gland; fr, foot retractor; i, intestine; la, lateral antenna; ls, lorica spine; m, mastax; o, oesophagus;
ov, ovary; p, protonephridium; s, sensory cirrus; sg, stomach gland; st, stomach, to, toe; tr, trophi.
108 Freshwater Invertebrates of the Malaysian Region

protonephridial system with ducts opening into a cloaca, or a bladder. A number of

taxa (e.g. Cupelopagis, Collotheca) have excretophores, specialized amoeboid
cells accumulating excretion products. A pair of sensory papillae, the lateral anten-
nae, is present in the postero-dorsal region of the trunk. The foot is mostly retractile
or pseudosegmented, or reduced in many pelagic taxa. It may possess a pair of
cement glands, which open terminally in an adhesive disk, near the base or on the
tips of one or two toes, and produce a secretion for attaching the rotifer to a substra-
tum. This adhesion is mostly temporary, but permanent and strong in the case of
sessile animals; the latter are unable to reattach after being dislodged (Wallace 1980).
The tegument of trunk and foot secrete a gelatinous cuticle without skeletal func-
tion. This cuticle can be very thick and completely envelop the animal, especially
(but not only) in sessile animals.
The integument of monogonont rotifers contains a dense intracytoplasmic filament
layer, which serves as skeletal lamina. The development of this layer varies greatly in
different species, even within a single family or genus (e.g. Cephalodella). Species
are called illoricate (soft-bodied) or loricate, depending on the thickness of this lamina.
The skeletal layer serves for muscle attachment and defence, either passively or
actively. The arrangement of stiff plates in the tegument of loricate monogononts is
taxonomically significant. In many genera, easily discernable lorica plates are present
in the thoracic region, occasionally also on the head and/or foot. They may encircle
the body, forming telescopic pseudosegments. The skeletal lamina is thin and supple
at the articulation between plates or at joints. The tegument between lorica plates
may be deeply invaginated, forming lateral (e.g. Lecane) or dorsal sulci (e.g. Mytilina).
In some taxa (e.g. Brachionus, Testudinella), the thorax is completely covered by an
undivided lorica, forming a defensive barrel in which both head and foot can be
retracted. Many monogononts are provided with lorica projections; these are either
movable, and are used in escape-responses (e.g. Filinia, Polyarthra), or immobile,
and serve as passive defensive devises against predators (e.g. Brachionus, Keratella),
or regulate the animal’s floating capacity (e.g. Kelicottia).

GENERAL BIOLOGY
In general, monogononts are small animals. Most are in the 200–500 µm size range;
only a few reach over 1 mm. They constitute one of the main groups of primary
freshwater Metazoa, occurring in a wide range of habitats, from the open water of
lakes to the interstitial habitats of sandy beaches or the minute water reservoirs in
plants (phytotelmata) or empty gastropod shells. Some monogononts inhabit brackish
water or marine environments. About 50 species are exclusively marine, 20 of which
belong to a single genus, Synchaeta (Nogrady 1982). Most species are free-living or
sessile, solitary or colonial, while some are epizoic or parasitic.
Rotifera: Monogononta 109

