Dragonfly

A dragonfly is a flying insect belonging to

the infraorder Anisoptera below the order
Odonata. About 3,000 extant species of
true dragonflies are known. Most are
tropical, with fewer species in temperate
regions. Loss of wetland habitat threatens
dragonfly populations around the world.
Adult dragonflies are characterized by a
pair of large, multifaceted, compound
eyes, two pairs of strong, transparent
wings, sometimes with coloured patches,
and an elongated body. Many dragonflies
have brilliant iridescent or metallic colours
produced by structural colouration, making
them conspicuous in flight. An adult
dragonfly's compound eyes have nearly
24,000 ommatidia each.
 Dragonfly
Temporal range: Early Jurassic to Recent

Yellow-winged darter
Sympetrum flaveolum

Scientific classification

Kingdom: Animalia

Phylum: Arthropoda

Class: Insecta

Order: Odonata

Suborder: Epiprocta
Infraorder: Anisoptera
Selys, 1854[1]

Families

Aeshnoidea
Aeshnidae (hawkers or darners)
Austropetaliidae
Gomphidae (clubtails)
Petaluridae (petaltails)
Cordulegastroidea
Chlorogomphidae
Cordulegastridae (spiketails)
Neopetaliidae
Libelluloidea
Libellulidae (skimmers, etc)
 Corduliidae (skimmers, etc)
Macromiidae (cruisers)
Synthemistidae (tigertails)
$Not a clade

Dragonflies can be mistaken for the

closely related damselflies, which make up
the other odonatan infraorder (Zygoptera)
and are similar in body plan, though usually
lighter in build; however, the wings of most
dragonflies are held flat and away from the
body, while damselflies hold their wings
folded at rest, along or above the
abdomen. Dragonflies are agile fliers, while
damselflies have a weaker, fluttery flight.
Dragonflies are predatory insects, both in
their aquatic nymphal stage (also known
as "naiads") and as adults. In some
species, the nymphal stage lasts up to five
years, and the adult stage may be as long
as 10 weeks, but most species have an
adult lifespan in the order of five weeks or
less, and some survive for only a few
days.[2] They are fast, agile fliers capable
of highly accurate aerial ambush,
sometimes migrating across oceans, and
often live near water. They have a uniquely
complex mode of reproduction involving
indirect insemination, delayed fertilization,
and sperm competition. During mating, the
male grasps the female at the back of the
head, and the female curls her abdomen
under her body to pick up sperm from the
male's secondary genitalia at the front of
his abdomen, forming the "heart" or
"wheel" posture.

Fossils of very large dragonfly-like insects,

sometimes called griffinflies, are found
from 325 million years ago (Mya) in Upper
Carboniferous rocks; these had wingspans
up to about 750 mm (30 in), though they
were only distant relatives, not true
dragonflies.

Dragonflies are represented in human

culture on artefacts such as pottery, rock
paintings, statues, and Art Nouveau
jewellery. They are used in traditional
medicine in Japan and China, and caught
for food in Indonesia. They are symbols of
courage, strength, and happiness in Japan,
but seen as sinister in European folklore.
Their bright colours and agile flight are
admired in the poetry of Lord Tennyson
and the prose of H. E. Bates.

Etymology
The infraorder Anisoptera comes from
Greek ἄνισος anisos "unequal"[3] and
πτερόν pteron "wing"[4] because
dragonflies' hindwings are broader than
their forewings.[5]
Evolution

The giant Upper Carboniferous dragonfly relative, Meganeura monyi, attained a wingspan around 680 mm (27 in).[6]
Museum of Toulouse

Mesurupetala, Late Jurassic (Tithonian), Solnhofen limestone, Germany

Dragonflies in Oze National Park

Dragonflies and their relatives are similar in

structure to an ancient group, the
Meganisoptera, from the 325 Mya Upper
Carboniferous of Europe, a group that
included the largest insect that ever lived,
Meganeuropsis permiana from the Early
Permian, with a wingspan around 750 mm
(30 in).[7] Known informally as "griffinflies",
their fossil record ends with the Permian–
Triassic extinction event (about 247 Mya).
The Protanisoptera, another ancestral
group that lacks certain wing-vein
characters found in modern Odonata, lived
from the Early to Late Permian age until
the end Permian event, and are known
from fossil wings from current-day United
States, Russia, and Australia, suggesting
they might have been cosmopolitan in
distribution. While both of those groups
are sometimes referred to as "giant
dragonflies", in fact true
dragonflies/odonata are more modern
insects that had not evolved yet.

