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Insecta: Odonata
Freshwater Invertebrates of the Malaysian Region 409

Insecta: Odonata
Albert G. Orr1, Stephen G. Butler2, Matti Hämäläinen3
and Robert G. Kemp4
1.
Cooperative Research Centre for Tropical Rainforest Ecology and Management,
Environmental Sciences, Griffith University, Nathan, Queensland 4111, Australia
(Email: agorr@universal.net.au)
2.
Red Willow, All Stretton, Shropshire, SY6 6HN, United Kingdom
(Email: sgbutler15@btopenworld.com)
3.
Department of Applied Biology, P.O. Box 27, FI-00014 University of Helsinki, Finland
(Email: matti.hamalainen@helsinki.fi)
4.
Tree Tops, 5 Maltings Close, Ackleton, Wolverhampton, WV6 7WB, United Kingdom
(Email: 10063.503@compuserve.com)

INTRODUCTION
With over 5000 described species worldwide, the Odonata is, after the Trichoptera,
the second largest insect order with an obligatory aquatic stage in the life history.
Only very rarely are the larvae secondarily terrestrial. Like many invertebrate groups
they are especially well represented in the tropics, about 500 described species
occurring in Sundaland. Of these, 350 are found within the territorial limits of
Malaysia, including species which are among the largest and smallest of their order in
the world.
Odonates are a very primitive and ancient group. Fossil wings of extant families
are known from the Triassic. Gigantic Protodonata flew in the Carboniferous. Today
two suborders are known, Anisoptera (unequal wings) or true dragonflies, and
Zygoptera (equal wings) often referred to as damselflies. Two relic Epiophlebia
species from the Himalayas and Japan, formerly included in the ancient suborder
Anisozygoptera, are at present placed in Anisoptera. In Malaysia Anisoptera and
Zygoptera are about equally represented. The fauna is quite well known, owing to
pioneering studies by F.F. Laidlaw and M.A. Lieftinck especially, but there remains
considerable scope for new discoveries, especially in Borneo. The larval stages remain
poorly known.
All odonates are predators, both as larvae and adults. Being near the top of the
food chain they are mostly relatively large insects occurring at only moderate
densities. They are of little economic significance. Both adults and larvae undoubtedly
prey extensively on mosquitoes, but it is not established whether this can affect the
incidence of arthropod-borne diseases in human populations. Occasionally in
Southeast Asia they are used as food and the larvae have been implicated in the
transmission of trematode parasites. However, perhaps most significantly, odonates
410 Freshwater Invertebrates of the Malaysian Region

are immensely interesting in their behaviour, ecology and zoogeography. They also
have considerable potential as indicators of environmental disturbance following
logging or pollution. This includes both stages of the life history, some groups being
more readily sampled as larvae, others as adults.

Figure 1. General view of A – an adult zygopteran (Coeliccia macrostigma – Platycnemididae) and

B – an anisopteran (Hemicordulia tenera – Corduliidae) showing principal features.
Insecta: Odonata 411

Structure of adults
Generalized adult structure is shown in Figure 1. The head is equipped with biting
mouthparts and bears very well developed compound eyes affording excellent, almost
360º vision. On top of the head are three small ocelli, or primitive eyes, arranged in
a triangle in Zygoptera. The antennae are very short, usually 6–7 segmented, with
the distal 4–5 segments forming an extremely thin flagellum. The thorax is divided
into a short narrow prothorax, bearing the front pair of legs, and a large synthorax,
consisting of the fused second and third thoracic segments (meso- and metathorax),
bearing the middle and hind pairs of legs and the wings. The prothorax, though
inconspicuous, is often highly modified in Zygoptera, especially in females. The synthorax
is mostly rather lightly built in the Zygoptera, more heavily so in the Anisoptera. The
wings of Zygoptera are narrow, particularly at the base, and fore- and hind-wings
are essentially similar (Fig. 1A). In Anisoptera the hind-wing is always expanded at
the base and the venation of the two wings differs substantially (Fig. 1B). The general
structure and venation of the wings are very different in the two suborders and the
details of venation are very important in classification and identification. There are
three pairs of legs, often modified in ways that assist with prey capture. The long, thin
abdomen consists always of 10 segments. In males the ventral side of the 2nd
segment bears a complex secondary copulatory organ. Frequently this is flanked by
small projecting flaps termed auricles or pseudoauricles which are of use in
classification. From the rear of the 10th segment arise: in male Zygoptera, two pairs
of anal appendages, upper (superior) and lower (inferior); in male Anisoptera, two
superior appendages and a single fused inferior; and in all females, two small dorsal
cerci. A ventral ovipositor is present in all Zygoptera and some Anisoptera.
Structure of larvae
The larvae are much stouter than their respective adults, and lack wings. In both
suborders the head is closely attached to the thorax and of limited mobility. The eyes
are relatively small and never touch. Behind the eyes are the often large, post-
ocular lobes, the shape and pattern of which may be useful in the determination of
species. The antennae usually have 3–7 segments and exhibit a variety of forms.
For example, the first segment in Calopterygidae may be as long as the other six
added together. In Gomphids the antennae have four segments that are short and
broad. All odonate larvae possess a hinged, prehensile labium, unique to the Order.
It is thrust forwards very rapidly by hydrostatic pressure and is used to capture prey.
It consists of the submentum and prementum on either side of a hinged articulation
and a pair of labial palps each with a movable hook. The structure and shape of this
organ and the number and arrangement of premental and palpal setae are extremely
useful in identification. The thorax, to which the legs are attached, consists of a
prothorax and meso-metathorax. The prothorax may have lateral outgrowths known
as the supracoxal armature, structures often used in the identification of aeshnid
412 Freshwater Invertebrates of the Malaysian Region

larvae. On the dorsum, four wing sheaths appear after the 3rd or 4th moult
(Fig.10A,B) and steadily increase in size with maturity. The abdomen is composed
of 10 segments with a vestigial eleventh. Frequently spines are situated on the mid-
dorsal and lateral aspects of the abdomen and can be taxonomically useful. Where
the adult female possesses an ovipositor, i.e. all Zygoptera and Aeshnidae, a
rudimentary structure is clearly visible on the ventral surface of segments 8 and 9.
Larvae of the two suborders can be separated easily on general structure and
the form of the anal appendages (Figs. 8, 10). Zygoptera are typically of light to
medium build with the abdomen tapered uniformly and two or three large caudal
lamellae present posteriorly. These may be sac-like, flattened or thin and elongate
and are very important in family classification. They function as gills and in
locomotion. Anisoptera are more robust, some being very broad and flattened. The
abdomen is widened in the middle before tapering posteriorly to a pointed anal pyramid,
consisting of a collection of short appendages including a central epiproct, two
lateral paraprocts, two smaller dorsal cerci, and in male Anisoptera a dorsal male
process which becomes the inferior appendage in the adult. The anus leads to a
rectal basket, concealed from the outside, which is lined with gills. The rapid expulsion
of water from the anus allows a form of jet propulsion, which serves as a secondary
form of locomotion, especially when escaping predators.

