Dragonfly

A dragonfly is a flying insect belonging to the order Odonata,

infraorder Anisoptera (from Greek ἄνισος anisos, "unequal" Dragonfly
and πτερόν pteron, "wing", because the hindwing is broader Temporal range: Early Jurassic to
than the forewing). Adult dragonflies are characterized by a pair Recent
of large, multifaceted compound eyes, two pairs of strong,
transparent wings, sometimes with coloured patches, and an
elongated body. 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. Many Yellow-winged darter
dragonflies have brilliant iridescent or metallic colours produced Sympetrum flaveolum
by structural colouration, making them conspicuous in flight. An
adult dragonfly's compound eyes have nearly 24,000 ommatidia Scientific classification
each.
Kingdom: Animalia
Fossils of very large dragonfly-like insects, sometimes called Phylum: Arthropoda
griffinflies, are found from 325 million years ago (Mya) in
Upper Carboniferous rocks; these had wingspans up to about Class: Insecta
750 mm (30 in), but were only distant ancestors, not true Order: Odonata
dragonflies. About 3,000 extant species of true dragonfly are
known. Most are tropical, with fewer species in temperate Suborder: Epiprocta
regions. Loss of wetland habitat threatens dragonfly populations Infraorder: Anisoptera
around the world.
Selys, 1854 [1]
Dragonflies are predatory insects, both in their aquatic nymphs Families
stage (also known as naiads) and as adults. In some species, the
nymphal stage lasts for up to five years, and the adult stage may Aeshnoidea
be as long as ten weeks, but most species have an adult lifespan
in the order of five weeks or less, and some survive for only a Aeshnidae (hawkers or
few days.[2] They are fast, agile fliers capable of highly accurate
darners)
aerial ambush, sometimes migrating across oceans, and often live
near water. They have a uniquely complex mode of reproduction Austropetaliidae
involving indirect insemination, delayed fertilization, and sperm
Gomphidae (clubtails)
competition. During mating, the male grasps the female at the
back of the head, and the female curls her abdomen under her Petaluridae (petaltails)
body to pick up sperm from the male's secondary genitalia at the
Cordulegastroidea
front of his abdomen, forming the "heart" or "wheel" posture.
Chlorogomphidae
Dragonflies are represented in human culture on artefacts such as
pottery, rock paintings, statues and Art Nouveau jewellery. They Cordulegastridae
are used in traditional medicine in Japan and China, and caught (spiketails)
for food in Indonesia. They are symbols of courage, strength,
Neopetaliidae
and happiness in Japan, but seen as sinister in European folklore. Libelluloidea
Their bright colours and agile flight are admired in the poetry of
Lord Tennyson and the prose of H. E. Bates. Corduliidae (emeralds)$
Libellulidae (skimmers,
etc)
Contents
Macromiidae (cruisers)
Evolution
Synthemistidae (tigertails)
Distribution and diversity $Not a clade
General description
Colouration
Biology
Ecology
Behaviour
Reproduction
Life cycle
Sex ratios
Flight
Flight speed
Motion camouflage
Temperature control
Feeding
Eyesight
Predators
Parasites
Dragonflies and humans
Conservation
In culture
In poetry and literature
In technology
Notes
References
Sources
External links

Evolution
Dragonflies and their relatives are similar in structure to an ancient group, 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);[4]. 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 palaeoptera, ancient-
winged. They, like the gigantic pre-dinosaur griffinflies, lack the
ability to fold their wings up against their bodies in the way modern The giant Upper Carboniferous
insects do, although some evolved their own different way to do so. dragonfly ancestor, Meganeura
The forerunners of modern Odonata are included in a clade called monyi, attained a wingspan around
the Panodonata, which include the basal Zygoptera (damselflies) 680 mm (27 in).[3] Museum of
and the Anisoptera (true dragonflies).[5] Today, some 3,000 species Toulouse
are extant around the world.[6][7]