Modern dragonflies do retain some traits

of their distant predecessors, and are in a
group known as the Palaeoptera, ancient-
winged. They, like the gigantic predinosaur
griffinflies, lack the ability to fold their
wings up against their bodies in the way
modern insects do, although some evolved
their own different way to do so. The
forerunners of modern Odonata are
included in a clade called the Panodonata,
which include the basal Zygoptera
(damselflies) and the Anisoptera (true
dragonflies).[8] Today, some 3,000 species
are extant around the world.[9][10]

The relationships of anisopteran families

are not fully resolved as of 2013, but all
the families are monophyletic except the
Corduliidae; the Gomphidae are a sister
taxon to all other Anisoptera, the
Austropetaliidae are sister to the
Aeshnoidea, and the Chlorogomphidae are
sister to a clade that includes the
Synthemistidae and Libellulidae.[11] On the
cladogram, dashed lines indicate
unresolved relationships; English names
are given (in parentheses):
 ⁠Gomphidae (clubtails)
⁠
⁠Austropetaliidae
⁠
⁠ ⁠Aeshnoidea (hawkers)
⁠⁠
⁠Petaluridae (petaltails)
⁠
⁠Macromiidae (cruisers)
⁠
⁠Neopetaliidae
⁠
⁠Anisoptera ⁠Libelluloidea
⁠ ⁠ ⁠Cordulegastridae
⁠
⁠ ⁠(goldenrings)
⁠ Libellulidae
⁠ ⁠
⁠ ⁠ ⁠ (skimmers)
⁠⁠⁠⁠
⁠⁠ "Corduliidae" [not
⁠ a clade]
⁠ (emeralds)

⁠Synthemistidae (tigertails)
⁠
⁠Chlorogomphidae
⁠
Distribution and diversity
About 3,012 species of dragonflies were
known in 2010; these are classified into
348 genera in 11 families. The distribution
of diversity within the biogeographical
regions are summarized below (the world
numbers are not ordinary totals, as
overlaps in species occur).[12]
 Family Oriental Neotropical Australasian Afrotropical Palaearctic Nearctic

Aeshnidae 149 129 78 44 58 40

Aust ropet aliidae 7 4

Pet aluridae 1 6 1 2

Gomphidae 364 277 42 152 127 101

Chlorogomphidae 46 5

Cordulegast ridae 23 1 18

Neopet aliidae 1

Corduliidae 23 20 33 6 18 51

Libellulidae 192 354 184 251 120 105

Macromiidae 50 2 17 37 7 10

Synt hemist idae 37

Incertae sedis 37 24 21 15 2

An aggregation of globe skimmers, Pantala flavescens, during migration

Dragonflies live on every continent except
Antarctica. In contrast to the damselflies
(Zygoptera), which tend to have restricted
distributions, some genera and species are
spread across continents. For example,
the blue-eyed darner Rhionaeschna
multicolor lives all across North America,
and in Central America;[13] emperors Anax
live throughout the Americas from as far
north as Newfoundland to as far south as
Bahia Blanca in Argentina,[14] across
Europe to central Asia, North Africa, and
the Middle East.[15] The globe skimmer
Pantala flavescens is probably the most
widespread dragonfly species in the world;
it is cosmopolitan, occurring on all
continents in the warmer regions. Most
Anisoptera species are tropical, with far
fewer species in temperate regions.[16]

Some dragonflies, including libellulids and

aeshnids, live in desert pools, for example
in the Mojave Desert, where they are active
in shade temperatures between 18 and
45 °C (64 and 113 °F); these insects were
able to survive body temperatures above
the thermal death point of insects of the
same species in cooler places.[17]

Dragonflies live from sea level up to the

mountains, decreasing in species diversity
with altitude.[18] Their altitudinal limit is
about 3700 m, represented by a species of
Aeshna in the Pamirs.[19]

Dragonflies become scarce at higher

latitudes. They are not native to Iceland,
but individuals are occasionally swept in
by strong winds, including a Hemianax
ephippiger native to North Africa, and an
unidentified darter species.[20] In
Kamchatka, only a few species of
dragonfly including the treeline emerald
Somatochlora arctica and some aeshnids
such as Aeshna subarctica are found,
possibly because of the low temperature
of the lakes there.[21] The treeline emerald
also lives in northern Alaska, within the
Arctic Circle, making it the most northerly
of all dragonflies.[22]

General description

Damselflies, like this Ischnura senegalensis, are slenderer in build than dragonflies, and most hold their wings closed over
their bodies.

Dragonflies (suborder Anisoptera) are

heavy-bodied, strong-flying insects that
hold their wings horizontally both in flight
and at rest. By contrast, damselflies
(suborder Zygoptera) have slender bodies
and fly more weakly; most species fold
their wings over the abdomen when
stationary, and the eyes are well separated
on the sides of the head.[12][23]

An adult dragonfly has three distinct

segments, the head, thorax, and abdomen,
as in all insects. It has a chitinous
exoskeleton of hard plates held together
with flexible membranes. The head is large
with very short antennae. It is dominated
by the two compound eyes, which cover
most of its surface. The compound eyes
are made up of ommatidia, the numbers
being greater in the larger species. Aeshna
interrupta has 22650 ommatidia of two
varying sizes, 4500 being large. The facets
facing downward tend to be smaller.
Petalura gigantea has 23890 ommatidia of
just one size. These facets provide
complete vision in the frontal hemisphere
of the dragonfly.[24] The compound eyes
meet at the top of the head (except in the
Petaluridae and Gomphidae, as also in the
genus Epiophlebia). Also, they have three
simple eyes or ocelli. The mouthparts are
adapted for biting with a toothed jaw; the
flap-like labrum, at the front of the mouth,
can be shot rapidly forward to catch
prey.[25][26] The head has a system for
locking it in place that consists of muscles
and small hairs on the back of the head
that grip structures on the front of the first
thoracic segment. This arrester system is
unique to the Odonata, and is activated
when feeding and during tandem flight.[12]