GENERAL BIOLOGY
Adults
Adult odonates typically emerge from the larval stage in the early morning. Some
clamber well away from the water, presumably to escape detection in the vulnerable
teneral stage; a few emerge very close to the larval habitat. Usually after an hour or
two the first flight is possible, but the newly emerged insect is not fully hardened for
at least a day and may take many days to achieve full adult colours and sexual
maturity. All species are exceptionally agile fliers and possess very keen vision.
Immature individuals spend much of their time feeding, usually away from the breeding
site, although in many Zygoptera this never seems to take them far from the water.
Both sexes of some species appear to form territories along forest paths or on tree
trunks which may be related to feeding. Even when sexually active, males of many
species feed during a set part of the day, sometimes at dawn and dusk, or feed on
alternate days. Prey taken are very diverse but poorly understood. Some of the
largest Anisoptera, such as Gynacantha (Aeshnidae), take very small Diptera and
other components of the aerial plankton, especially at dusk, whereas small Zygoptera
such as Ceriagrion cerinorubellum (Coenagrionidae) have been observed feeding
on teneral Archibasis (Coenagrionidae) substantially larger than themselves. In the
case of small forest Zygoptera which move little from their breeding sites, clearing
of waterside vegetation may contribute directly to their demise by denying them a
source of adult nutrition.
Insecta: Odonata 413

Mating mostly takes place at the breeding site which is regarded as the primary
habitat. Almost all bodies of water imaginable are utilized, from phytotelmata in the
forest, torrential mountain streams, lakes and permanent ponds to temporary puddles.
Malaysia, especially Sarawak, is rich in swamp forest habitats ranging from alluvial
freshwater swamp to peat swamp forest and mangrove. In general, primary forest,
especially mixed dipterocarp, freshwater swamp and kerangas, tends to support
shade loving stenotopic species with a high number of endemics, whereas open and
disturbed habitats as well as peat swamp forest and mangrove are characterized by
wide ranging, sun loving, eurytopic species. Only a few species are specialized to
the harsher environments of highly acidic water in peat swamps, or oligohaline pools
on the landward margins of mangrove. A greater proportion of stenotopic, stream
dwelling species among Zygoptera are found in the calopterygoid families, the
Platystictidae, Platycnemididae and Protoneuridae, and among Anisoptera in the
Gomphidae and Corduliidae. Eurytopic species typically include the Zygopteran
Coenagrionidae and the Anisopteran Libellulidae but there are many exceptions.
Both stenotopic and eurytopic species show promise as indicators of environmental
disturbance as the latter tend to invade pristine forest areas following perturbation,
while the former disappear.
Mating behaviour and the act of mating are complex. Males frequently defend
territories which include the oviposition site and either patrol the area (Corduliidae,
some Aeshnidae and Gomphidae), or perch nearby (most other groups). The steps
leading to mating may involve aerial capture, perhaps denying a female access to an
oviposition site without mating, or elaborate courtship and display of ornamentation
(Calopterygidae, Chlorocyphidae). Insemination is achieved by a curious indirect
mechanism. The male bears on his second segment a secondary sexual apparatus
with seminal vesicle and intromittent organ. Before mating he must charge the seminal
vesicle on segment 2 with sperm from the sperm reservoir opening on the underside
of the 9th segment. Prior to copulation the male grasps the head (Anisoptera) or the
prothorax (Zygoptera) of the female with his anal appendages. These may be of a
very specific form and designed to ensure species recognition. With this connection
made the pair are said to be in tandem. For the female to be inseminated, she must
bring her genital pore located on the 9th segment forward to engage with the male’s
secondary genitalia at the base of his abdomen. Once joined the pair are said to be
in the wheel. Copulation can last a few seconds or many minutes, part of the function
of prolonged copulation being to remove from the female’s spermatheca the sperm
of previous mates, frequently using microscopic recurved barbs on the penis (Waage
1984). Following insemination the pair disengage from the wheel but may remain in
tandem while the female lays her eggs. This is termed contact guarding. Non-
contact guarding occurs when the male releases the female, but follows her closely
as she oviposits in his territory. In many forest species females oviposit alone. All
414 Freshwater Invertebrates of the Malaysian Region

Zygoptera and the Anisopteran family Aeshnidae insert their eggs into a substrate
using the ovipositor. When the substrate is living plant tissue these may be very
firmly wedged. Other Anisoptera drop their eggs onto the water in masses,
frequently dipping their abdomen as they do so. In this case it is common for eggs to
be agglomerated in gelatinous strings or in small blobs which stick to vegetation.
The lifespan of most adult odonates is not well known, but small Zygoptera such
as Libellago species (Chlorocyphidae) may live no longer than a week or two. Large
Anisoptera, especially Aeshnidae, are generally considered to live up to several months.
Natural enemies are little known, but predation by birds may be a significant source
of mortality. Dolomedes spiders may take chlorocyphids sitting on logs near water
level and significant numbers of Zygoptera especially are taken in spider webs in the
vegetation around ponds and drains. Parasites include gregarines, water mites, and
ceratopogonid midges (Orr and Cranston 1997). These parasites have been shown to
increase mortality (Åbro 1990). An exhaustive account of all aspects of odonate
biology is given by Corbet (1999).
Larvae
Most of the odonate life cycle is spent as a larva in an aquatic environment and it is in
this stage we find the greatest diversity of form. Time taken for eggs to hatch varies
from just a few days to several months depending upon latitude and species. The
first stage in larval development is the prolarva. Here the young larva is totally
encapsulated within a cuticular sac that probably protects it during its escape from the
egg shell/plant material. The time spent in this stage before shedding the cuticle is
generally very brief, often less than a minute, however it may last longer if hatching
occurs out of water. The young larva then undergoes from 8 to 15 moults, a process
also known as ecdysis, before leaving the water and emerging as an adult. With
each successive moult after the early instars the wing cases grow proportionally larger
and their development is a guide to the maturity of the insect. Metamorphosis,
involving the absorption of many larval tissues and development of adult structures,
takes place within the final instar larval exoskeleton. There is no pupal stage.
Habitat selection is principally an adult activity and must satisfy the requirements
of all stages in the life cycle. Females sometimes select their oviposition site with
considerable care e.g. Pericnemis species (Coenagrionidae) breed only in water
filled cavities in plants, including buttress pans, rot holes and bamboo stumps (Orr
1994). Other species, notably Orthetrum (Libellulidae), are less specific and oviposit
in a wide variety of sites in still water or slowly moving water. Their larvae are
tolerant of wide fluctuations in environmental conditions e.g. temperature, oxygenation,
pH etc. It is interesting to note however that many larvae, including those from
specific habitat types, can be collected and reared under artificial conditions that are
very different to those experienced in nature. It is often quite simple to breed species
on a diet of Daphnia, mosquito larvae, tubifex worms or tadpoles of appropriate
Insecta: Odonata 415

