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In this page we describe the typical sequence of stages during mating
and reproductions of dragonflies and damselflies.
Their reproduction and
associated behaviour is unique among the animal world. The behaviour is complex
but universal within dragonflies and damselflies, although with difference spices there could be some minor variations.
Territorial Behaviour
Some dragonflies and damselflies species show very
strong territory behaviour while some species do not. A territory is defined as a fixed area that an animal defends against intrusion from
other of its own species. In dragonflies and damselflies the territory will usually
be a good egg-laying site.
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- Above pictures show the hovering Australian Emperor
and the perching Fiery Skimmer guarding their
territory.
The male dragonflies and damselflies will return to their breeding ground and
stay there when
they mature and ready for mating. Most male Skimmer Dragonflies
perch on their favorite spot and overlook their territory. The male Hawker
and Emerald will usually fly, non-stop within a
fixed path over their territory. When there is the intruders of the same species
male (sometime other species as well), the territory holder will driver away the
intruder, by showing better body colour, better flying skill, larger in size or
some other advantages. Sometime the very close and violent contact are occurred. The
males will stay in their territory and wait for the female.
In most dragonflies and some damselflies, females only visit their breeding
site to mate and lay eggs. They will leave the breeding sit after laying eggs,
until next batch of eggs are ready.
When the female enter the male territory
(or when a male meet a female in the non-territory species), they will start the
next mating sequence, the courtship and recognition.
Courtship and Recognition
- Courtship and recognition includes the male showing his ovipositing
site to the female. By courtship, the male and female can make sure they are the same species and are suitable mates. The female may reject the mating if she thinks the
ovipositing site is not good enough. There are different courtship pattern for different species.
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- A female Fiery Skimmer
come into a male territory,
they just start the
courtship and recognition.
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- However, courtship is not so common in dragonflies and damselflies. For
most species, no courtship can be seen.
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- For some species, after the female enters the male territory, she will
immediately be grasped and clasped by the male and the couple will mate.
Some species males just snatch unwary females while they're warming in the
sun. Some species males even grab the immature ones, shimmer-fresh after emergence.
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For the non-territory species, there will also be the courtship and recognition
stages. They may swarm and several males may pursuit the same female. This
is common in damselflies.
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- If the female do not accept the male, various signal will
be made, depends on species. The most common is the downward curving of her
abdomen. If the female indicates acceptance, the male will grasp her thorax
with his legs.
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- No matter there is the courtship or without courtship,
after the male grasp the female, they start the
next mating sequence, Grasping and Clasping.
Grasping and Clasping
The male grasps the female head or thorax and curves his abdomen to
clasp her prothorax or head with his anal appendages to form the tandem pair. This action usually takes less than a second.
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- The first picture shows a male Fiery Skimmer
overtakes a female from above. They may do this
in air or when the female is on vegetation. Sometimes the male may force the female to land or even knock flying female onto the
water (as shown in the second picture, the Blue Skimmer).
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- The above pictures show the anal appendages at the abdomen tips of a Australian
Flatwing Damselfly male and a Blue Skimmer
Dragonfly male.
Damselflies have a pair of inferior appendages and a pair superior
appendages, i.e.., totally four appendages. Dragonflies have a pair of superior appendages and
a single inferior one, totally three. Those specialized appendages are used to clasp the female for copulation. In damselflies the
male clasp the prothorax of the female. In
dragonflies the male clasp the female back of the head. Those appendages,
like lock and key, will only fit into the same species female.
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- At this stage, the couple is said to be 'in tandem position'.
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- Tandem position is the male seizure of the female head
(dragonflies) or prothorax (damselflies). The male and female will be in tandem for a while, from seconds to minutes.
They may fly in tandem or find a suitable place to settle.
Intramale Sperm translocation
All insect males have their genital opening for sperm at the
ninth abdominal segment. Dragonflies/damselflies males have (beside the first genitalia)
their secondary genitalia, which is the accessory
organ on the second abdominal. Sperm is moved from the first genitalia into the secondary genitalia
just
before copulation.
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- The picture shows the male transferring sperm to his genitalia (or accessory
genitalia). The male bend his abdomen forward so that the first genitalia touches his secondary genitalia. This may take several seconds.
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- For dragonflies the males do this before he finds a female.