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.[8] On the cladogram, dashed lines
indicate unresolved relationships; English names are given (in
parentheses):
Anisoptera Mesurupetala, Late Jurassic
Gomphidae (clubtails) (Tithonian), Solnhofen limestone,
Germany

Austropetaliidae

Aeshnoidea (hawkers)

Petaluridae (petaltails)

Play media
Macromiidae (cruisers)
Dragonflies in Oze National Park

Neopetaliidae
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).[9]

Family Oriental Neotropical Australasian Afrotropical Palaearctic Nearctic Pacific World

Aeshnidae 149 129 78 44 58 40 13 456

Austropetaliidae 7 4 11
Petaluridae 1 6 1 2 10
Gomphidae 364 277 42 152 127 101 980
Chlorogomphidae 46 5 47
Cordulegastridae 23 1 18 46
Neopetaliidae 1 1
Corduliidae 23 20 33 6 18 51 12 154
Libellulidae 192 354 184 251 120 105 31 1037
Macromiidae 50 2 17 37 7 10 125
Synthemistidae 37 9 46
Incertae sedis 37 24 21 15 2 99

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;[10] emperors Anax live throughout the Americas from
as far north as Newfoundland to as far south as Bahia Blanca in
Argentina,[11] across Europe to central Asia, North Africa, and the Middle
East.[12] 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.[13]

Some dragonflies, including libellulids and aeshnids, live in desert pools, An aggregation of globe
for example in the Mojave Desert, where they are active in shade skimmers, Pantala
temperatures between 18 and 45 °C (64.4 to 113 °F); these insects were flavescens, during migration
able to survive body temperatures above the thermal death point of insects
of the same species in cooler places.[14]

Dragonflies live from sea level up to the mountains, decreasing in species diversity with altitude.[15] Their
altitudinal limit is about 3700 m, represented by a species of Aeshna in the Pamirs.[16]

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.[17] 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.[18] The treeline emerald also lives in northern Alaska, within the
Arctic Circle, making it the most northerly of all dragonflies.[19]

General description
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.[9][20]

An adult dragonfly has three distinct segments, the head, thorax,

Damselflies, like this Ischnura and abdomen, as in all insects. It has a chitinous exoskeleton of
senegalensis, are more slender in hard plates held together with flexible membranes. The head is
build than dragonflies, and most hold large with very short antennae. It is dominated by the two
their wings closed over their bodies. 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.[21] 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.[22][23] 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.[9]

The thorax consists of three segments as in all insects. The

prothorax is small and is 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.[24] 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.[25] 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.[23] The legs Head of a southern hawker
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.[24]
 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 Anatomy of a dragonfly
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,
Migrant hawker, Aeshna
mixta, has the long
and is transferred to the secondary genitalia prior to mating. The male holds
slender abdomen of
the female behind the head using a pair of claspers on the terminal segment. In
aeshnid dragonflies. 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.[24][26][27][28]

Dragonfly nymphs vary in form with species, and are loosely classed into claspers, sprawlers, hiders, and
burrowers.[9] The first instar is known as a prolarva, a relatively inactive stage from which it quickly moults
into the more active nymphal form.[29] 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.[9] Whereas damselfly nymphs have three
feathery external gills, dragonfly nymphs have internal gills, located around the fourth and fifth abdominal
segments. 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.[30]

Colouration

Many adult dragonflies have brilliant iridescent or metallic colours

produced by structural colouration, making them conspicuous in
flight. Their overall colouration 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-
coloured and obtain their typical colours after a few days,[25] some
have their bodies covered with a pale blue, waxy powderiness
Iridescent structural colouration in a
dragonfly's eyes
called pruinosity; it wears off when scraped during mating, leaving
darker areas.[31]

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.[32]
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.[33]

Dragonfly nymphs are usually a well-camouflaged blend of dull

brown, green, and grey.[30] Male green darner, Anax junius has
noniridescent structural blue; the
female (below) lacks the colour.
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.[34] 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.[34]

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.[34]