Anatomy of a dragonfly

The thorax consists of three segments as

in all insects. The prothorax is small and
flattened dorsally into a shield-like disc,
which has two transverse ridges. The
mesothorax and metathorax are fused into
a rigid, box-like structure with internal
bracing, and provide a robust attachment
for the powerful wing muscles inside.[27]
The thorax bears two pairs of wings and
three pairs of legs. The wings are long,
veined, and membranous, narrower at the
tip and wider at the base. The hindwings
are broader than the forewings and the
venation is different at the base.[28] The
veins carry haemolymph, which is
analogous to blood in vertebrates, and
carries out many similar functions, but
which also serves a hydraulic function to
expand the body between nymphal stages
(instars) and to expand and stiffen the
wings after the adult emerges from the
final nymphal stage. The leading edge of
each wing has a node where other veins
join the marginal vein, and the wing is able
to flex at this point. In most large species
of dragonflies, the wings of females are
shorter and broader than those of
males.[26] The legs are rarely used for
walking, but are used to catch and hold
prey, for perching, and for climbing on
plants. Each has two short basal joints,
two long joints, and a three-jointed foot,
armed with a pair of claws. The long leg
joints bear rows of spines, and in males,
one row of spines on each front leg is
modified to form an "eyebrush", for
cleaning the surface of the compound
eye.[27]

Migrant hawker, Aeshna mixta, has the long, slender abdomen of aeshnid dragonflies.

The abdomen is long and slender and

consists of 10 segments. Three terminal
appendages are on segment 10; a pair of
superiors (claspers) and an inferior. The
second and third segments are enlarged,
and in males, on the underside of the
second segment has a cleft, forming the
secondary genitalia consisting of the
lamina, hamule, genital lobe, and penis.
There are remarkable variations in the
presence and the form of the penis and the
related structures, the flagellum, cornua,
and genital lobes. Sperm is produced at
the 9th segment, and is transferred to the
secondary genitalia prior to mating. The
male holds the female behind the head
using a pair of claspers on the terminal
segment. In females, the genital opening is
on the underside of the eighth segment,
and is covered by a simple flap (vulvar
lamina) or an ovipositor, depending on
species and the method of egg-laying.
Dragonflies having simple flaps shed the
eggs in water, mostly in flight. Dragonflies
having ovipositors use them to puncture
soft tissues of plants and place the eggs
singly in each puncture they
make.[27][29][30][31]

Dragonfly nymphs vary in form with

species, and are loosely classed into
claspers, sprawlers, hiders, and
burrowers.[12] The first instar is known as a
prolarva, a relatively inactive stage from
which it quickly moults into the more
active nymphal form.[32] The general body
plan is similar to that of an adult, but the
nymph lacks wings and reproductive
organs. The lower jaw has a huge,
extensible labium, armed with hooks and
spines, which is used for catching prey.
This labium is folded under the body at
rest and struck out at great speed by
hydraulic pressure created by the
abdominal muscles.[12] Both damselsfly
and dragonfly nymphs ventilate the rectum,
but just some damselfly nymphs have a
rectal epithelium that is rich in trachea,
relying mostly on three feathery external
gills as their major source of respiration.
Only dragonfly nymphs have internal gills,
called a branchial chamber, located
around the fourth and fifth abdominal
segments. These internal gills consist
originally of six longidudinal folds, each
side supported by cross-folds. But this
system has been modified in several
families. Water is pumped in and out of the
abdomen through an opening at the tip.
The naiads of some clubtails (Gomphidae)
that burrow into the sediment, have a
snorkel-like tube at the end of the
abdomen enabling them to draw in clean
water while they are buried in mud. Naiads
can forcefully expel a jet of water to propel
themselves with great rapidity.[33][34][35]
Coloration

Iridescent structural colouration in a dragonfly's eyes

Many adult dragonflies have brilliant

iridescent or metallic colours produced by
structural colouration, making them
conspicuous in flight. Their overall
coloration is often a combination of
yellow, red, brown, and black pigments,
with structural colours. Blues are typically
created by microstructures in the cuticle
that reflect blue light. Greens often
combine a structural blue with a yellow
pigment. Freshly emerged adults, known
as tenerals, are often pale, and obtain their
typical colours after a few days.[28] Some
have their bodies covered with a pale blue,
waxy powderiness called pruinosity; it
wears off when scraped during mating,
leaving darker areas.[36]

Male green darner, Anax junius has noniridescent structural blue; the female (below) lacks the colour.
Some dragonflies, such as the green
darner, Anax junius, have a noniridescent
blue that is produced structurally by
scatter from arrays of tiny spheres in the
endoplasmic reticulum of epidermal cells
underneath the cuticle.[37]

The wings of dragonflies are generally

clear, apart from the dark veins and
pterostigmata. In the chasers
(Libellulidae), however, many genera have
areas of colour on the wings: for example,
groundlings (Brachythemis) have brown
bands on all four wings, while some
scarlets (Crocothemis) and dropwings
(Trithemis) have bright orange patches at
the wing bases. Some aeshnids such as
the brown hawker (Aeshna grandis) have
translucent, pale yellow wings.[38]

Dragonfly nymphs are usually a well-

camouflaged blend of dull brown, green,
and grey.[33]