size. Because of the risk of cannibalism it is important to separate larvae and as they
reach maturity they must be supplied with a twig on which to emerge.
Larvae are voracious carnivores, using vision and/or mechanoreceptors in the
detection of their prey. Typically, the prehensile labium is shot out very rapidly to
capture the target organism. Many odonates are sit-and-wait (ambush) predators
but some, especially lestids and aeshnids are more active and often stalk and chase
their prey. Anax guttatus (Aeshnidae) larvae may swim rapidly for several centi-
metres to reach a large prey item and young Indaeschna grubaueri (Aeshnidae)
kept in containers swim up to take mosquito larvae on the water surface. Pantala
flavescens (Libellulidae) swim-hover just below the water surface, supporting them-
selves with tiny pulses of water expelled from the anus, and actively pursue tadpoles
(Corbet 1999). These more active predators are often characterized by a longer
prementum than other forms. Adaptations for prey selection also depend upon the
size of the larva and, because they are opportunistic foragers, the diet reflects the
prey items available within their microhabitat. Large larvae are known to tackle small
fish, tadpoles, ovipositing adult Zygoptera and other odonate larvae including their
own species.
The larval habitat and mode of feeding may be related to the general form of the
larva, respiratory structures and behaviour and the structure of the mask. Respiratory
adaptations include the development of a long snorkel in the posterior segments of
Macrogomphus species (Gomphidae), allowing it to burrow to a depth of at least 5
cm and probably deeper while still taking clean water into its rectal basket. Species
living in environments very low in oxygen such as phytotelmata may have larger
breathing surfaces, as in the expanded caudal lamellae of Pericnemis, or in the case
of Indaeschna grubaueri, they may compensate in their behaviour, resting with
their anal pyramid just penetrating the water surface, apparently absorbing oxygen
from the atmosphere. Larvae of Tetracanthagyna (Aeshnidae) also almost leave
the water at night, sitting with only their head submerged, but this appears to be a
method of ambushing surface swimming fish. Larvae of Euphaea (Euphaeidae)
have auxiliary filamentous gills along the sides of the abdomen but given that they inhabit
well aerated riffles in clear streams this is evidently not an adaptation to low oxygen.
Larvae may be burrowers, clingers or simply live concealed within bottom detritus
or aquatic vegetation. Burrowers include riverine species e.g. Paragomphus
(Gomphidae), Chlorogomphus (Chlorogomphidae) and probably many chlorocyphids,
whose larvae lie in wait at least partially concealed by fine sand and mud.
Chlorocyphids have two tough elongate caudal lamellae, perhaps developed to
protect them from abrasion. Gomphids are often dorso-ventrally flattened, perhaps
as an aid to shallow burrowing, whereas Chlorogomphus larvae are covered in setae
able to trap fine substrate particles to aid concealment. Macrogomphus species that
burrow deep in silt are rounded and elongate. Many more species live deep in leaf
packs in and around torrents, including some chlorocyphids, Devadatta
416 Freshwater Invertebrates of the Malaysian Region

(Amphipterygidae) and Sieboldius (Gomphidae), the latter being in appearance a

perfect dead leaf mimic. It is not known from direct observation to what extent
burrowing types actively seek prey but it seems likely that they must forage for
items such as small worms and Diptera larvae, especially deep burrowers in silt.
Although the mask is typically extended explosively using hydrostatic pressure (the
mechanism of which is different in Anisoptera and Zygoptera) it can also be
extended at least partially by muscular effort. Burrowing forms feeding in sand,
mud or detritus on relatively sedentary small prey items detected by tactile or
chemoreceptors probably need only extend the labium slightly in order to capture
and hold their prey. A few odonate larvae are known to scavenge dead animal
material and it is possible this behaviour is more widespread than generally appreciated
among species living in leaf packs and bottom detritus.
Clingers include most Zygoptera, aeshnids and some libellulid species e.g. Zygonyx
and Onychothemis, which cling to rocks in fast flowing riffle areas of streams.
Some Drepanosticta (Platystictidae) species and Euphaea are found under stones
in riffles. Most other Zygoptera and the aeshnids that fall within this category cling to
vegetation and other structures above the bottom substrate. Most libellulid and corduliid
larvae tend to hide among roots, leaf trash and other bottom debris but may emerge
into the open at night to forage. Among these forms aeshnids especially are armed
with a strongly hooked labial palp and long, strong movable hook. These forms in the
later stadia attack large prey such as tadpoles when available. Species feeding
habitually on smaller prey, including many libellulids, corduliids and coenagrionids
tend to have broader scoop like labial palps with numerous fine setae. However
many species take a wide range of prey sizes and the form of the mask can at best
give a rough guide to dietary preferences. Cratilla metallica (Libellulidae), which
breeds in leafy forest pools, regularly takes tadpoles half its own size or larger,
although based on the structure of its mask it appears best adapted to smaller prey.
Some other libellulids including Camacinia, Tramea and Hydrobasileus cling to
vegetation near the water surface and swim actively. Their elongate fore and mid
tarsi bear a dense tuft of hairs which presumably helps them gain purchase on the
water. Such forms are also distinguished by their particularly broad heads.
The growth rate of tropical odonates is probably mostly dependent on available
food. For example Indaeschna grubaueri reared with super-abundant food developed
from egg to adult in 107 days (Orr 1994). These conditions are often approximated
in nature but individuals living in crowded tree holes may take two years to develop.
Smaller Zygoptera and extreme opportunists among the Libellulidae such as Pantala
flavescens breeding in temporary puddles may develop in as little as three or four
weeks; rapid development is essential if the larva is to mature before the water dries
up, although at least in some species partial desiccation can be endured between rains.
Odonate larvae can represent a high proportion of the standing stock (biomass)
of an aquatic community. The extreme situation occurs in phytotelmata where very
Insecta: Odonata 417

slowly growing aeshnid larvae feed on productive but short lived detritivores, and
the top level of the trophic pyramid of biomass is actually more than 50% of the
total. In less extreme situations, in small streams the total biomass of odonate larvae
may range from 10–30% of total, again supported by more productive lower trophic
levels. This means that actual number of larvae can be quite high, although it is
exceptional for routine stream surveys to discover more than a fraction of the larvae
actually present, mostly because many are either buried or superbly concealed.

REGIONAL TAXA
The Malaysian odonate fauna comprises 342 named species. (Present collections
also include half a dozen further species from Peninsular Malaysia and many more
from Borneo, as yet unidentified and/or undescribed and not included here; also it is
believed that many species still remain undiscovered in nature.) These include 161
species of Zygoptera in 10 families and 181 species of Anisoptera in 5 families.
Overall 239 species are known from Sabah, Sarawak and Brunei, and 226 from
Peninsular Malaysia (including Singapore). A total of 123 species, or 36%, are shared
between Peninsular Malaysia and Sabah–Sarawak. As expected the family
compositions of the two areas are very similar, but much higher levels of endemicity
are found in Borneo (40%) than the peninsula (11%) which shares much of its non-
Bornean fauna with either Sumatra or Thailand. In this context we define endemic
species as either those restricted to the Malay Peninsula or those occurring in Sabah,
Sarawak and Brunei and restricted to Borneo. To define eastern endemics as those
species found only within the political boundaries of Sabah and Sarawak would be
misleading and probably very inaccurate owing to our limited knowledge of the
fauna of the border regions of Kalimantan which surely include many species thus
far recorded only from Sabah or Sarawak. There is, in particular, a small but distinct
montane fauna normally associated with Gunung Kinabalu and possibly endemic
to this area but given the paucity of collecting in neighbouring ranges extending into
northern Kalimantan the wider distribution of many of these elements cannot be
excluded. About 40 species are known from Indonesian Borneo (Kalimantan) but
not yet located in Sabah or Sarawak. Many of these are definitely restricted to the
south but others may be found, especially in Sarawak, with increased collecting.
Within the peninsular species, the highest levels of endemicity occur in the family
Platystictidae, in which the nine members of the widespread genera Protosticta and
Drepanosticta are all endemic. No other family exhibits an endemicity level of
more than 33% and in seven families there are no endemic species at all. Nevertheless
the genera Sundacypha (Chlorocyphidae), Podolestes (Megapodagrionidae),
Pericnemis (Coenagrionidae), Brachygonia, Chalybeothemis and Pornothemis
(Libellulidae) all occur in both regions of Malaysia but are elsewhere restricted to
Sundaland (which also includes southern Thailand, Sumatra, Java, Bali and smaller
418 Freshwater Invertebrates of the Malaysian Region

Table 1. Species richness and endemicity of odonate families in Peninsular Malaysia and
Sabah–Sarawak. Significantly high proportions of endemics, 40% and above are in bold.
Endemicity is defined as species occurring exclusively in the Malay Peninsula or species
restricted to Borneo.