Monogonont communities can be very diverse: tropical lakes may contain over
210 species of monogonont rotifer (Dumont and Segers 1996), with densities commonly
reaching up to about 1000 individuals per litre. Under natural conditions, densities as
high as 12,000 (sewage ponds: Seaman et al 1986) or even more than 100,000
(soda-water, Chad: Iltis and Riou-Duvat 1971) individuals per litre have been recorded.
In mass aquaculture systems, densities of 50,000 to 500,000 individuals per litre are
routinely attained (Lubzens 1987).
Many species play an important role in lacustrine food webs because they have a
rapid turnover rate and metabolism. They feed largely on detritus and bacteria be-
sides algae, and are important prey for small carnivores such as copepods, insects
and fish larvae. Most monogononts are filter feeders, or feed by browsing the
bacterial film on substrata. Food particles are directed to the mouth by the
metachronous beating of the coronal cilia. Many, however, are active predators or
feed on large algae. They either completely engulf their prey or suck out their contents.
Food is transported to a muscular masticatory apparatus, the mastax, via a short,
ciliated pharynx. This mastax contains hard, jaw-like structures, the trophi, and
associated muscles. The trophi are one of the most important features used in the
classification and identification of monogononts. From there, particles are transported
through a short oesophagus to a stomach, intestine and anus situated dorsally of the
foot. The terminal part of the intestine is a cloaca into which the oviducts and bladder
or protonephridial ducts open. Two sets of glands are associated with the digestive
system, the mastax or salivary glands, and the gastric glands. In a number of taxa, the
stomach is a blind sac, and food remnants are extruded by the trophi (Asplanchna)
or accumulated in defaecation bodies in the trunk (Ascomorpha). Endosymbiotic
zoochlorellae occur in several taxa (e.g. Encentrum, Ascomorpha).
Many monogononts are morphologically variable, mostly as a consequence of
phenotypic plasticity. Seasonal changes in phenotype (cyclomorphosis) in successive
generations are related to changes in physical, chemical or biological features of the
environment. Predator-induced polymorphism occurs in many species, especially
Brachionus and Keratella, and affects presence and length of lorica spines in response
to the presence of predators in the environment. When Brachionus calyciflorus
were cultured in media that previously held the predatory species Asplanchna,
offspring of the former developed lorica spines, which deterred predation by
Asplanchna (Gilbert 1966; 1967). Further experiments revealed that predator-
produced kaeromones induce spine development in other Brachionidae as well,
although some genetic variation is evident (Gilbert 1999, 2001). Additional phenotypic
variation can result from differences in the stage of population cycle. The first
generation individuals of Polyarthra and some Filinia hatching from resting eggs
lack the characteristic paddles or jumping bristles of their parthenogenetic conspecifics.
110 Freshwater Invertebrates of the Malaysian Region

Life cycle
Monogononts are cyclic parthenogens. Population increase results mostly from
asexual reproduction, which prevails during the life cycle. Sexual reproduction is
initiated under the influence of certain environmental or endogenous clues (e.g.
changes in photoperiod, population density and food availability). Mictic females
that are not fertilized within a short time after birth (four hours in Brachionus
plicatilis: Snell and Childress 1987), will produce haploid eggs developing into males.
Fertilized females produce a diploid, encysted diapausing embryo, called a resting
egg. In many species, the resting egg is retained within the female, which dies after
its production (e.g. many Filinia, Lecane). In other species, however, several resting
eggs or parthenogenetic as well as sexual (male or resting) eggs can be produced
(amphoteric females). In addition, pseudosexual eggs, diapausing eggs produced via
parthenogenesis, have been reported (Gilbert and Schreiber 1998). Resting eggs
typically possess a thick, often characteristically sculptured wall (Fig. 2). They are
very resistant, and can remain vital for long periods (up to 65 years: Kotani et al.
2001) of adverse environmental conditions. After receiving the proper hatching
stimulus, a single amictic female will hatch from the resting egg, and this single
specimen can initiate a new population through parthenogenesis. Resting eggs
can be dispersed over large distances by wind or waterfowl, and may be responsible
for the wide distribution of many monogonont taxa.
Monogononts have a single gonad, a feature which is reflected in the name of
the taxon (Greek: monos, single; gonos, seed). They are generally oviparous, and
either carry their eggs (e.g. Brachionus, Pompholyx), attach them to a substratum
(many benthic forms) or release them into the plankton (e.g. Notholca). Some
species are ovoviviparous or even viviparous. Males are generally much smaller
than females and occur only during the sexual phase of the life cycle. They have a
short life span, are easily overlooked and, consequently, are unknown in many species.
Males are functionally reduced; in one species (Trochosphaera solstitialis) the
male is not released but lives within its mother (McCullough and Lee 1980).