Dragonflies as a group occupy a considerable variety of habitats,

but many species, and some families, have their own specific
environmental requirements.[35] 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.[35] 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 Habitat preference: A four-spotted
pools, also often benefiting from the absence of predators there.[35] chaser, Libellula quadrimaculata on
Vegetation and its characteristics including submerged, floating, an emergent plant, the water violet
emergent, or waterside are also important. Adults may require Hottonia palustris, with submerged
emergent or waterside plants to use as perches; others may need vegetation in the background
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.[35] 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;[35] a few species
such as Sympetrum danae (black darter) and Libellula quadrimaculata (four-spotted chaser) prefer acidic
waters such as peat bogs,[36] while others such as Libellula fulva (scarce chaser) need slow-moving,
eutrophic waters with reeds or similar waterside plants.[37][38]

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.[39]
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.[40] 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.[39] 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.[41] In some species females have evolved behavioural responses such as feigning
death to escape the attention of males.[42] 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.[43]

Reproduction

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.[44] 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
Mating pair of marsh
the "heart" or "wheel";[34][45] the pair may also be described as being "in
skimmers, Orthetrum
cop".[46]
luzonicum, forming a "heart"
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,[47] something made possible by delayed
fertilisation[34][45] and driven by sexual selection.[44] 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.[48] 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.[47]

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.[48] 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.[30]

Life cycle

Dragonflies are
hemimetabolous insects; they
do not have a pupal stage and
undergo an incomplete
metamorphosis with a series of
Illustration of a naiad with mask
nymphal stages from which
extended
the adult emerges.[49] Eggs
laid inside plant tissues are
Nymph of emperor dragonfly, Anax usually shaped like grains of rice, while other eggs are the size of a
imperator 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
[9]
(depending on species) as they grow. 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.[49] They breathe
through gills in their rectum, and can rapidly propel themselves by suddenly expelling water through the
anus.[50] Some naiads, such as the later stages of Antipodophlebia asthenes, hunt on land.[51]

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 skin begins to split at a weak spot
behind the head. The adult dragonfly crawls out of its nymph skin, 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.[52]

Dragonflies in temperate areas can be categorized into two groups, an early Ecdysis: Emperor dragonfly,
group and a later one. In any one area, individuals of a particular "spring Anax imperator, newly
species" emerge within a few days of each other. The springtime darner emerged and still soft,
(Basiaeschna janata), for example, is suddenly very common in the spring, holding on to its dry exuvia,
but disappears a few weeks later and is not seen again until the following and expanding its wings
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.[53]
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.[54]

Flight

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.[55] They have four different styles of flight:[56] A number of
flying modes are used that include 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 Brown hawker, Aeshna grandis in
flight: The hindwings are about 90°
of lift; phased-stroking, with the hindwings beating 90° ahead of
out of phase with the forewings at
the forewings, is used for fast flight. This style creates more thrust,
this instant, suggesting fast flight.
but less lift than counter-stroking; synchronised-stroking, with
forewings and hindwings beating together, is used when changing
direction rapidly, as it maximises thrust; and 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.[56]

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.[56]

Southern hawker, Aeshna cyanea: its Dragonflies generate lift in at least

wings at this instant are four ways at different times,
Red-veined darters
synchronised for agile flight. including classical lift like an
(Sympetrum fonscolombii)
aircraft wing; supercritical lift with flying "in cop" (male ahead)
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.[56]

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).[57] However, the greatest reliable flight speed records are for other types of insects.[58] 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).[59] Dragonflies can travel at 100 body-lengths per
second in forward flight, and three lengths per second backwards.[22]

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][60][61] 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.[62] The principle of motion camouflage

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.[63]

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.[63]

Feeding

Adult dragonflies hunt on the wing using their exceptionally acute eyesight and strong, agile flight.[45]
They are almost exclusively carnivorous, eating a wide variety of insects ranging from small midges and
mosquitoes to butterflies, moths, damselflies, and smaller dragonflies.[59] 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.[64] A dragonfly may consume as much
as a fifth of its body weight in prey per day.[65] Dragonflies are also
some of the insect world's most efficient hunters, catching up to
95% of the prey they pursue.[66]