Taxon Peninsular Malaysia Sabah and Sarawak Total % common to

No. spp % endemic No. spp % endemic species Peninsula-Borneo
ZYGOPTERA 88 16 105 66 161 20
Amphipterygidae 1 0 1 100 2 0
Calopterygidae 6 0 9 78 13 15
Chlorocyphidae 10 0 18 67 23 22
Euphaeidae 3 0 8 75 9 22
Lestidae 5 0 3 33 6 33
Megapodagrionidae 4 25 5 80 8 13
Coenagrionidae 33 9 28 43 47 30
Platycnemididae 10 10 11 82 19 11
Platystictidae 9 100 11 100 20 0
Protoneuridae 7 14 11 55 14 31
ANISOPTERA 138 8 134 19 181 50
Gomphidae 32 19 24 46 46 21
Chlorogomphidae 3 33 1 0 3 33
Aeshnidae 15 13 27 15 30 43
Corduliidae 14 14 13 46 18 47
Libellulidae 74 0 69 7 84 69
Total 226 11 239 40 342 3

adjacent islands). Surprisingly, a few Bornean species restricted to southern

Kalimantan, hence not counted as Sabah–Sarawak species in this discussion, occur
in Peninsular Malaysia. These include Platylestes heterostylus (Lestidae),
Prodasineura interrupta (Protoneuridae) and Pseudagrion coomansi
(Coenagrionidae).
By contrast Sabah and Sarawak have (Bornean) endemicity rates exceeding
70% for the Zygopteran families Amphipterygidae, Chlorocyphidae, Euphaeidae,
Calopterygidae, Megapodagrionidae, Platystictidae and Platycnemididae. An astonishing
66% of all Zygoptera are endemic. The corresponding figure for the Anisoptera is
only 19% although rates of 46% occur in the Gomphidae and Corduliidae, still
considerably higher than in Peninsular Malaysia. Endemic genera include Rhinoneura
(Chlorocyphidae), Matronoides (Calopterygidae), Bornargiolestes
(Megapodagrionidae), Linaeshna (Aeshnidae) and Pseudagrionoptera
(Libellulidae). Additionally the coenagrionid genus Stenagrion is restricted to northern
Borneo and Palawan, a weak faunistic affinity present in other groups. To emphasize
the poor knowledge of the fauna of this area it should be noted that Linaeshna is
known from two specimens only (collected nearly 100 years apart) and
Bornargiolestes from two, dubiously conspecific specimens.
Insecta: Odonata 419

Widespread genera occurring in Borneo but not in the peninsula include Indolestes
(Lestidae), Xiphiagrion (Coenagrionidae), Anaciaeschna (Aeshnidae) and
Metaphya (Corduliidae). Conversely, genera present in Peninsular Malaysia, but
lacking in Borneo include Echo (Calopterygidae), Aristocypha (Chlorocyphidae),
Calicnemia, Indocnemis (both Platycnemididae), Cephalaeschna (Aeshnidae),
Gomphidictinus, Merogomphus, Nepogomphus, Onychogomphus, Paragomphus
(all Gomphidae), Brachythemis, Indothemis, Macrodiplax and Phyllothemis (all
Libellulidae). These are all widespread in Asia or at least found as far as Thailand.
The Malaysian odonate fauna forms part of a larger Sundaland fauna, numbering
over 500 species. Its components may be divided into a general Indo-Australian
element (including a small pan tropical group), a tropical Asia component, normally
arrested either by Wallace’s line or Weber’s line, a small Australian component,
including the corduliine genera Metaphya, Hemicordulia and Procordulia, and an
endemic Sundaland component already noted. The regional Caloptera damselflies
(i.e. the superfamily Calopterygoidea, containing Amphipterygidae, Chlorocyphidae,
Euphaeidae, Calopterygidae), although including representatives found in Thailand
and further west, are by and large endemic although ultimately the fauna is derived
from tropical Asian elements and genera such as Libellago and Rhinocypha
(Chlorocyphidae) are widespread. The group is poorly represented to the east beyond
Sulawesi by Neurobasis (Calopterygidae) and Rhinocypha. The Lestidae are poorly
represented by tropical Asian or ubiquitous (Lestes) genera. The small
megapodagrionid fauna is largely endemic, sharing little in common with the extensive
argiolestine fauna to the east. The Coenagrionidae are mostly Indo-Australian, at
least at the generic level, but only a small fraction of species are widespread. The
Platycnemididae and Protoneuridae are both represented by wide ranging Asian
genera none of which is found east of Wallace’s line, although other, mostly endemic
genera in these families occur in the Australian region. Platystictidae occur from Sri
Lanka to New Guinea, everywhere showing very high levels of species endemicity
but low generic diversity. Among the Anisoptera are very widespread genera such
as Anax, Gynacantha (Aeshnidae), Macromia (Corduliidae), Orthetrum,
Crocothemis, Trithemis, Tramea, Pantala, and Tholymis (Libellulidae). The
Gomphidae are mostly tropical Asian in origin without representatives to the east,
even at the generic level. An exception is the old-world tropical Ictinogomphus. The
Aeshnidae and Libellulidae are, in addition to the endemic elements mentioned above,
mostly Indo-Australian at the generic level, but largely tropical Asian at the species level.
Literature
M.A. Lieftinck’s (1904–1985) activity focused mainly on the Indo-Australian odonate
fauna. His publications, which contribute very much to our knowledge of Malaysian
odonates are listed and comprehensively indexed by van Tol (1992). Lieftinck’s
(1954) “Handlist of Malaysian Odonata”, which covers the fauna of the whole of
Sundaland, is still the major reference work for the area. Its references cover the
earlier literature thoroughly. Besides taxonomic data this publication also provides
420 Freshwater Invertebrates of the Malaysian Region