Trophi morphology
The trophi are a complex set of extracellular cuticular sclerites. Each trophi element
is made up of tubular structures with a dense osmiophilic core embedded in a matrix.
They are formed by a number of different epithelial cells, which can be appreciated
in the structure of the rami and manubria containing compartments. The mastax
and associated trophi are highly adaptive, and structurally and functionally complex
in conjunction to food type and feeding habits. The primary function of the mastax is
to grind food particles, however, common adaptations are for grasping or sucking in
complete food particles or the contents of large prey. Basically, the trophi consists of
five sets of elements (Fig. 3). An unpaired median fulcrum connects two rami at their
Rotifera: Monogononta 111

joint, and serves as insertion for abductor muscles for the rami. The fulcrum is
mostly rectangular in lateral view, but can be elongate, and/or provided with a basal
plate. The rami are hollow, more or less triangular structures. Their inner margin is
provided with numerous elongate elements, the rami scleropillii, which may be fused
forming a reinforced ridge and/or a series of blunt or sharp tooth-like projections.
Commonly, the rami are provided with antero-lateral projections, the alulae. Fulcrum
and rami are joined in a functional unit, the incus (Latin, anvil). The unci are plates
formed by connection of a variable number of teeth into one rigid structure. The

Figure 2. Some monogonont resting eggs. A–C superorder Gnesiotrocha, order Flosculariacea, A –
Conochilus natans (Seligo), B – Hexarthra mira (Hudson), C – Filinia longiseta (Ehrenberg); D, E, F
superorder Pseudotrocha, order Ploima, D – Asplanchna sieboldi (Leydig), E – Brachionus durgae
Dhanapathi, F – Keratella valga (Ehrenberg) (scanning electron microscopic photographs. Scale
bars = 10 µm).
112 Freshwater Invertebrates of the Malaysian Region

teeth are mostly unequal, with the ventral ones the largest, and consist of a shaft and
an arrow-shaped head. Minute subuncinal teeth are mostly present, and are situated
under the unci plate. The manubria are crescent-shaped and compartmented supports
of the unci with (Ploimida) or without (Gnesiotrocha) an elongate shaft. Unci and
manubria together form the malleus (Latin, hammer) of the trophi. Diverse accessory
sclerites occur in monogononts (e.g. Fig. 3I, J).
The trophi can be distinguished in almost any female monogonont, but are
completely or strongly reduced in most males. The basic pattern is considerably
modified in the different families or species according to their mode of life and their
feeding habits. Seven trophi types are recognized in monogononts:
- Malleoramate: (Fig. 3A, B) Fulcrum short, rami more or less triangular, flat; unci teeth numerous,
occasionally resembling striations; manubria crescent-shaped, without shaft.
- Uncinate: (Fig. 3C) Similar to the malleoramate type, but all trophi elements except the unci
strongly reduced. Unci teeth few (2–5 teeth per uncus), elongate and curved supporting rods for
the mastax. In Atrochidae and Collothecidae only.
- Malleate: (Fig. 3D, E) Similar to the malleoramate type: fulcrum short, rami more or less
triangular or lyriform, in a straight angle with the fulcrum. The unci plates have several teeth, but
generally less than in the malleoramate type (4–12 teeth), and the connection between the teeth
is more strongly developed. The manubria are provided with a shaft, which is typically fairly
short (more elongate in the submalleate trophi of Lecanidae and some Proalidae).
- Virgate: (Fig. 3F–H) Fulcrum elongate, rami bent towards dorsally, unci with few teeth or re-
duced, and manubria mostly with elongate shafts. The virgate trophi type is the most variable of
all trophi types; occasionally it is asymmetrical (e.g. Trichocerca: Fig. 3H). Virgate trophi
occasionally consist of thin, hence hardly discernable elements (e.g. many Synchaetidae).
- Cardate: (Fig. 3I) Fulcrum short, rami lyriform, manubria with a shaft and a characteristic,
additional ventral projection. Accessory trophi elements are mostly present, and numerous.
Restricted to Lindiidae.
- Forcipate: (Fig. 3J, K) Fulcrum mostly short, rami elongate, long pincers usually armed with
anterior and median teeth. Unci strong, but with a single or few teeth only, dagger- or sword-
shaped. Manubria long, thin, often with intramallei (Fig. 3J). In Dicranophoridae and Ituridae.
- Incudate: (Fig. 3L) Fulcrum short, rami elongate, pincer-shaped. Manubria and unci strongly
reduced. In Asplanchnidae.
A few monogonont taxa have trophi that can hardly be placed in any of these types
(e.g. Tetrasiphon).

