Dragonfly

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"Anisoptera" redirects here. For other uses, see Anisoptera (disambiguation).
This article is about the insect. For other uses, see Dragonfly (disambiguation).

Dragonfly
Temporal range: Jurassic–recent

PreЄ

Pg

Yellow-winged darter

Sympetrum flaveolum

Scientific classification

Kingdom: Animalia
 Clade: Euarthropoda

Class: Insecta

Order: Odonata

Suborder: Epiprocta

Infraorder: Anisoptera
Selys, 1854

Families

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

A dragonfly is an insect belonging to the

order Odonata, infraorder Anisoptera (from Greek ἄνισος anisos, "uneven" and πτερόν pteron,
"wing", because the hindwing is broader than the forewing). Adult dragonflies are characterized
by large, multifaceted eyes, two pairs of strong, transparent wings, sometimes with coloured
patches, and an elongated body. Dragonflies can be mistaken for the related
group, damselflies (Zygoptera), which are similar in structure, though usually lighter in build;
however, the wings of most dragonflies are held flat and away from the body, while damselflies
hold the wings folded at rest, along or above the abdomen. Dragonflies are agile fliers, while
damselflies have a weaker, fluttery flight. Many dragonflies have brilliant iridescent or metallic
colours produced by structural coloration, making them conspicuous in flight. An adult
dragonfly's compound eyes have nearly 24,000 ommatidia each.
Fossils of very large dragonfly ancestors in the Protodonata are found from 325 million years ago
(Mya) in Upper Carboniferous rocks; these had wingspans up to about 750 mm (30 in).
About 3000 species of Anisoptera are in the world today. Most are tropical, with fewer species
in temperate regions.
Dragonflies are predators, both in their aquatic larval stage, when they are known as nymphs or
naiads, and as adults. Several years of their lives are spent as nymphs living in fresh water; the
adults may be on the wing for just a few days or weeks. They are fast, agile fliers, sometimes
migrating across oceans, and are often found 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 or on the prothorax, 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.
Loss of wetland habitat threatens dragonfly populations around the world. Dragonflies are
represented in human culture on artifacts such as pottery, rock paintings, 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 Alfred, Lord Tennyson and the prose of H. E. Bates.

Contents
[hide]

 1Phylogeny
 2Distribution and diversity
 3General description
o 3.1Coloration
 4Biology
o 4.1Ecology
o 4.2Behaviour
o 4.3Reproduction
o 4.4Life cycle
o 4.5Sex ratios
o 4.6Flight
 4.6.1Flight speed
 4.6.2Motion camouflage
o 4.7Temperature control
o 4.8Feeding
 5Predators and parasites
 6Dragonflies and humans
o 6.1Conservation
o 6.2In culture
o 6.3In poetry and literature
o 6.4In technology
 7Notes
 8References
 9Further reading
 10External links

Phylogeny[edit]
The giant Upper Carboniferousdragonfly ancestor, Meganeura monyi, attained a wingspan of about
680 mm (27 in).[1] Museum of Toulouse

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

Dragonflies and their relatives are an ancient group. The oldest fossils are of
the Protodonata 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);[2] their fossil record ends with the Permian–Triassic extinction
event (about 247 Mya). The Protanisoptera, another ancestral group which 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. The forerunners of
modern Odonata are included in a clade called the Panodonata, which include the basal
Zygoptera (damselflies) and the Anisoptera (true dragonflies)[3] Today there are some 3000
species extant around the world.[4][5]
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.[6] On the cladogram, dashed
lines indicate unresolved relationships; English names are given (in parentheses):
Anisoptera
Gomphidae (clubtails)

Austropetaliidae

Aeshnoidea (hawkers)

Petaluridae (petaltails)

Macromiidae (cruisers)

Libelluloidea
Neopetaliidae

Cordulegastridae (goldenrings)

Libellulidae (skimmers)

"Corduliidae" [not a clade] (emeralds)

Synthemistidae (tigertails)

Chlorogomphidae
Distribution and diversity[edit]
About 3012 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
summarised below (the world numbers are not ordinary totals, as overlaps in species occur).[7]

[hide]Famil Orien Neotrop Australa Afrotrop Palaear Nearc Paci Wor

y tal ical sian ical ctic tic fic ld

Aeshnidae 149 129 78 44 58 40 13 456

Austropetalii
7 4 11
dae

Petaluridae 1 6 1 2 10

Gomphidae 364 277 42 152 127 101 980

Chlorogomp
46 5 47
hidae

Cordulegastri
23 1 18 46
dae

Neopetaliida
1 1
e

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

Synthemistid
37 9 46
ae

Incertae
37 24 21 15 2 99
sedis
An aggregation of globe skimmers, Pantala flavescens, during migration