brief general comments on the habitats of each species. From the older publications,
a synopsis of Malay Peninsula dragonflies by Laidlaw (1931a) and a checklist of
Bornean dragonflies by Laidlaw (1931b) are of special importance. Since Lieftinck’s
handlist numerous, more specialized taxonomic and faunistic papers have been
published, many of which are listed in the references. Some are useful synopses of
the faunae of limited areas within the region, such as Murphy (1997) for Singapore,
Kitagawa (1977) for Penang and Orr (2001) for Brunei. Two papers focusing on
the general ecology of Malaysian odonates are Furtado (1969) and Thompson and
van Tol (1993). Orr (2003) provides an illustrated guide to many Bornean species and
is generally useful within the region.
Fraser’s (1933–1936) volumes on the Odonata of the former British India,
including Myanmar continue to be extremely useful in identification of adults of
many Malaysian species, especially those from the peninsula. Asahina’s (1993)
treatment of the Thai fauna is also very useful for identifying Peninsular Malaysian
dragonflies. The recent books on adult dragonflies of Thailand (Pinratana et al.
1988; Ek-Amnuay 1996; Hämäläinen and Pinratana 1999), Hong Kong (Wilson
1995, 2003) and Taiwan (Wang 2000) also illustrate in colour several Malaysian
species. Publications for identification of larvae of Malaysian species are harder to
find. Ishida’s (1996) richly illustrated book of Japanese dragonfly larvae is also useful
for identifying the Malaysian fauna at family and generic level and some of Lieftinck’s
published and unpublished drawings are collected in van Tol (1992).
However, as a first introduction to dragonflies for the beginner, two publications
are especially recommended. Miller (1995) is a concise, but many sided treatment
of dragonfly biology, and Watson and O’Farrell (1991) provides an excellent brief
introduction to dragonfly structure.
For more specialized taxonomic matters, the detailed world Odonata catalogue
by Bridges (1994) is an invaluable source of taxonomic and biographic information,
and the world list of Odonata by Tsuda (2000) includes the known species with their
distributions. Excellent general accounts of other faunas which include detailed
accounts of biology and morphology include Askew (1988) on the European fauna,
Watson et al. (1991) on the Australian fauna which shares some elements with
Malaysia, Westfall and May (1996) and Needham et al. (2000) on the North
American fauna. Finally, a most comprehensive synopsis of present knowledge of
the ecology and behaviour of Odonata is provided by Corbet (1999).

KEYS TO ADULTS (by A.G. Orr)

Key to Suborders: adults
1. Eyes well separated by a space greater than their own diameter. Fore- and hind-wings are
essentially similar in shape and venation, narrow at base (Fig.1A) ............................ Zygoptera
- Eyes usually confluent across the middle line, or if separated, never by a space greater than their
own diameter. Hind-wing expanded at base and venation differs from that in fore-wing (Fig. 1B)
............................................................................................................................... Anisoptera
Insecta: Odonata 421

Key to Zygoptera families: adults

1. Numerous (5–70+) antenodal crossveins in the costal space of both wings (Fig. 2A,B). Wings
often with opaque markings, sometimes iridescent .................................................................. 2
- Only two antenodal crossveins in costal space (Fig. 3F). Wings hyaline .............................. 5
2. Wings strongly petiolated just basal to level of arculus (Fig. 2A). Mostly drab species of
medium size and build (hind-wing 28–34 mm*); wings hyaline ....... AMPHIPTERYGIDAE
* Philoganga, which may occur in Peninsular Malaysia, is larger.
- Wings not petiolated, or only weakly petiolated (Fig. 2B) ................................................... 3
3. Antenodals in costal and subcostal space not aligned distal to level of arculus (Fig. 2C). Only
5–15 antenodals in costal space. Ante- and postclypeus produced into a long upturned rostrum
(Fig. 2F). Mostly small species, HW 15–30 mm, abdomen shorter than hind-wing except in
Rhinoneura (montane Borneo) and often brightly coloured ................ CHLOROCYPHIDAE
- Most of the numerous (15–70+) antenodals in costal and subcostal space aligned except distally
near the subnodus (Fig. 2D, E). Clypeus not produced into a rostrum. Abdomen longer ......... 4
4. Only one crossvein in cubital space basal to arculus (Fig. 3A). Fore-wing markedly longer than
hind-wing. Males often with small pseudoauricles on abdominal segment 2 (Fig. 2G). Head,
synthorax and abdomen never bright metallic green. Robust species with short legs; hind-wing
24–32 mm ....................................................................................................... EUPHAEIDAE
- Several crossveins in cubital space basal to arculus (Fig. 3B). Wings of nearly equal length, broad
and paddle shaped. Males lacking pseudoauricles. Head, synthorax and abdomen mostly bright
metallic green. Lightly built long legged species; hind-wing 30–40 mm .... CALOPTERYGIDAE

5. Numerous supplementary veins inserted between the main veins from the distal wing margin to
the level of the pterostigma (Fig. 3C) ...................................................................................... 6
- Supplementary veins absent from the wing tip, or if present, to the depth of 1–2 cells only (Fig.
3D) ......................................................................................................................................... 7

6. IR3 and R 4+5 originating closer to arculus than to subnodus (Fig. 3E). Fore-wing quadrilateral
with costal side very short; almost triangular. Small to medium sized lightly built species resting
with wings open (HW 21–31 mm) ....................................................................... LESTIDAE
- IR3 and R 4+5 originating closer to subnodus than to arculus (Fig. 3G). Fore-wing quadrilateral
longer, parallel sided or diverging, costal side nearly as long as anal side. Fairly robust species
resting with wings open (HW 20–30 mm) .................................... MEGAPODAGRIONIDAE
7. Anal vein absent or very poorly developed, never extending beyond distal end of quadrilateral
(Fig. 4D,E). CuP meeting wing margin less than halfway along its length ............................. 8
- Anal vein well developed, extending at least two cells beyond distal end of quadrilateral (Figs. 4F,
5A,B). CuP meeting wing margin at least halfway along its length, usually well beyond this.... 9
8. Wings narrow, terminally falcate (Fig. 4D). Small sub-basal crossvein between CuP and wing
margin (Fig. 4A). Sectors of arculus fused basally to form a short stalk (Fig. 4A). Small very
fine built species (HW 16–26 mm), often metallic, male appendages thin elongate and branched
............................................................................................................... PLATYSTICTIDAE
- Wings not falcate. No sub-basal crossvein between CuP and wing margin. Sectors of arculus
separated basally (Fig. 4B). Small fine non-metallic species (HW 17–22 mm); male appendages
squat ....................................................................................................... PROTONEURIDAE
422 Freshwater Invertebrates of the Malaysian Region

Figure 2. A – hind-wing of Devadatta podolestoides (Amphipterygidae) showing venation and

petiolation. B – hind-wing of Rhinocypha aurofulgens (Chlorocyphidae) showing venation and general
wing shape. C – detail of costal and subcostal space in hind-wing of R. aurofulgens showing mostly
non-aligned distal antenodals (Ax). D – same for Vestalis beryllae (Calopterygidae) showing mostly
aligned distal antenodals. E – same for Euphaea subcostalis (Euphaeidae) showing aligned distal
antenodals. F – lateral and dorsal view of head of Libellago semiopaca (Chlorocyphidae) showing
rostrum. G – ventral view of second abdominal segment of Euphaea subcostalis showing pseudauricles.
Insecta: Odonata 423