REGIONAL TAXA
Until recently, monogonont rotifer taxa were considered cosmopolitan or, at least,
potentially cosmopolitan. Although this is probably correct for many taxa, recent
studies reveal an increasing number of species with geographically circumscribed
areas, especially in littoral or benthonic taxa (Segers 1996). However, strong
indications exist that cryptic diversity is high in this group (e.g. see Snell 1989). This,
in conjunction with poor taxonomic resolution, and fuzzy taxonomy (Segers 1998)
indicates that existing records should be interpreted with caution.
Rotifera: Monogononta 113

At present, some 220 monogonont taxa are on record for Malaysia, and c. 170
from Singapore (Sudzuki 1989; Segers 2001). These numbers are relatively high,
but result from a few major faunistic contributions (Karunakaran and Johnson 1978;
Fernando and Zankai 1981; Sudzuki 1989; Green 1995). On the other hand, a
comparison with the more intensively studied Thai fauna (320+ species on record:
see Sanoamuang et al. 2001) reveals that these numbers are nowhere near a reliable
estimate of the total diversity of the Malaysian monogonont fauna. Analysis of the
data is further hampered by the low reliability of non-illustrated records, and the fact
that the only contribution that includes records from Sabah and Sarawak (Fernando
and Zankai 1981), does not separate records from Borneo and Peninsular Malaysia.
The Malaysian rotifer fauna is particularly rich in species of Lecane, Brachionus
and Trichocerca. Whereas a dominance of littoral-periphytonic Lecane and
Trichocerca species is well-documented for natural, slightly acidic water bodies in
tropical regions, the high proportion of Brachionus species on record results from a
research focus on freshwater habitats with fisheries prospects: Brachionus is
particularly abundant in highly productive waters, and the successful use of B.
plicatilis-group as live food for fish fry in marine pisciculture has drawn intense
research interest towards the diversity of this genus. Predominantly cold-water genera
like Ploesoma and Synchaeta are evidently rare in Malaysia, but can be expected to
occur at high altitudes. Surprisingly, none of several Oriental endemic monogononts
(Segers 2001) have so far been recorded from Malaysia, although this is the case for
Thailand, Indonesia, and Singapore.

Identification
Rotifer identification is tedious because of the small size of most species, and the
requirement to examine trophi morphology. In most illoricate species, only living
specimens can be identified, as they will contract into an unidentifiable mass upon
fixation. Some, however, may still be recognizable by their trophi. Studying live
rotifers can be difficult, especially in the case of fast-moving species, and it may be
necessary to immobilize the animals by compressing them with the cover glass.
Alternatively, narcotics can be used to immobilize agile specimens. Several drugs
work well for this (Nogrady and Rowe 1993), but no single anaesthetic has been
shown to be universally effective. A convenient alternative is to slowly add ethyl
alcohol (96%, not denaturated) to a sample. This method works rather well with
Notommatidae. The hot-water fixation technique (Edmondson 1959: fast addition of
boiling water to a petri-dish containing the sample) gives satisfying results with some
taxa, notably collothecids but also bdelloids. The identification of most loricate
monogononts, on the other hand, relies on lorica morphology of well-contracted
specimens. Such material is best obtained by fixation, and preservation in 4%
formaldehyde or formol.
114 Freshwater Invertebrates of the Malaysian Region