Biology

Ecology

Dragonflies and damselflies are predatory

both in the aquatic nymphal and adult
stages. Nymphs feed on a range of
freshwater invertebrates and larger ones
can prey on tadpoles and small fish.[39]
One species, Gomphus militaris, even live
as a parasite, feeding on the gills of gravid
mussels.[40] Adults capture insect prey in
the air, making use of their acute vision
and highly controlled flight. The mating
system of dragonflies is complex, and they
are among the few insect groups that have
a system of indirect sperm transfer along
with sperm storage, delayed fertilization,
and sperm competition.[39]

Adult males vigorously defend territories

near water; these areas provide suitable
habitat for the nymphs to develop, and for
females to lay their eggs. Swarms of
feeding adults aggregate to prey on
swarming prey such as emerging flying
ants or termites.[39]

Habitat preference: A four-spotted chaser, Libellula quadrimaculata on an emergent plant, the water violet Hottonia
palustris, with submerged vegetation in the background

Dragonflies as a group occupy a

considerable variety of habitats, but many
species, and some families, have their own
specific environmental requirements.[41]
Some species prefer flowing waters, while
others prefer standing water. For example,
the Gomphidae (clubtails) live in running
water, and the Libellulidae (skimmers) live
in still water.[41] Some species live in
temporary water pools and are capable of
tolerating changes in water level,
desiccation, and the resulting variations in
temperature, but some genera such as
Sympetrum (darters) have eggs and
nymphs that can resist drought and are
stimulated to grow rapidly in warm,
shallow pools, also often benefiting from
the absence of predators there.[41]
Vegetation and its characteristics
including submerged, floating, emergent, or
waterside are also important. Adults may
require emergent or waterside plants to
use as perches; others may need specific
submerged or floating plants on which to
lay eggs. Requirements may be highly
specific, as in Aeshna viridis (green
hawker), which lives in swamps with the
water-soldier, Stratiotes aloides.[41] The
chemistry of the water, including its trophic
status (degree of enrichment with
nutrients) and pH can also affect its use
by dragonflies. Most species need
moderate conditions, not too eutrophic,
not too acidic;[41] a few species such as
Sympetrum danae (black darter) and
Libellula quadrimaculata (four-spotted
chaser) prefer acidic waters such as peat
bogs,[42] while others such as Libellula
fulva (scarce chaser) need slow-moving,
eutrophic waters with reeds or similar
waterside plants.[43][44]

Behaviour

Many dragonflies, particularly males, are

territorial. Some defend a territory against
others of their own species, some against
other species of dragonfly and a few
against insects in unrelated groups. A
particular perch may give a dragonfly a
good view over an insect-rich feeding
ground; males of many species such as
the Pachydiplax longipennis (blue dasher)
jostle other dragonflies to maintain the
right to alight there.[45] Defending a
breeding territory is common among male
dragonflies, especially in species that
congregate around ponds. The territory
contains desirable features such as a
sunlit stretch of shallow water, a special
plant species, or the preferred substrate
for egg-laying. The territory may be small
or large, depending on its quality, the time
of day, and the number of competitors,
and may be held for a few minutes or
several hours. Dragonflies including
Tramea lacerata (black saddlebags) may
notice landmarks that assist in defining the
boundaries of the territory. Landmarks
may reduce the costs of territory
establishment, or might serve as a spatial
reference.[46] Some dragonflies signal
ownership with striking colours on the
face, abdomen, legs, or wings. The
Plathemis lydia (common whitetail) dashes
towards an intruder holding its white
abdomen aloft like a flag. Other
dragonflies engage in aerial dogfights or
high-speed chases. A female must mate
with the territory holder before laying her
eggs.[45] There is also conflict between the
males and females. Females may
sometimes be harassed by males to the
extent that it affects their normal activities
including foraging and in some dimorphic
species females have evolved multiple
forms with some forms appearing
deceptively like males.[47] In some species
females have evolved behavioural
responses such as feigning death to
escape the attention of males.[48] Similarly,
selection of habitat by adult dragonflies is
not random, and terrestrial habitat patches
may be held for up to 3 months. A species
tightly linked to its birth site utilises a
foraging area that is several orders of
magnitude larger than the birth site.[49]
Reproduction

Mating pair of marsh skimmers, Orthetrum luzonicum, forming a "heart"

Mating in dragonflies is a complex,

precisely choreographed process. First,
the male has to attract a female to his
territory, continually driving off rival males.
When he is ready to mate, he transfers a
packet of sperm from his primary genital
opening on segment 9, near the end of his
abdomen, to his secondary genitalia on
segments 2–3, near the base of his
abdomen. The male then grasps the
female by the head with the claspers at the
end of his abdomen; the structure of the
claspers varies between species, and may
help to prevent interspecific mating.[50] The
pair flies in tandem with the male in front,
typically perching on a twig or plant stem.
The female then curls her abdomen
downwards and forwards under her body
to pick up the sperm from the male's
secondary genitalia, while the male uses
his "tail" claspers to grip the female behind
the head: this distinctive posture is called
the "heart" or "wheel";[39][51] the pair may
also be described as being "in cop".[52]