Dragonflies are found on every continent except Antarctica. In contrast to the damselflies
(Zygoptera), which tend to have restricted distributions, some genera and species are found
across continents. For example, the blue-eyed darner Rhionaeschna multicolor is found all
across North America, and in Central America;[8] emperors Anax are found throughout the
Americas from as far north as Newfoundland to as far south as Bahia Blanca in
Argentina,[9] across Europe to central Asia, North Africa, and the Middle East.[10] 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.[11]
Dragonflies including libellulids and aeshnids are found in desert pools, for example in
the Mojave Desert, where they are active in shade temperatures between 18 and 45 °C (64.4 to
113 °F); these insects were able to survive body temperatures above the thermal death point of
insects of the same species found in cooler places.[12]
Dragonflies can be found from sea level up to the mountains, decreasing in species diversity with
altitude.[13] Their altitudinal limit is about 3700 m, represented by a species of Aeshna in
the Pamirs.[14]
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.[15] 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.[16] The treeline emerald is also found
in northern Alaska, within the Arctic Circle, making it the most northerly of all dragonflies.[17]

General description[edit]

Damselflies, like this Ischnura senegalensis, are more slender 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.[7][18]
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.[19] 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.[20][21] 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.[7]

Anatomy of a dragonfly

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 provides a robust attachment for the powerful wing muscles inside it.[22] 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.[23] 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.[21]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.[22]
Migrant hawker, Aeshna mixta, has the long slender abdomen of aeshnid dragonflies.

The abdomen is long and slender and consists of 10 segments and a terminal appendage-
bearing segment. The second and third segments are enlarged, and in males, a cleft on the
underside of the second segment contains a pair of claspers and the penis. The spermaries open
on the 9th segment. In females, the genital opening is on the underside of the eighth segment
and is covered by a simple flap or an ovipositor, depending on species and the method of egg-
laying.[22]
Dragonfly nymphs vary in form with species and are loosely classed into claspers, sprawlers,
hiders, and burrowers.[7] The first instar is known as a prolarva, a relatively inactive stage from
which it quickly moults into the more active nymphal form.[24] 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.[7]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.[25]
Coloration[edit]

Iridescent structural coloration in a dragonfly's eyes

Many adult dragonflies have brilliant iridescent or metallic colours produced by structural
coloration, 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-coloured and obtain
their typical colours after a few days,[23] some have their bodies covered with a pale blue, waxy
powderiness called pruinosity; it wears off when scraped during mating, leaving darker areas.[26]
Noniridescent structural blue occurs in the green darner, Anax junius; the female (below) lacks blue.

Some dragonflies, such as the green darner, Anax junius, have a noniridescent blue which is
produced structurally by scatter from arrays of tiny spheres in the endoplasmic reticulum of
epidermal cells underneath the cuticle.[27]
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.[28]
Dragonfly nymphs are usually a well-camouflaged blend of dull brown, green, and grey.[25]

Biology[edit]
Ecology[edit]
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 tadpolesand
small fish.[29] 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.[29]
Adult males vigorously defend territories near water; these areas provide suitable habitat for the
larvae 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.[29]

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.[30] 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.[30] Some species are found 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 larvae that can resist drought and are stimulated to grow rapidly in warm, shallow pools, also
often benefiting from the absence of predators there.[30] 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.[30] 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 acid;[30] a
few species such as Sympetrum danae (black darter) and Libellula quadrimaculata (four-spotted
chaser) prefer acidic waters such as peat bogs,[31] while others such as Libellula fulva (scarce
chaser) need slow-moving, eutrophic waters with reeds or similar waterside plants.[32][33]
Behaviour[edit]
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,
and the blue dasher (Pachydiplax longipennis) jostles other dragonflies to maintain the right to
alight there.[34]
Defending a breeding territory is fairly common among male dragonflies, especially among
species that congregate around ponds in large numbers. The territory contains desirable features
such as a sunlit stretch of shallow water, a special plant species, or a
particular substrate necessary 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. Some dragonflies signal ownership with striking colours on the face, abdomen,
legs, or wings. The common whitetail (Plathemis lydia) 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.[34] 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.[35] In
some species females have evolved behavioural responses such as feigning death to escape the
attention of males.[36]
Reproduction[edit]