Identification of monogonont specimens mostly requires microscopic examination

under the highest available magnification (1000x) using immersion oil, but it may be
useful to check specimens under 400x magnification without a cover slide in order to
assess their cross section. Reliable identification of many, especially illoricate,
monogononts requires the examination of trophi morphology. In the case of large
specimens or for routine identification light microscopy can suffice, but for some small
species or taxonomic studies scanning electron microscopy (S.E.M.) may be required.
For light microscopy, animals are transferred to a slide containing only a minute amount
of liquid. After examination of the complete animal, its trophi can be isolated by adding
a drop of commercial sodium hypochloride (NaOCl, ‘bleach’), KOH or NaOH (4%)
near the edge of the cover slide. Through capillarity the solution will spread under the
cover slide and clear out the trophi, which can then be studied. For the preparation of
trophi for S.E.M. examination, the following method can be used:
- A small number of clean (rinsed in distilled water) specimens (3–10) are placed on a clean
(degreased in alcohol, rinsed in distilled water and dried) cover slip or circular cover slip, placed
on a microscope slide. A minute quantity of glycerol placed between cover slip and slide is used
to prevent movement of the cover slip during the preparation process.
- By addition of a small drop of sodium hypochloride the specimens are dissolved. This process
will last, depending on the species and the concentration of the reagent, from 5 to 20 minutes. It
may be necessary to remove the trophi from the lorica, by gently squeezing the lorica with a fine
needle (e.g. Tungsten needles sharpened by electrolysis in a KOH solution) after most soft
tissues are dissolved.
- As soon as the trophi are clean, the remaining reagent is removed by using the capillary suction
of an ultrafine pipette, and a minute quantity of distilled water is added.
- Trophi are rinsed by adding, and subsequently removing distilled water. This is done 3–6 times,
until no crystal formation can be observed anymore when some of the removed rinsing water is
dried on a different slide. When necessary, the orientation of trophi can be changed by moving a
fine needle near the trophi.
- The preparation is dried by evaporation, and the trophi will automatically stick to the cover slip.
The cover slip can then be transferred to a S.E.M. stub, sputter-coated and examined.

This method can also be used to prepare trophi for inclusion in a permanent micro-
scope slide, in which case the cover slip is inverted on some mounting medium. Critical
point drying is not necessary for the preparation of trophi for S.E.M.; it is, however,
required when complete monogononts are prepared, even in loricate species.
As many monogonont rotifers are widely distributed, most identification works
treat all extant species of the taxa considered, and can therefore be used to identify
Malaysian monogononts. A classic identification guide is the German book by Koste
(1978), which is now dated. Recently, a number of English identification guides
treating selected taxa have become available (De Smet 1996; De Smet and Pourriot
1997; Nogrady et al. 1993, 1995; Nogrady and Segers 2002; Segers 1995), and
more are in preparation.
Rotifera: Monogononta 115

Figure 3. Trophi types in Monogononta. A, B – malleoramate type; C – uncinate type; D, E – malleate

type; F, G, H – virgate type; I – cardate type; J, K – forcipate type; L – incudate type. a, alulus; e,
ephipharynx; f, fulcrum; i: intramalleus; m, manubrium; ol, oral lamella; pi, pseudointromalleus; pm,
pseudomanubrium; r, ramus; u, uncus. (A, B, C, D frontal, E, I, J dorsal, F lateral, G, H, K ventral view.
Scale bars: A–F, H, I, K, L =10µm; G = 5µm; J = 1µm).
S.E.M. photographs. A – Sinantherina semibullata (Thorpe); B – Conochilus hippocrepis (Schrank);
C – Cupelopagis vorax (Leidy); D – Brachionus calyciflorus Pallas; E – Proales fallaciosa Wulfert;
F – Cephalodella gibba (Ehrenberg); G – C. ventripes (Dixon-Nutall); H – Trichocerca chattoni (de
Beauchamp); I – Lindia torulosa Dujardin; J – Encentrum longidens Donner; K – Dicranophorus
robustus Harring and Myers; L – Asplanchna brightwelli Gosse.
116 Freshwater Invertebrates of the Malaysian Region