Egg-laying (ovipositing) involves not only

the female darting over floating or
waterside vegetation to deposit eggs on a
suitable substrate, but also the male
hovering above her or continuing to clasp
her and flying in tandem. The male
attempts to prevent rivals from removing
his sperm and inserting their own,[53]
something made possible by delayed
fertilisation[39][51] and driven by sexual
selection.[50] If successful, a rival male
uses his penis to compress or scrape out
the sperm inserted previously; this activity
takes up much of the time that a
copulating pair remains in the heart
posture.[54] Flying in tandem has the
advantage that less effort is needed by the
female for flight and more can be
expended on egg-laying, and when the
female submerges to deposit eggs, the
male may help to pull her out of the
water.[53]

Egg-laying takes two different forms

depending on the species. The female in
some families (Aeshnidae, Petaluridae)
has a sharp-edged ovipositor with which
she slits open a stem or leaf of a plant on
or near the water, so she can push her
eggs inside. In other families such as
clubtails (Gomphidae), cruisers
(Macromiidae), emeralds (Corduliidae),
and skimmers (Libellulidae), the female
lays eggs by tapping the surface of the
water repeatedly with her abdomen, by
shaking the eggs out of her abdomen as
she flies along, or by placing the eggs on
vegetation.[54] In a few species, the eggs
are laid on emergent plants above the
water, and development is delayed until
these have withered and become
immersed.[33]
Life cycle

Nymph of emperor dragonfly, Anax imperator

Illustration of a naiad with mask extended

Dragonflies are hemimetabolous insects;

they do not have a pupal stage and
undergo an incomplete metamorphosis
with a series of nymphal stages from
which the adult emerges.[55] Eggs laid
inside plant tissues are usually shaped like
grains of rice, while other eggs are the size
of a pinhead, ellipsoidal, or nearly
spherical. A clutch may have as many as
1500 eggs, and they take about a week to
hatch into aquatic nymphs or naiads which
moult between six and 15 times
(depending on species) as they grow.[12]
Most of a dragonfly's life is spent as a
nymph, beneath the water's surface. The
nymph extends its hinged labium (a
toothed mouthpart similar to a lower
mandible, which is sometimes termed as a
"mask" as it is normally folded and held
before the face) that can extend forward
and retract rapidly to capture prey such as
mosquito larvae, tadpoles, and small
fish.[55] They breathe through gills in their
rectum, and can rapidly propel themselves
by suddenly expelling water through the
anus.[56] Some naiads, such as the later
stages of Antipodophlebia asthenes, hunt
on land.[57]

Ecdysis: Emperor dragonfly, Anax imperator, newly emerged and still soft, holding on to its dry exuvia, and expanding its
wings
The nymph stage of dragonflies lasts up to
five years in large species, and between
two months and three years in smaller
species. When the naiad is ready to
metamorphose into an adult, it stops
feeding and makes its way to the surface,
generally at night. It remains stationary
with its head out of the water, while its
respiration system adapts to breathing air,
then climbs up a reed or other emergent
plant, and moults (ecdysis). Anchoring
itself firmly in a vertical position with its
claws, its exoskeleton begins to split at a
weak spot behind the head. The adult
dragonfly crawls out of its nymph
exoskeleton, the exuvia, arching
backwards when all but the tip of its
abdomen is free, to allow its exoskeleton
to harden. Curling back upwards, it
completes its emergence, swallowing air,
which plumps out its body, and pumping
haemolymph into its wings, which causes
them to expand to their full extent.[58]

Dragonflies in temperate areas can be

categorized into two groups, an early
group and a later one. In any one area,
individuals of a particular "spring species"
emerge within a few days of each other.
The springtime darner (Basiaeschna
janata), for example, is suddenly very
common in the spring, but disappears a
few weeks later and is not seen again until
the following year. By contrast, a "summer
species" emerges over a period of weeks
or months, later in the year. They may be
seen on the wing for several months, but
this may represent a whole series of
individuals, with new adults hatching out
as earlier ones complete their lifespans.[59]

Sex ratios

The sex ratio of male to female

dragonflies varies both temporally and
spatially. Adult dragonflies have a high
male-biased ratio at breeding habitats. The
male-bias ratio has contributed partially to
the females using different habitats to
avoid male harassment. As seen in Hine's
emerald dragonfly (Somatochlora hineana),
male populations use wetland habitats,
while females use dry meadows and
marginal breeding habitats, only migrating
to the wetlands to lay their eggs or to find
mating partners. Unwanted mating is
energetically costly for females because it
affects the amount of time that they are
able to spend foraging.[60]
Brown hawker, Aeshna grandis in flight: The hindwings are about 90° out of phase with the forewings at this instant,
suggesting fast flight.

Flight

Red-veined darters (Sympetrum fonscolombii) flying "in cop" (male ahead)

Dragonflies are powerful and agile fliers,

capable of migrating across the sea,
moving in any direction, and changing
direction suddenly. In flight, the adult
dragonfly can propel itself in six directions:
upward, downward, forward, backward, to
left and to right.[61] They have four
different styles of flight.[62]

Counter-stroking, with forewings beating

180° out of phase with the hindwings, is
used for hovering and slow flight. This
style is efficient and generates a large
amount of lift.
Phased-stroking, with the hindwings
beating 90° ahead of the forewings, is
used for fast flight. This style creates
more thrust, but less lift than counter-
stroking
Synchronised-stroking, with forewings
and hindwings beating together, is used
when changing direction rapidly, as it
maximises thrust
Gliding, with the wings held out, is used
in three situations: free gliding, for a few
seconds in between bursts of powered
flight; gliding in the updraft at the crest
of a hill, effectively hovering by falling at
the same speed as the updraft; and in
certain dragonflies such as darters,
when "in cop" with a male, the female
sometimes simply glides while the male
pulls the pair along by beating his
wings.[62]
Southern hawker, Aeshna cyanea: its wings at this instant are synchronised for agile flight.