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.[37] 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";[29][38] the pair may also be described
as being "in cop".[39]
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,[40] something made possible by delayed fertilisation[29][38] and driven by sexual
selection.[37] 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.[39]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.[40]
Egg-laying takes two different forms depending on the species. The female in some families 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.[39] 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.[25]
Life cycle[edit]

Nymph of emperor dragonfly, Anax imperator

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.[41] 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.[7] 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.[41] They breathe through gills in their rectum, and
can rapidly propel themselves by suddenly expelling water through the anus.[42] Some naiads,
such as the later stages of Antipodophlebia asthenes, hunt on land.[43]
Ecdysis: Emperor dragonfly, Anax imperator, newly emerged and still soft, holding on to its dry exuvia, and
expanding its wings

Parts of a dragonfly nymph including the labial "mask"

The larval 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 larval 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.[44]
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 has disappeared 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 short lifespans which is
an average of 7 months.[45][46]

Sex ratios[edit]
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.[47]

Brown hawker, Aeshna grandis in flight: The hindwings are about 90° out of phase with the forewings at
this instant, suggesting fast flight.

Flight[edit]

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

Dragonflies are powerful and agile fliers, capable of migrating across oceans, moving in any
direction, and changing direction suddenly. In flight, the adult dragonfly can propel itself in six
directions: upward, downward, forward, back, to left and to right.[48] They have four different styles
of flight:[49] 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 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; 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.[49]

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.[49]
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; creating vortices; and vortex shedding. 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.[49]
Flight speed[edit]
Old and unreliable claims are made that dragonflies such as the southern giant darner can fly up
to 60 miles per hour (97 km/h).[50] However, the greatest reliable flight speed records are for other
types of insects.[51] In general, large dragonflies like the hawkers have a maximum speed of 10–
15 metres per second (22–34 mph) with average cruising speed of about 4.5 metres per second
(10 mph).[52]Dragonflies can fly at 100 body-lengths per second, and three lengths per second
backwards.[20]
Motion camouflage[edit]

The principle of motion camouflage

Further information: 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][53][54] 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.[55]
Temperature control[edit]
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.[56]
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.[56]

Blue dasher eating honey bee

Common clubtail, Gomphus vulgatissimus, with prey

Feeding[edit]
Adult dragonflies hunt on the wing using their exceptionally acute eyesight and strong, agile
flight.[38] They are almost exclusively carnivorous, eating a wide variety of insects ranging from
small midges and mosquitoes to butterflies, moths, damselflies, and smaller dragonflies.[52] 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.[57] A dragonfly may
consume as much as a fifth of its body weight in prey per day.[58]
The larvae 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.[52] A few species, especially those that live in temporary waters, are likely to leave
water. Nymphs of Cordulegaster bidentata sometimes hunt small arthropods on the ground at
night.[7]
Predators and parasites[edit]

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,[59] and
the hobby;[60] 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 larvae[59] and
they are also preyed on by newts, frogs, fish, and water spiders.[61] 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.[62]
Dragonflies are affected by three major groups of parasites: water mites, gregarine protozoa,
and trematode flatworms (flukes). Water mites, Hydracarina, can kill smaller dragonfly larvae,
and may also be seen on adults.[63] Gregarines infect the gut and may cause blockage and
secondary infection.[64] 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.[65]

Dragonflies and humans[edit]

Woodcut on paper, after Kitagawa Utamaro, 1788

Conservation[edit]
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.[66]
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.[66]
Habitat degradation has reduced dragonfly populations across the world, for example in
Japan.[67] 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.[68][69] Dragonflies are steadily declining in Africa, and represent a
conservation priority.[70]
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.[71]
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.[72][73]
In culture[edit]
A blue-glazed faience dragonfly amulet was found by Flinders Petrie at Lahun, from the
Late Middle Kingdom of ancient Egypt.[74]

Dragonfly symbol on a Hopi bowl from Sikyátki, Arizona

For some Native American tribes, dragonflies represent swiftness and activity; for the Navajo,
they symbolize pure water. They are a common motif in Zuni pottery; stylized as a double-barred
cross, they appear in Hopi rock art and on Pueblo necklaces.[75]:20–26 They have been used in
traditional medicine in Japan and China. In Indonesia, adults are caught on poles made sticky
with birdlime, then fried in oil as a delicacy.[76]