KEY TO FAMILIES
Sixty genera and 26 families have been recorded from the Malaysian region, but
many other taxa (e.g. Microcodon clavus, monotype of Microcodonidae;
Trochosphaera) have been found in Southeast Asia and can therefore be expected to
occur in Malaysia. For this reason, the following key includes all extant families of
monogonont rotifers, except the North American endemic Birgeidae. Due to the high
morphological diversity of many monogonont families, an exclusive family key
appears unpractical. To remedy this, characteristic genera are included in the following
key. An asterisk indicates taxa, which are not yet recorded from Malaysia, but can be
expected there.
1. Trophi malleoramate or uncinate; foot, when present, without toes ......................................... 2
- Trophi otherwise; foot, when present, with toes ...................................................................... 8
2. Coronal ciliae short, trophi malleoramate ................................................................................. 3
- Coronal ciliae, when present, modified to elongate setae; anterior region a wide infundibulum;
trophi uncinate ......................................................................................................................... 7
3. Body conical with six thick, arm-like and setose appendages .................... HEXARTHRIIDAE
(One genus of common pelagic monogononts: Hexarthra)
- Body without such appendages or with bristle-like setae ......................................................... 4
4. Body loricate, more or less dorso-ventrally flattened ............................. TESTUDINELLIDAE
(Three genera: the monotypic and Oriental endemic *Anchitestudinella – A. mekongensis Berzins;
pelagic Pompholyx; and most species-rich, predominantly littoral Testudinella)
- Body illoricate, animals occasionally inhabiting a gelatinous matrix or stiff tube ...................... 5
5. Foot absent ..................................................................................... TROCHOSPHAERIDAE
(Three genera: common, pelagic Filinia; pelagic Horaella, and littoral *Trochosphaera)
- Foot present, retracted in preserved specimens ...................................................................... 6
6. Right unci with interdigitating accessory teeth proximally (Fig. 3B); corona round, horseshoe or
U-shaped. Adult females free-swimming, solitary or in small to large colonies (> 150 individuals)
................................................................................................................... CONOCHILIDAE
(Two genera including the common, pelagic Conochilus)
- Trophi with symmetrical unci or right unci without accessory teeth; corona round, heart-shaped,
elliptical or with lobes. Adult females sessile, rarely building large, free-floating colonies ......
............................................................................................................... FLOSCULARIIDAE
(Seven genera of solitary or colonial, sessile –*Beauchampia; Floscularia; Limnias; *Octotrocha,
and Ptygura, sessile or pelagic – Lacinularia; and Sinantherina – animals)

7. Corona with lobes, tentacles or points carrying long setae ...................... COLLOTHECIDAE
(Two genera of mostly sessile monogononts: Collotheca, Stephanoceros)
- Corona funnel-shaped without ciliae or setae ................................................. ATROCHIDAE
(Three genera of sessile monogononts including *Acyclus, and Cupelopagis)

8. Trophi cardate, animals illoricate ......................................................................... LINDIIDAE

(One genus of littoral animals: Lindia)
- Trophi otherwise, or animals loricate ................................................................................... 9
Rotifera: Monogononta 117

9. Trophi forcipate, animals illoricate ..................................................................................... 10

- Trophi otherwise, or animals loricate .................................................................................. 11
10. Trophi with beak-shaped allulae, not protrosible; stomach and intestine mostly greenish, filled
with zoochlorellae ................................................................................................ ITURIDAE
(One genus of littoral animals: Itura)
- Trophi without allulae or with differently shaped allulae; stomach and intestine rarely with
zoochlorellae .................................................................................... DICRANOPHORIDAE
(A diverse and species-rich family containing 19 genera of mostly free-living, creeping and
crawling freshwater monogononts; some species are brackish water or marine, some are
interstitial, epizoic or parasitic – *Albertia and *Balatro. Genera recorded from the Oriental
region are Dicranophoroides, Dicranophorus, Encentrum, *Paradicranophorus,
*Parencentrum, and Wierzejskiella. The endoparasitic Claria segmentata – monotype of
family Clariaidae – described from Vietnam, will key out here)