9. Costal side of quadrilateral in fore-wing two fifths or less length of anal side (Fig. 5B,C,D).
Tiny to large species (hind-wing 11–36 mm) .......................... (most) COENAGRIONIDAE
- Costal side of quadrilateral in fore-wing at least half length of anal side (Figs. 4F, 5A,E,F) ...... 10
10. Hind tibia with 4–8 short stout spines; legs short, hind tibia less than length of humeral suture
(Fig. 5H). Male inferior appendage rarely as long as segment 10, generally squat and often
shorter than superior (Fig. 5J,K). Hind-wing sometimes less than 15mm (range 9–22 mm) ......
............................................................................................................. COENAGRIONIDAE
(some Agriocnemis, some Amphicnemis, Argiocnemis, some Mortonagrion, Stenagrion)
- Hind tibia with 10–16 long filamentous spines; mostly longer than humeral suture (Fig. 5G). Male
inferior appendage thin, tapered, longer than superior and segment 10 (Fig. 5I). Hind-wing never
less than 15 mm (range 16–35 mm) .................................................... PLATYCNEMIDIDAE

Key to Anisoptera families: adults

Note. Females of some Corduliidae and Libellulidae may be difficult to separate on family characteristics.
1. Eyes well separated (Fig. 6A) or close but not quite touching when viewed from above. If eyes
fairly close then large (HW > 40 mm) green and yellow banded species, usually with auricles in
the male .................................................................................................................................. 2
- Eyes broadly contiguous (Fig. 6C) or at least touching viewed from above. If eyes just touching
(Fig. 6B) then medium sized (HW < 40 mm) red or brownish species with the male lacking
auricles ................................................................................................................................... 3
2. Median space with two or more crossveins (Fig. 6D). Large black and yellow or green banded
species (HW > 40 mm) ..................................................................... CHLOROGOMPHIDAE
- Median space lacking crossveins (Fig. 6E); Eyes always well separated. Small to large species
(HW 22–55 mm) ............................................................................................... GOMPHIDAE

3. Triangles similar in both wings, elongate along wing axis with costal side much longer than basal
side (Fig. 7A). Female always with well developed ovipositor. Medium sized to huge species
(HW 36–80 mm) ................................................................................................ AESHNIDAE
- Triangles dissimilar, at least in the sense that they cannot be superimposed without rotation; If
elongate along wing axis in hind-wing, then elongate parallel to body in fore-wing (Fig. 7B);
female lacking ovipositor ......................................................................................................... 4
4. Ground colour of thorax and abdomen almost always metallic; most species with well defined
projection on the hind margin of the compound eye (Fig. 7D,E); males of most species with hind-
wing anal angle acute (Fig. 7G) and auricles present ventrally on segment 2 of abdomen (Fig.
6F). Males always with tibial keels on the inner (flexor) surface (Fig. 7C), at least on fore and
hind legs. Small to large species (HW 22–55 mm) ......................................... CORDULIIDAE
(Males of Hemicordulia (Fig. 1B) are distinguished only by presence of tibial keels and metallic
green colouration)
- Ground colour of thorax mostly non-metallic and abdomen only very rarely metallic—more
commonly dorso ventrally flattened and red or blue; never with well defined projection on hind
margin of eye—at most a marked sinuousness (Fig. 7F); males without auricles and with anal
angle of hind-wing always rounded (Fig. 7B); tibial keels never present on any legs. Tiny to
large species (hind-wing 12–53 mm) ............................................................ LIBELLULIDAE
(In Zygonyx the abdomen is metallic but males lack tibial keels and other corduliid
features)
424 Freshwater Invertebrates of the Malaysian Region

Figure 3. A – hind-wing base of Euphaea subcostalis (Euphaeidae) showing single crossvein in cubital
space basal to arculus (Ac). B – same of Vestalis beryllae (Calopterygidae) showing several crossveins
in cubital space basal to arculus (Ac). C – wing tip of Podolestes orientalis (Megapodagrionidae)
showing supplementary veins between main veins (labelled), beyond level of pterostigma. D – same of
Pericnemis triangularis (Coenagrionidae) showing supplementary veins no more than two cells deep.
E – fore-wing of Lestes praemorsus (Lestidae) showing venation, especially position of IR3 and R4+5
arising closer to arculus than to subnodus. F – detail of costal and sub-costal spaces in L. praemorsus.
G – fore-wing of Podolestes orientalis (Megapodagrionidae) showing venation, especially position of
IR3 and R4+5 arising closer to subnodus than to arculus.
Insecta: Odonata 425

Figure 4. A – hind-wing base of Drepanosticta rufostigma (Platystictidae) showing sub-basal cubito-

anal crossvein, anal crossing (ac), stalked sectors of the arculus and very short CuP. B – hind-wing base
of Prodasineura verticalis (Protoneuridae) showing ac, anal vein reduced to short anal bridge (ab) and
separated sectors of the arculus. C – lateral view of male anal appendages of Prodasineura sp. showing
short, squat superiors and inferiors. D – hind-wing of D. rufostigma showing general venation and
falcate wing tip. E – hind-wing of P. verticalis showing general venation and detail of long, thin
quadrilateral. F – fore-wing of Copera vittata (Platycnemididae) showing venation with well developed
anal vein (A1) and detail of long thin quadrilateral.
426 Freshwater Invertebrates of the Malaysian Region

Figure 5. A – fore-wing of Coeliccia macrostigma (Platycnemididae) showing venation with well

developed anal vein (A1) and detail of quadrilateral: sides labelled as; ‘a’, anal; ‘b’, basal; ‘c’, costal; ‘d’,
distal. B – fore-wing of Pericnemis triangularis (Coenagrionidae) showing venation with well developed
anal vein (A1) and detail of quadrilateral: sides labelled as; ‘a’, anal; ‘b’, basal; ‘c’, costal; ‘d’, distal.
C – quadrilateral in fore-wing of Onychargia (Coenagrionidae). D – quadrilateral in fore-wing of
Ischnura (Coenagrionidae). E – quadrilateral in fore-wing of Agriocnemis (Coenagrionidae). F –
quadrilateral in fore-wing of Indocnemis (Platycnemididae). G – lateral view of head (twisted) and
thorax of Coeliccia macrostigma (Platycnemididae) showing humeral suture on synthorax, legs including
meta-tibia with spines. H – same of Stenagrion dubium (Coenagrionidae). I – lateral view of male anal
appendages in C. macrostigma. J – lateral view of male anal appendages in S. dubium. K – lateral view
of male anal appendages in Amphicnemis remiger (Coenagrionidae).
Insecta: Odonata 427

Figure 6. A – dorsal view of head of Ictinogompus decoratus (Gomphidae). B – dorsal view of head of
Rhodothemis rufa (Libellulidae). C – dorsal view of head of Anax guttatus (Aeshnidae). D – base of
hind-wing of male Chlorogomphus sp. (Chlorogomphidae). E – base of hind-wing of male Ictinogomphus
decoratus (Gomphidae). F – lateral view of male abdominal segments 1–2 showing auricles in Macromidia
fulva (Corduliidae).
428 Freshwater Invertebrates of the Malaysian Region

Figure 7. A – base of fore- and hind-wings of Indaeschna grubaueri (Aeshnidae) illustrating similar
triangles. B – base of fore- and hind-wings of Orthetrum pruinosum (Libellulidae) illustrating dissimilar
triangles and the foot-shaped anal loop and rounded anal angle on the hind-wing. C – hindleg of
Hemicordulia tenera (Corduliidae) showing the larger divisions of the leg and the membranous tibial
keel. D – lateral view of compound eye of male Macromia corycia (Corduliidae) showing projection on
hind margin. E – the same for female Idionyx yolanda (Corduliidae) showing a less prominent but still
definite projection. F – the same for Zygonyx iris (Libellulidae) showing a sinuous margin with no
definite projection. G – base of fore- and hind-wings of Macromia cincta (Corduliidae) illustrating
dissimilar triangles and the acute anal angle on the hind-wing.
Insecta: Odonata 429