For damselflies the males do this immediately afterwards.
Copulation invitation
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- After forming a tandem and the sperm in the male secondary genitalia
is ready, male damselfly invites copulation by wing flapping and flexing the
abdomen. The female responds with bending up her abdomen to the
genitalia of the male and form the wheel position.
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- Dragonflies usually start copulation in flight, the male swing up the
female.
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- The female then curls her abdomen forwards to contact with the secondary
genitalia in order to receive sperm. The male and female form the heart
shape wheel and it is known as Wheel Position. Notice that in damselfly male is holding by female's neck (prothorax)
while in dragonfly is holding the back her head.
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- Most damselflies settle and perched for copulation. Most dragonflies start
copulation in flight, some may continue in flight and some may settle.
Settle copulation usually last longer, from minutes to hours. Air
copulation may last only a few seconds.
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- For some species, it is found that before transferring his sperm, the male
will remove the existing sperm from the female genitalia which is from earlier mate, then
replace it with his own.
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Tandem after Copulation
After the copulation the couple may separate or remain in tandem, depends on
species.
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- A long period of Tandem Position may be occurred after copulation. The couple
may be at rest or in flight and there is no genital contact. The reason could be
the male waiting for the female for the readiness of oviposition. By holding the
female he can make sure she does not mate with other male before laying eggs.
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Guarding Oviposition
Oviposition takes place usually immediately after copulation. The male often accompanies the female when she oviposits. Eggs are laid in
two general ways, some species females,
they have their sharp ovipositor which can cut
into plants and deposit eggs into there. Some species do not have their sharp
ovipositor and simply drop their eggs into waters.
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- There are different ways of male accompaniment, some species are still in tandem, with the
female laying eggs into the plants materials, using her sharp ovipositor to
cut open the plants materials. This is common in many damselflies and
some Hawker Dragonflies species.
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- Some species in tandem with the female dipping her abdomen tip quickly into the
water surface, wash away the eggs by water. This is can be seen in some skimmer
dragonflies.
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- Some species the male hovers around and guard the female when she is laying
eggs, which is common in many territorial skimmers, such as the Fiery Skimmer.
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- However, some species female may oviposits alone. This is common in
most Hawker and some damselflies.
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- The adult male and female may live 7-10 weeks. A female may lay several
batches of eggs. Each batch she may lay a few hundreds to a few thousands
eggs, depends on species. Those eggs may hatch after 5-40 days, depend on
species, temperature and other conditions. Generally there are about
10-15 instars stages, may take one to ten years living in waters, depends on
species. Then they emergence and become adults. A new life
cycle is started.
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How the complex mating behaviour pattern in dragonflies and
damselflies evolved?
There is still not known exactly why the dragonfly male have two sets of
reproduction organs and why they evolved such a complex mating behaviour.
The secondary genitalia is used as the intromittent organ
during copulation and have similar function as the pedipalps of a male spiders
introduce sperm into females.
All predator males have a common problem - how to avoid being
eaten by the females, before during and after mating? To a female point of
view, there is benefit and no disadvantage to prey on the male unless the male
has contribution to bring up their offspring.
Male dragonflies and damselflies successfully solved this
problem by clasping the female with their anal appendages before, during and
after mating. We seldom see male eaten by female. This could be one of the
reasons that they evolved such a complex mating behaviour.
Male spiders also solved this problem, although not as successful
as
the dragonflies. As we know male spiders are still sometime eaten by female.
Male spiders evolved their pedipalps to hold their sperm so that they can run
away a bit more easier after mating.
Praying Mantids, the relatively young order compares with
spiders and dragonflies, did not have enough time to evolve a solution yet.
So praying mantids male is known to be eaten by female from time to time.
This could be the reason why some assassin bug species breed
parthenogentically, i.e., reproduce by female without mating. And some assassin
bug species male help to look after their young.
- Reference:
- 1. Dragonflies - NATURALISTS' HANDBOOKS 7, Peter L Miller, Richmond Publishing, 1987
- 2. The Australian Dragonflies - CSIRO, Watson, Theisinger &
Abbey,1991, p7..
- 3. Field Guide to Dragonflies of Hong Kong - Keith DP Wilson, Cosmos Books, 2003,
p27.
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