11. Trophi incudate. Animals saccate, illoricate, mostly without foot, and/or intestine and anus
................................................................................................................ APLANCHNIDAE
(3 genera: the common, pelagic predatory Asplanchna; littoral predatory Asplanchnopus; and rare
littoral Harringia)
- Trophi otherwise, or animals loricate .................................................................................... 12
12. Both foot and toes elongate, together longer than the body ................................................... 13
- Foot less elongate, toes may be long ..................................................................................... 14
13. Trophi virgate ................................................................................................. SCARIDIIDAE
(One genus of common, littoral monogononts: Scaridium)
- Trophi malleate .............................................................................................. Beauchampiella
(Monotypic genus (single species: B. eudactylota (Gosse, 1886) of family Euchlanidae)

14. Body with numerous, feather- or leaf-shaped movable projections; body shape more or less
rectangular; trophi virgate ....................................................................................... Polyarthra
(Common, pelagic animals. One of four genera in Synchaetidae)
- Body without such projections ............................................................................................... 15
15. Body loricate ......................................................................................................................... 16
- Body illoricate ....................................................................................................................... 28
(It may be difficult to assess this feature in some soft-loricate animals, especially when studying live
animals. When in doubt, check for the presence of lorica projections (always absent in illoricate
animals, occasionally present in loricate taxa), or by looking at the cross section: sulci (longitudinal
furrows) can be seen even when the lorica is weak. Many typically loricate genera like Brachionus
and Lecane contain illoricate species, and also the opposite holds (e.g. Cephalodella)).
16. Lorica with ventral, dorsal and/or lateral longitudinal sulcus or sulci: lorica consists of several
separate plates ..................................................................................................................... 17
- Lorica without longitudinal sulci ............................................................................................ 23
17. Sulci laterally only; lorica consists of a ventral and a dorsal plate (a sharp dorsal keel may be
present) ................................................................................................................................ 18
- Sulci not restricted to lateral location ..................................................................................... 21
18. No foot ................................................................................................................................. 19
- Foot and toes present (occasionally indistinct in contracted specimens) ........................... 20
118 Freshwater Invertebrates of the Malaysian Region

19. Trophi malleate ........................................................................... BRACHIONIDAE (partly)

(Three of seven genera: common pelagic Anuraeopsis; rare, cold-water pelagic *Kellicottia;
and very common, species-rich, littoral, and pelagic Keratella)
- Trophi virgate ...................................................................................................... Ascomorpha
(Common, littoral. One of two genera of Gastropodidae)

20. Foot short, inserted ventrally, on the ventral plate ........................................... LECANIDAE
(A single, species-rich – 42 taxa recorded from Malaysia by Sudzuki 1989 – genus of very
common, littoral, benthic, and interstitial monogononts: Lecane)
- Foot short or elongate, inserted caudally; foot and toes project from between ventral and
dorsal plate ..................................................................................... EUCHLANIDAE (partly)
(Three of five genera: rare, littoral Dipleuchlanis; common, littoral and occasionally pelagic
Euchlanis; rare *Tripleuchlanis. The rare, monotypic genus *Diplois, characterized by an additional
dorsal longitudinal sulcus, has been recorded from Indonesia)
21. Lorica with a distinct dorsal sulcus .............................................................................. Mytilina
(Common, littoral. One of two genera of Mytilinidae. Loricate species of Cephalodella will key out here)
- Lorica with a ventral sulcus .................................................................................................. 22
22. Trophi malleate, lorica mostly smooth .......................................................... LEPADELLIDAE
(Two of four genera: common littoral, and interstitial Colurella, rare littoral Paracolurella)
- Trophi virgate, lorica ornamented .............................................................................. Ploesoma
(Rare, generally cold-water. One of four genera of Synchaetidae)