The wings are powered directly, unlike

most families of insects, with the flight
muscles attached to the wing bases.
Dragonflies have a high power/weight
ratio, and have been documented
accelerating at 4 G linearly and 9 G in
sharp turns while pursuing prey.[62]

Dragonflies generate lift in at least four

ways at different times, including classical
lift like an aircraft wing; supercritical lift
with the wing above the critical angle,
generating high lift and using very short
strokes to avoid stalling; and creating and
shedding vortices. Some families appear
to use special mechanisms, as for
example the Libellulidae which take off
rapidly, their wings beginning pointed far
forward and twisted almost vertically.
Dragonfly wings behave highly dynamically
during flight, flexing and twisting during
each beat. Among the variables are wing
curvature, length and speed of stroke,
angle of attack, forward/back position of
wing, and phase relative to the other
wings.[62]
Flight speed

Old and unreliable claims are made that

dragonflies such as the southern giant
darner can fly up to 97 km/h (60 mph).[63]
However, the greatest reliable flight speed
records are for other types of insects.[64] In
general, large dragonflies like the hawkers
have a maximum speed of 36–54 km/h
(22–34 mph) with average cruising speed
of about 16 km/h (9.9 mph).[65] Dragonflies
can travel at 100 body-lengths per second
in forward flight, and three lengths per
second backwards.[25]
Motion camouflage

The principle of motion camouflage

In high-speed territorial battles between

male Australian emperors (Hemianax
papuensis), the fighting dragonflies adjust
their flight paths to appear stationary to
their rivals, minimizing the chance of being
detected as they approach.[a][66][67] To
achieve the effect, the attacking dragonfly
flies towards his rival, choosing his path to
remain on a line between the rival and the
start of his attack path. The attacker thus
looms larger as he closes on the rival, but
does not otherwise appear to move.
Researchers found that six of 15
encounters involved motion
camouflage.[68]

Temperature control

The flight muscles need to be kept at a

suitable temperature for the dragonfly to
be able to fly. Being cold-blooded, they can
raise their temperature by basking in the
sun. Early in the morning, they may choose
to perch in a vertical position with the
wings outstretched, while in the middle of
the day, a horizontal stance may be
chosen. Another method of warming up
used by some larger dragonflies is wing-
whirring, a rapid vibration of the wings that
causes heat to be generated in the flight
muscles. The green darner (Anax junius) is
known for its long-distance migrations,
and often resorts to wing-whirring before
dawn to enable it to make an early
start.[69]

Becoming too hot is another hazard, and a

sunny or shady position for perching can
be selected according to the ambient
temperature. Some species have dark
patches on the wings which can provide
shade for the body, and a few use the
obelisk posture to avoid overheating. This
behaviour involves doing a "handstand",
perching with the body raised and the
abdomen pointing towards the sun, thus
minimising the amount of solar radiation
received. On a hot day, dragonflies
sometimes adjust their body temperature
by skimming over a water surface and
briefly touching it, often three times in
quick succession. This may also help to
avoid desiccation.[69]
Feeding

Common clubtail, Gomphus vulgatissimus, with prey

Adult dragonflies hunt on the wing using

their exceptionally acute eyesight and
strong, agile flight.[51] They are almost
exclusively carnivorous, eating a wide
variety of insects ranging from small
midges and mosquitoes to butterflies,
moths, damselflies, and smaller
dragonflies.[65] A large prey item is
subdued by being bitten on the head and is
carried by the legs to a perch. Here, the
wings are discarded and the prey usually
ingested head first.[70] A dragonfly may
consume as much as a fifth of its body
weight in prey per day.[71] Dragonflies are
also some of the insect world's most
efficient hunters, catching up to 95% of the
prey they pursue.[72]

The nymphs are voracious predators,

eating most living things that are smaller
than they are. Their staple diet is mostly
bloodworms and other insect larvae, but
they also feed on tadpoles and small
fish.[65] A few species, especially those
that live in temporary waters, are likely to
leave the water to feed. Nymphs of
Cordulegaster bidentata sometimes hunt
small arthropods on the ground at night,
while some species in the Anax genus have
even been observed leaping out of the
water to attack and kill full-grown tree
frogs.[12][73]

Eyesight

Dragonfly vision is thought to be like slow

motion for humans. Dragonflies see faster
than humans do; they see around 200
images per second.[74] A dragonfly can see
in 360 degrees, and nearly 80 percent of
the insect's brain is dedicated to its
sight.[75]

Predators

Southern red-billed hornbill with a captured dragonfly in its bill

Although dragonflies are swift and agile

fliers, some predators are fast enough to
catch them. These include falcons such as
the American kestrel, the merlin,[76] and the
hobby;[77] nighthawks, swifts, flycatchers
and swallows also take some adults;
some species of wasps, too, prey on
dragonflies, using them to provision their
nests, laying an egg on each captured
insect. In the water, various species of
ducks and herons eat dragonfly nymphs[76]
and they are also preyed on by newts,
frogs, fish, and water spiders.[78] Amur
falcons, which migrate over the Indian
Ocean at a period that coincides with the
migration of the globe skimmer dragonfly,
Pantala flavescens, may actually be
feeding on them while on the wing.[79]