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.[59] A few species, especially those that live in temporary Common clubtail, Gomphus
waters, are likely to leave the water to feed. Nymphs of vulgatissimus, with prey
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.[9][67]

Eyesight

Dragonfly vision is thought to be like slow motion for humans. Dragonflies see faster than we do; they see
around 200 images per second.[68] A dragonfly can see in 360 degrees, and nearly 80 percent of the
insect's brain is dedicated to its sight.[69]

Predators
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,[70] and the hobby;[71] 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,
Southern red-billed hornbill with a
various species of ducks and herons eat dragonfly nymphs[70] and
captured dragonfly in its bill
they are also preyed on by newts, frogs, fish, and water spiders.[72]
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.[73]

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.[74] Gregarines infect the gut and may cause blockage and secondary infection.[75] 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.[76]

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.[77]

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.[77]

Habitat degradation has reduced dragonfly populations across the world, for example in Japan.[78] 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.[79][80]
Dragonflies are steadily declining in Africa, and represent a conservation priority.[81]

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.[82]

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.[83][84]

In culture

A blue-glazed faience dragonfly amulet was found by Flinders

Petrie at Lahun, from the Late Middle Kingdom of ancient
Egypt.[85]

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
Flying Dragon, by Alexander Calder,
ground and healed them if they were injured.[86] For the Navajo,
1975, Art Institute of Chicago
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.[87]: 20–26

As a seasonal symbol in Japan, the dragonflies are associated with season of autumn.[88] 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.[87]: 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.[89]

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.[90]
Dragonfly - a close-up view.
Images of dragonflies are common in Art Nouveau, especially in
jewellery designs.[91] They have also been used as a decorative
motif on fabrics and home furnishings.[92] Douglas, a British motorcycle manufacturer based in Bristol,
named its innovatively designed postwar 350-cc flat-twin model the Dragonfly.[93]

Among the classical names of Japan are Akitsukuni ( 秋津国 秋津島

), Akitsushima ( ), Toyo-akitsushima ( 豊
秋津島 ). Akitsu is an old word for dragonfly, so one interpretation of Akitsushima is "Dragonfly
Island".[94] 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.[95][96]

In Europe, dragonflies have often been seen as sinister. Some English

vernacular names, such as "horse-stinger",[97] "devil's darning needle", and
"ear cutter", link them with evil or injury.[98] Swedish folklore holds that the
devil uses dragonflies to weigh people's souls.[87]: 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".[98] 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.[99][100] Interestingly, the
Dragonfly pictured in the
Hungarian name for dragonfly is szitakötő ("sieve-knitter"). coat of arms of Kiili
Parish
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][101]

More recently, dragonfly watching has become popular in America as some birdwatchers seek new groups
to observe.[102]

In heraldry, like other winged insects, the dragonfly is typically depicted tergiant (with its back facing the
viewer), with its head to chief.[103]
 Woodcut on paper, after
Kitagawa Utamaro, 1788
Dragonfly symbol on a Hopi
bowl from Sikyátki, Arizona, Accurately drawn
c. 1400–1625 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.

Tiffany dragonfly pendant

lamp, designed c. 1903
Tiffany & Co. Japonism Japanese tsuba with a
vase with dragonfly dragonfly, 1931: Shibuichi
handles, c. 1879, Walters with gold and silver, Walters
Art Museum 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."[104] 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.[105] Hori Bakusui (1718–1783) similarly wrote
"Dyed he is with the / Colour of autumnal days, / O red dragonfly."[104]

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."[106]
The novelist H. E. Bates described the rapid, agile flight of dragonflies in his 1937 nonfiction book[107]
Down the River:[108]

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.[109]

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".[110][111]

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.[101]
c. Optrode is a portmanteau of "optical electrode".

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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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