23. Head shield present (retractable in Lepadella) ........................................... LEPADELLIDAE

(Two of four genera: common, species-rich, and littoral Lepadella; littoral Squatinella)
- No head shield ...................................................................................................................... 24
24. Trophi virgate ........................................................................................................................ 25
- Trophi malleate ..................................................................................................................... 26
25. Foot annulated, projects through a ventral aperture in the lorica ............................... Gastropus
(Uncommon. One of two genera of Gastropodidae)
- Foot inserted terminally, consisting of a single, short foot pseudosegment bearing elongate,
occasionally curved and/or asymmetrical spine-like toes ........................ TRICHOCERCIDAE
(Three genera, including the common, species-rich and ecologically diverse Trichocerca; the rare,
littoral or interstitial Elosa has been recorded from Singapore)

26. Lorica fairly homogeneous, relatively flexible, covering head, trunk and foot; toes occasionally
fairly long ................................................................................................... TRICHOTRIIDAE
(Three genera of littoral monogononts, including the common Macrochaetus, and Trichotria).
- Lorica covers trunk only, foot projects through a ventral or distal aperture ........................... 27
27. Lorica broader than high ............................................................................ BRACHIONIDAE
(Four of seven genera: the very common, species-rich and variable Brachionus, very common Plationus
patulus, and common benthic Platyias. One species of the cold-water *Notholca, N. striata, has been
recorded from Indonesia. Loricate species of Epiphanidae, e.g. Epiphanes brachionus, will key out here)
- Lorica with a dorsal keel ...................................................................................... Lophocharis
(Uncommon, littoral. One of two genera of Mytilinidae)

28. Trophi unique: fulcrum short, rami elongate, bent to dorsally, unci and manubria weakly developed.
Stomach filled with large unicellular algae. Vitellarium elongate with 20–30 nuclei arranged in a
row, wound around the intestine. A large, transparent species ................ TETRASIPHONIDAE
(monotypic family, with a single, periphytonic species: Tetrasiphon hydrocora)
Rotifera: Monogononta 119

- Not this combination of characters ...................................................................................... 29

29. Trophi virgate ...................................................................................................................... 30
- Trophi malleate .................................................................................................................... 32
30. With purple plate anterior to the mastax; corona somewhat circular. Foot pseudosegmented,
with a single toe .............................................................................. *MICROCODONIDAE
(Monotypic family, single, littoral species: Microcodon clavus)
- Not this combination of characters ...................................................................................... 31
31. Corona apical, consisting of isolated groups of ciliae, with four prominent sensory setae.
Mostly free-swimming, pelagic species .................................................................. Synchaeta
(Common, pelagic but predominantly cold-water; few tropical species known to date. Freshwater,
brackish water or marine. One of four genera of Synchaetidae)
- Corona ventral, well developed. Mostly creeping or crawling, littoral species .......................
................................................................................................................. NOTOMMATIDAE
(A very diverse and species-rich family containing 17 genera of mostly free-living, creeping and
crawling freshwater monogononts, including the largest rotifer genus, Cephalodella – c. 190 species.
Other common genera are, Eosphora; Monommata; Notommata; and Taphrocampa. Also
Enteroplea; Eothinia; *Pleurotrocha; *Resticula may be found)

32. Mouth situated at the end of a wide vestibulum; corona apical, mostly with specialized ciliae ...
...................................................................................................................... EPIPHANIDAE
(Five genera, including the common Epiphanes and Proalides – recorded as Liliferotrocha. The
monotypic *Cyrtonia – C. tuba – and *Microcodides are also expected to occur in the region).
- Mouth superficial; corona oblique ..................................................................... PROALIDAE
(Four genera of littoral and interstitial taxa: Bryceella; Proales; *Proalinopsis; *Wulfertia. Illoricate
Lecane species will key out here, and can be identified by the foot consisting of a single, short
pseudosegment only)

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