Tiffany dragonfly pendant lamp

In the United States, dragonflies and damselflies are sought out as a hobby similar to birding and
butterflying, known as oding, from the Latin name of the dragonfly order, Odonata. Oding is
especially popular in Texas, where 225 different species of odonates have been observed. With
care, and with dry fingers, dragonflies can be handled and released by oders, as can be done
with butterflies, though it is not encouraged.[77]

Tiffany & Co. Japonismvase with dragonfly handles, circa 1879, Walters Art Museum

Images of dragonflies are common in Art Nouveau, especially in jewellery designs.[78] They have
also been used as a decorative motif on fabrics and home furnishings.[79] Douglas, a British
motorcycle manufacturer based in Bristol, named its innovatively designed postwar 350-cc flat-
twin model the Dragonfly.[80]
Among the classical names of Japan are Akitsukuni (秋津国), Akitsushima (秋津島), Toyo-
akitsushima (豊秋津島). Akitu or akidu are archaic or dialectal Japanese words for dragonfly, so
one interpretation of Akitsushima is "Dragonfly Island".[81] This is attributed to a legend in which
Japan's mythical founder, Emperor Jinmu, was bitten by a mosquito, which was then eaten by a
dragonfly.[82][83]
As a seasonal symbol in Japan, the dragonfly is associated with autumn.[84] More generally,
dragonflies are symbols of courage, strength, and happiness, and they often appear in art and
literature, especially haiku. 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.[75]:38

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.

In Europe, dragonflies have often been seen as sinister. Some English vernacular names, such
as "horse-stinger",[85] "devil's darning needle", and "ear cutter", link them with evil or
injury.[86] Swedish folklore holds that the devil uses dragonflies to weigh people's souls.[75]: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".[86] The Southern United States term "snake
doctor" refers to a folk belief that dragonflies follow snakes around and stitch them back together
if they are injured.[87]
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 larva (at lower left) with the mask extended
appears to be plagiarised.[b][88]
In poetry and literature[edit]
Japanese tsuba with a dragonfly, 1931: Shibuichiwith gold and silver, Walters Art Museum

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."[89] 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.[90] Hori Bakusui (1718–1783) similarly wrote "Dyed he is with the / Colour of autumnal
days, / O red dragonfly."[89]
The poet Alfred, 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."[91]
The novelist H. E. Bates described the rapid, agile flight of dragonflies in his 1937 nonfiction
book[92] Down the River:[93]
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.[94]

In technology[edit]
A dragonfly has been genetically modified with light-sensitive "steering neurons" in its nerve cord
to create a cyborg-like "DragonflEye". The neuronscontain 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".[95][96]

Notes[edit]
1. Jump up^ This is not to say that other species may not use the
same technique, only that this species has been studied.
2. Jump up^ 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 larva 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.[88]
3. Jump up^ Optrode is a portmanteau of "optical electrode".

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94. Jump up^ Bates, H. E. (12 February 1937). "Country Life: Pike
and Dragonflies". The Spectator (5668): 269 (online p. 17).
95. Jump up^ "Equipping Insects for Special Service". Draper. 19
January 2017.
96. Jump up^ Ackerman, Evan (1 June 2017). "Draper's Genetically
Modified Cyborg DragonflEye Takes Flight". IEEE Spectrum.

Further reading[edit]
 Berger, Cynthia (2004). Dragonflies. Stackpole
Books. ISBN 978-0-8117-2971-0.
 Corbet, Phillip S. (1999). Dragonflies: Behavior and Ecology of
Odonata. Ithaca, NY: Cornell University Press. pp. 559–
561. ISBN 0-8014-2592-1.
 Dijkstra, Klaas-Douwe B. (2006). Field Guide to the Dragonflies
of Britain and Europe. British Wildlife Publishing. ISBN 0-
9531399-4-8.
 Meister, Cari (2001). Dragonflies. ABDO. p. 16. ISBN 978-1-
57765-461-2.
 Powell, Dan (1999). A Guide to the Dragonflies of Great Britain.
Arlequin Press. ISBN 1-900-15901-5.
 Trueman, John W. H.; Rowe, Richard J. (2009). "Odonata". Tree
of Life. Retrieved 25 February 2015.

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

[show]

Extant Odonata families

[show]

Insects in culture

66

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paea: 11840

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Categories:
 Insects acting as insect pest control agents
 Dragonflies
 Odonata
 Extant Pennsylvanian first appearances
 Insects in culture
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