Parasites
Dragonflies are affected by three major
groups of parasites: water mites, gregarine
protozoa, and trematode flatworms
(flukes). Water mites, Hydracarina, can kill
smaller dragonfly nymphs, and may also
be seen on adults.[80] Gregarines infect the
gut and may cause blockage and
secondary infection.[81] Trematodes are
parasites of vertebrates such as frogs,
with complex life cycles often involving a
period as a stage called a cercaria in a
secondary host, a snail. Dragonfly nymphs
may swallow cercariae, or these may
tunnel through a nymph's body wall; they
then enter the gut and form a cyst or
metacercaria, which remains in the nymph
for the whole of its development. If the
nymph is eaten by a frog, the amphibian
becomes infected by the adult or fluke
stage of the trematode.[82]

Dragonflies and humans

Conservation

Most odonatologists live in temperate

areas and the dragonflies of North
America and Europe have been the subject
of much research. However, the majority of
species live in tropical areas and have
been little studied. With the destruction of
rainforest habitats, many of these species
are in danger of becoming extinct before
they have even been named. The greatest
cause of decline is forest clearance with
the consequent drying up of streams and
pools which become clogged with silt. The
damming of rivers for hydroelectric
schemes and the drainage of low-lying
land has reduced suitable habitat, as has
pollution and the introduction of alien
species.[83]

In 1997, the International Union for

Conservation of Nature set up a status
survey and conservation action plan for
dragonflies. This proposes the
establishment of protected areas around
the world and the management of these
areas to provide suitable habitat for
dragonflies. Outside these areas,
encouragement should be given to modify
forestry, agricultural, and industrial
practices to enhance conservation. At the
same time, more research into dragonflies
needs to be done, consideration should be
given to pollution control and the public
should be educated about the importance
of biodiversity.[83]

Habitat degradation has reduced dragonfly

populations across the world, for example
in Japan.[84] Over 60% of Japan's wetlands
were lost in the 20th century, so its
dragonflies now depend largely on rice
fields, ponds, and creeks. Dragonflies feed
on pest insects in rice, acting as a natural
pest control.[85][86] Dragonflies are steadily
declining in Africa, and represent a
conservation priority.[87]

The dragonfly's long lifespan and low

population density makes it vulnerable to
disturbance, such as from collisions with
vehicles on roads built near wetlands.
Species that fly low and slow may be most
at risk.[88]

Dragonflies are attracted to shiny surfaces

that produce polarization which they can
mistake for water, and they have been
known to aggregate close to polished
gravestones, solar panels, automobiles,
and other such structures on which they
attempt to lay eggs. These can have a
local impact on dragonfly populations;
methods of reducing the attractiveness of
structures such as solar panels are under
experimentation.[89][90]

In culture

A blue-glazed faience dragonfly amulet

was found by Flinders Petrie at Lahun,
from the Late Middle Kingdom of ancient
Egypt.[91]

Many Native American tribes consider

dragonflies to be medicine animals that
had special powers. For example, the
southwestern tribes, including the Pueblo,
Hopi, and Zuni, associated dragonflies with
transformation. They referred to
dragonflies as "snake doctors" because
they believed dragonflies followed snakes
into the ground and healed them if they
were injured.[92] For the Navajo, dragonflies
symbolize pure water. Often stylized in a
double-barred cross design, dragonflies
are a common motif in Zuni pottery, as
well as Hopi rock art and Pueblo
necklaces.[93]: 20–26 

As a seasonal symbol in Japan,

dragonflies are associated with season of
autumn.[94] In Japan, they are symbols of
rebirth, courage, strength, and happiness.
They are also depicted frequently in
Japanese art and literature, especially
haiku poetry. Japanese children catch
large dragonflies as a game, using a hair
with a small pebble tied to each end, which
they throw into the air. The dragonfly
mistakes the pebbles for prey, gets
tangled in the hair, and is dragged to the
ground by the weight.[93]: 38 

In Chinese culture, dragonflies symbolize

both change and instability. They are also
symbols in the Chinese practices of Feng
Shui, where placements of dragonfly
statues and artwork in parts of a home or
office are believed to bring new insights
and positive changes.[95]

In both China and Japan, dragonflies have

been used in traditional medicine. In
Indonesia, adult dragonflies are caught on
poles made sticky with birdlime, then fried
in oil as a delicacy.[96]

Images of dragonflies are common in Art

Nouveau, especially in jewellery
designs.[97] They have also been used as a
decorative motif on fabrics and home
furnishings.[98] Douglas, a British
motorcycle manufacturer based in Bristol,
named its innovatively designed postwar
350-cc flat-twin model the Dragonfly.[99]

Among the classical names of Japan are

秋津国), Akitsushima (秋津島),
Akitsukuni (
Toyo-akitsushima (豊秋津島). Akitsu is an
old word for dragonfly, so one
interpretation of Akitsushima is "Dragonfly
Island".[100] This is attributed to a legend in
which Japan's mythical founder, Emperor
Jimmu, was bitten by a mosquito, which
was then eaten by a dragonfly.[101][102]