KEYS TO LARVAE (by S.G. Butler and A.G. Orr)

In using this key to Malaysian odonate larvae it must be noted that some families
and many genera have not yet been satisfactorily separated. There has been little
focused research on odonate early stages in Malaysia, some artificial groupings
(e.g. Megapodagrionidae) are yet to be resolved and the earlier keys of Needham
and Gyger (1937, 1939) have not yet been updated. Cross-generic characteristics
occur in larval Gomphidae especially, the result of parallel evolution of structures
serving similar functions, in a stage which may occupy up to 90% of their life cycle.
Also, recent major studies in phylogeny are producing changes to the classification
of the order. This key can only therefore be tentative and incomplete, but its main
value will hopefully be to provide a basis for further work, based upon the rearing
out of larvae. The key is mainly a family key, but owing to the closeness of certain
members of the oriental Libellulidae and Corduliidae, the latter are separated on
generic characteristics.

Key to Suborders: larvae

1. Slender, usually with cylindrical abdomen which tapers distally. Posterior of abdomen with 2
or 3 caudal appendages (gill lamellae) projecting from end of abdomen (Figs. 10A, 13) ...............
................................................................................................................................ Zygoptera
- Stout, abdomen variable, but always widening somewhat distally, before then tapering to anal
triangle. Posterior of abdomen without conspicuous caudal appendages or lamellae, but with 5
short spinous appendages (Figs. 10B, 14) ............................................................... Anisoptera

Key to Zygoptera families: larvae

1. Antennal segment 1 almost as long as others combined (Fig. 10C) ....................................... 2
- Antennal segment 1 much shorter than above (Fig. 10D) ...................................................... 3
2. Body cylindrical in cross-section; lamellate central gill shorter than triquetral laterals (Fig.
10E); labium with tiny scattered premental setae and large premental cleft (Figs 10F, 13G)...
............................................................................................................. CALOPTERYGIDAE
(Larvae are elongate with long spidery legs. They live mainly in forest streams in clear
flowing water in leafpacks, among stones and root masses, or they may be in slower
rivulets in swamp forests)
- Compact body, rather depressed; labium flat, lacking premental and palpal setae; central gill
reduced to conical process, laterals spiniform and sharply triquetral (Figs 10G, 13D) ..........
............................................................................................................ CHLOROCYPHIDAE
(Larvae are somewhat stout, with the caudal lamellae modified to form 2 long and thin
lateral cerci, the median lamella being reduced to a tiny remnant. Individuals can be
difficult to locate and may be buried deeply in detritus or silt)

3. Abdomen with lateral gills (Figs 10H, 13B) .................................................. EUPHAEIDAE

(Larvae are robust, with broad triquetral caudal lamellae, as well as their unique (in Odonata)
abdominal gills. They are mainly found under stones or amongst detritus in riffles)
- Abdomen without lateral gills ................................................................................................ 4
430 Freshwater Invertebrates of the Malaysian Region

4. Caudal gills saccoidal, or with saccoidal parts (Fig. 10 I) ...................................................... 5

- Caudal gills lamellate (Fig. 10 J,K). ....................................................................................... 6
5. Caudal lamellae inflated, all similar, in the form of simple, elongate-oval sacs with long,
filamentous tips (Figs 10L, 13C) ........................................................... PLATYSTICTIDAE
(Larvae are small and slender, with inflated (saccoidal) filamentous lamellae. The labium
is square-shaped, lacking in large premental setae and with a pronounced median cleft on
the distal margin)
- Caudal lamellae bearing scale-like setae, median lamella very different in size and shape from
laterals (Figs 11A, 13A) .................................................................... AMPHIPTERYGIDAE
(The genus Devadatta has stout larvae, which are found in leaf packs, amongst root
systems or under stones in small, rocky forest streams)

6. Labium distal margin with median cleft (Fig. 11E) ................................................................ 7

- Labium with distal margin entire (Fig. 11D) .......................................................................... 8
7. Setae present on moveable hook of labial palps, proximal portion of labial prementum greatly
narrowed (Fig. 11E), lamellae with secondary trachea arising at right angles to main stems (Figs
11F, 13H) .............................................................................................................. LESTIDAE
(Slender, rather elongate, active predators. They are found in shallow marshy habitats,
ranging from pristine alluvial forest and the verges of peat swamp forest to open drains
and permanent ponds)
- Labium with median cleft and with a single palpal seta, but without premental setae (Figs 11B,
13E) ............................................................................................ MEGAPODAGRIONIDAE
(Stout, often long legged forms, found in leaf litter in streams and stagnant pools in swamp
forest and among leaves in trickles from forest springs. Rhinagrion mima, the only described
larva of this family, has unusual lamellae, the central and laterals being very different from
each other. Fig. 11C)

8. Lamellae with thickened proximal half and with membranous distal half (Fig. 11 I) often ovoid
or subquadrate in shape (Fig. 11H) ........................................................ PROTONEURIDAE
(Larvae are short and stout, with broad heads and long legs. The caudal lamellae are short
and stout, the central gill being shorter than the laterals. Larvae are found in silty areas
among root masses in slowly flowing swampy areas, backwaters and stagnant edges of
streams.)
- Lamellae not thickened basally and variable in shape, sometimes filamentous (Fig. 11K) .... 9

9. Lamellae nodate or subnodate (Fig. 11G), lateral margins of occiput rounded (Figs 11L, 13F)
............................................................................................................. COENAGRIONIDAE
- Lamellae denodate, lateral margins of occiput variable ......................................................... 10
10. Lamellae with a filamentous fringe (Fig. 11K), lateral margins of occiput slightly angled (Fig.
11J) ................................................................ some PLATYCNEMIDIDAE (most Copera)
(The larvae of Platycnemididae vary considerably generically and currently cannot be grouped
together. They occur in leaf packs on streams and in leaf litter in stagnant forest pools)
- Lamellae lacking filamentous fringe and denodate, lateral margins of occiput variable ............
................................................... some COENAGRIONIDAE, some PLATYCNEMIDIDAE
(These larvae of the more common coenagrionid species are often abundant in open ditches,
ponds, pools and lakes amongst aquatic vegetation. Some coenagrionid and most platycnemidid
genera are found in more pristine habitats such as swamp forest, whilst one coenagrionid genus
specialises in phytotelmata).
Insecta: Odonata 431

Figure 8. Lateral view of larva: A – Zygoptera; B – Anisoptera; Ventral view of larva: C – Aeshnidae
male. D – Aeshnidae female.
432 Freshwater Invertebrates of the Malaysian Region

Figure 9. Labium dorsal view: A – Libellulidae/Corduliidae; B – Aeshnidae; Head frontal

view: C – Libellulidae/Corduliidae: LHS lamella lateral view: D – Coenagrionidae; Lamella
dorsal view: E – Coenagrionidae; Anal appendages dorsal view: F – Aeshnidae X; G – Aeshnidae C;
LHS Antenna dorsal view: H – Calopterygidae; I – Coenagrionidae; J – Gomphidae; K – Gomphidae.
Insecta: Odonata 433