In Europe, dragonflies have often been

seen as sinister. Some English vernacular
names, such as "horse-stinger",[103] "devil's
darning needle", and "ear cutter", link them
with evil or injury.[104] Swedish folklore
holds that the devil uses dragonflies to
weigh people's souls.[93]: 25–27  The
Norwegian name for dragonflies is
Øyenstikker ("eye-poker"), and in Portugal,
they are sometimes called tira-olhos
("eyes-snatcher"). They are often
associated with snakes, as in the Welsh
name gwas-y-neidr, "adder's servant".[104]
The Southern United States terms "snake
doctor" and "snake feeder" refer to a folk
belief that dragonflies catch insects for
snakes or follow snakes around and stitch
them back together if they are
injured.[105][106] Interestingly, the Hungarian
name for dragonfly is szitakötő ("sieve-
knitter").

The watercolourist Moses Harris (1731–

1785), known for his The Aurelian or natural
history of English insects (1766), published
in 1780, the first scientific descriptions of
several Odonata including the banded
demoiselle, Calopteryx splendens. He was
the first English artist to make illustrations
of dragonflies accurate enough to be
identified to species (Aeshna grandis at
top left of plate illustrated), though his
rough drawing of a nymph (at lower left)
with the mask extended appears to be
plagiarised.[b][107]
More recently, dragonfly watching has
become popular in America as some
birdwatchers seek new groups to
observe.[108]

In heraldry, like other winged insects, the

dragonfly is typically depicted tergiant
(with its back facing the viewer), with its
head to chief.[109]
Dragonfly symbol on a Hopi bowl from
Sikyátki, Arizona, c. 1400–1625
 Accurately drawn dragonflies by Moses
Harris, 1780: At top left, the brown hawker,
Aeshna grandis (described by Linnaeus,
1758); the nymph at lower left is shown with
the "mask" extended.
Woodcut on paper, after Kitagawa Utamaro,
1788

Tiffany & Co. Japonism vase with dragonfly

handles, c. 1879, Walters Art Museum
Tiffany dragonfly pendant lamp, designed c.
1903
 Japanese tsuba with a dragonfly, 1931:
Shibuichi with gold and silver, Walters Art
Museum

In poetry and literature

Lafcadio Hearn wrote in his 1901 book A

Japanese Miscellany that Japanese poets
had created dragonfly haiku "almost as
numerous as are the dragonflies
themselves in the early autumn."[110] The
poet Matsuo Bashō (1644–1694) wrote
haiku such as "Crimson pepper pod / add
two pairs of wings, and look / darting
dragonfly", relating the autumn season to
the dragonfly.[111] Hori Bakusui (1718–
1783) similarly wrote "Dyed he is with the /
Colour of autumnal days, / O red
dragonfly."[110]

The poet Lord Tennyson, described a

dragonfly splitting its old skin and
emerging shining metallic blue like
"sapphire mail" in his 1842 poem "The Two
Voices", with the lines "An inner impulse
rent the veil / Of his old husk: from head to
tail / Came out clear plates of sapphire
mail."[112]

The novelist H. E. Bates described the

rapid, agile flight of dragonflies in his 1937
nonfiction book[113] Down the River:[114]

I saw, once, an endless

procession, just over an area of
water-lilies, of small sapphire
dragonflies, a continuous play of
blue gauze over the snowy
flowers above the sun-glassy
water. It was all confined, in
true dragonfly fashion, to one
 small space. It was a continuous
turning and returning, an
endless darting, poising,
striking and hovering, so swift
that it was often lost in
sunlight.[115]

In technology

A dragonfly has been genetically modified

with light-sensitive "steering neurons" in its
nerve cord to create a cyborg-like
"DragonflEye". The neurons contain genes
like those in the eye to make them
sensitive to light. Miniature sensors, a
computer chip and a solar panel were
fitted in a "backpack" over the insect's
thorax in front of its wings. Light is sent
down flexible light-pipes named
optrodes[c] from the backpack into the
nerve cord to give steering commands to
the insect. The result is a "micro-aerial
vehicle that's smaller, lighter and stealthier
than anything else that's
manmade".[116][117]

Explanatory notes
a. This is not to say that other species may
not use the same technique, only that this
species has been studied.
b. Reviewing his artwork, the odonatologists
Albert Orr and Matti Hämäläinen comment
that his drawing of a 'large brown' (Aeshna
grandis, top left of image) was "superb",
while the "perfectly natural colours of the
eyes indicate that Harris had examined
living individuals of these aeshnids and
either coloured the printed copper plates
himself or supervised the colourists."
However, they consider the nymph on the
same plate far less good, "a very stiff
dorso-lateral view of an aeshnid larva with
mask extended. No attempt has been
made to depict the eyes, antennae or hinge
on the mask or labial palps, all
inconceivable omissions for an artist of
Harris' talent had he actually examined a
 specimen", and they suggest he copied it
from August Johann Rösel von
Rosenhof.[107]
c. Optrode is a portmanteau of "optical
electrode".

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Berger, Cynthia (2004). Dragonflies (http

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External links
The dictionary definition of dragonfly at
Wiktionary
Media related to Anisoptera at
Wikimedia Commons
Data related to Anisoptera at
Wikispecies

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