Figure 10. A – Zygoptera (Coenagrionidae), dorsal view; B – Anisoptera (Libellulidae), dorsal view;
C – Calypterygidae LHS antenna, dorsal view; D – Libellulidae LHS antenna, dorsal view; E – Calopterygidae
lamellae lateral view LHS; F – Calopterygidae labium, dorsal view; G – Chlorocyphidae gills, dorsal
view; H – Euphaeidae abdominal gills ventral view; I – saccoidal gills dorsal view; J – lamellate gills RHS
lateral view; K – lamellate gills dorsal view; L – Drepanosticta sundana lamellae, dorsal view.
434 Freshwater Invertebrates of the Malaysian Region

Figure 11. A – Devadatta argioides base of epiproct and paraprocts dorsal view; B – Megapodagrionidae
labium in dorsal view; C – Megapodagrionidae (top)central lamella in lateral view, (bottom) LHS lamella
in lateral view; D – Coenagrionidae labium in dorsal view; E – Lestidae labium in dorsal view; F – Lestidae
RHS lamella, in lateral view; G – Coenagrionidae RHS lamella, lateral view; H – Protoneuridae lamella in
lateral view; I – Protoneuridae caudal lamellae in dorsal view; J – Platycnemididae head in dorsal view; K
– Platycnemididae LHS lamella in lateral view; L – Coenagrionidae head in dorsal view.
Insecta: Odonata 435

Key to Anisoptera families: larvae

1. Antennae 5–7 segmented (Fig. 10D), all tarsi 3-segmented (Fig. 12C) .................................. 2
- Antennae 4 segmented, usually flattened (Fig. 12A); pro- and mesotarsus 2-segmented (Figs
12B, 14A–C) ................................................................................................... GOMPHIDAE
(Extremely variable in form, ranging from leaf shaped (Sieboldius spp) to elongate with
extended segment 10 (Macrogomphus spp). Mainly ovate and flattened and normally
found amongst detritus, silt or under stones, predominately in running water)

2. Labium spoon-shaped (Fig. 12G,H,N,O), labial palp with moveable hook shorter than breadth
of palp and distal margin crenulated or serrated (Fig. 12G,H,N,O) ...................................... 3
- Prementum flat, labial palp with distal margin smooth and moveable hook longer than breadth
of palp (Fig. 12E), body elongated with hind leg not reaching the apex of abdomen (Figs 12D,
14E,H) ............................................................................................................... AESHNIDAE
(Elongate and active predators, taking a range of prey including large tadpoles and small
fish. Found in a range of habitats, including lakes, open ponds, forest pools, steams, drains
and phytotelmata)

3. Body short, squat and dorso-ventrally flattened, hind leg reaching beyond apex of abdomen
(Fig. 12J). Apex of prementum entire; labial palp with distal margin more evenly crenulated
(Fig. 12H,I,N,O) .................................................................................................................... 4
- Body elongated with tarsal claw of hind leg not reaching to apex of abdomen (Fig. 12F); Apex
of prementum bifid; labial palp with distal margin with deep, irregular, serrate dentations (Figs
12G, 14D) ...................................................................................... CHLOROGOMPHIDAE
(Chlorogomphid larvae are elongate and finely hirsute mud or sand burrowers. They can
be found in small forest streams, or in torrential sections of larger streams)

4. Head without extended frontal shelf, horn or ridge; variable palps, labium, anal appendages and
abdominal spines ................................... LIBELLULIDAE and some CORDULIIDAE (rare)
(Variable in both size and shape; larvae can be found in open lakes, paddy fields, ponds and
drains or forested swamps amongst waterweed or detritus. Others may be found at the
margins of rivers in silt, whilst one genus Zygonyx is modified to live on rocks in torrential
streams and another species Lyriothemis cleis breeds in phytotelmata in forest understorey)
- Frontal shelf between antennae, either extended frontally, or with elevated ridge or horn (Figs
12 L, M) ................................................................................................................................. 5
5. Prementum and palps without raptorial setae; distal margin of palps with very large crenulations
(Fig. 12K) ............................................................................. CORDULIIDAE: Epophthalmia
(Larvae are large, with a uniquely shaped labium and very long legs. They are found in the
margins of large lakes or drains in the vicinity of forests)
- Prementum and palps with setae, crenulations of palps smaller and more regular ................... 6
6. Long legged, with an abdomen that may be flattened or triangular in cross-section and is usually
strongly hooked dorsally. Labium lacking fields of tiny setae (Fig. 12N). Mature larva often
large. (Fig. 14F) .......................................................................... CORDULIIDAE: Macromia
(The head has a variably shaped frontal horn between the antennae. Larvae can be found
in torrential or slow sections of streams and rivers)
- Larva smaller and with warty setae on ventral surface of labial palps, prementum and palps
also bear irregular fields of tiny setae (Fig. 12O) ............................ CORDULIIDAE: Idionyx
(Comparatively little known, larvae are squat, hairy and somewhat cylindrical in shape.
Larvae have been found in small streams amongst detritus and algal growth)
436 Freshwater Invertebrates of the Malaysian Region

Figure 12. A – Gomphidae LHS antenna in dorsal view; B – Gomphidae RHS foreleg in dorsal view;
C – Aeshnidae RHS leg in dorsal view; D – Aeshnidae in dorsal view; E – Aeshnidae labium in dorsal
view; F – Chlorogomphus sp. in dorsal view; G – Chlorogomphus sp. labium in dorsal view; H –
Libellulidae sp. labium in dorsal view; I – Libellulidae sp. labium in dorsal view; J – Libellulidae in dorsal
view; K – Epophthalmia frontalis labium in dorsal view; L – Macromia moorei head in frontal view; M
– Macromia moorei head in LHS lateral view; N – Macromia sp. labium in dorsal view; O – Idionyx
stevensi (not Malaysian), labium in dorsal view.
Insecta: Odonata 437

Figure 13. Zygoptera larvae. A – Amphipterygidae: Devadatta argyoides; B – Euphaeidae: Euphaea

subcostalis; C – Platystictidae: Drepanosticta sp.; D – Chlorocyphidae: Libellago hyalina; E –
Megapodagrionidae: Rhinagrion borneense; F – Coenagrionidae: Pericnemis triangularis, with lateral
view of caudal lamellae; G – Calopterygidae: Neurobasis longipes; H – Lestidae: Orolestes wallacei, with
detail of mask and labial palp; I – Platycnemididae: Copera vittata, with lateral view of caudal lamella.
438 Freshwater Invertebrates of the Malaysian Region

Figure 14. Anisoptera larvae. A – Gomphidae: Megalogomphus sp.; B – Gomphidae: Sieboldius

japponicus; C – Gomphidae: Macrogomphus sp.; D – Chlorogomphidae: Chlorogomphus sp.; E –
Aeshnidae: Indaeschna grubaueri; F – Corduliidae: Macromia westwoodii; G – Libellulidae:
Hydrobasileus croceus; H – Aeshnidae: Tetracanthagyna degorsi; I – Libellulidae: Cratilla metallica.
Insecta: Odonata 439

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The Internet also provides an ever increasing amount of information on dragonflies. Links to
reliable websites may be found at the Odonate Information Network [http://www.afn.org/~iori].

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