Dragonfly - Wikipedia

2/1/23, 8:40 AM Dragonfly - Wikipedia
Contents
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1Taxonomy
2Evolution
3Distribution and diversity

4General description
4.1Coloration
5Biology
5.1Ecology
5.2Behaviour
5.3Reproduction
5.4Life cycle
5.5Sex ratios
5.6Flight
5.6.1Flight speed
5.6.2Motion camouflage
5.7Temperature control
5.8Feeding
5.9Eyesight
6Predators
7Parasites
8Dragonflies and humans
8.1Conservation
8.2In culture
8.3In poetry and literature
8.4In technology
9Explanatory notes
https://en.wikipedia.org/wiki/Dragonfly 1/23
10References
10.1Citations
10.2General and cited sources
11External links
Dragonfly
A dragonfly is a flying insect belonging to the infraorder
Anisoptera below the order Odonata. About 3,000 extant species of Dragonfly
true dragonfly are known. Most are tropical, with fewer species in Temporal range: Early Jurassic to
temperate regions. Loss of wetland habitat threatens dragonfly Recent
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.
Dragonflies can be mistaken for the closely related damselflies, which Yellow-winged darter
make up the other odonatan infraorder (Zygoptera) and are similar Sympetrum flaveolum
in body plan though usually lighter in build; however, the wings of
most dragonflies are held flat and away from the body, while Scientific classification
damselflies hold their wings folded at rest, along or above the Kingdom: Animalia
abdomen. Dragonflies are agile fliers, while damselflies have a
weaker, fluttery flight. Phylum: Arthropoda
Dragonflies are predatory insects, both in their aquatic nymphs stage Class: Insecta
(also known as naiads) and as adults. In some species, the nymphal Order: Odonata
stage lasts for up to five years, and the adult stage may be as long as
ten weeks, but most species have an adult lifespan in the order of five Suborder: Epiprocta
weeks or less, and some survive for only a few days.[2] They are fast, Infraorder: Anisoptera
agile fliers capable of highly accurate aerial ambush, sometimes Selys, 1854[1]
migrating across oceans, and often live near water. They have a
uniquely complex mode of reproduction involving indirect Families
insemination, delayed fertilization, and sperm competition. During
mating, the male grasps the female at the back of the head, and the Aeshnoidea
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 Aeshnidae (hawkers or
"heart" or "wheel" posture. darners)
Fossils of very large dragonfly-like insects, sometimes called Austropetaliidae
griffinflies, are found from 325 million years ago (Mya) in Upper Gomphidae (clubtails)
Carboniferous rocks; these had wingspans up to about 750 mm
(30 in), though they were only distant relatives, not true dragonflies. Petaluridae (petaltails)
Cordulegastroidea
Dragonflies are represented in human culture on artefacts such as
pottery, rock paintings, statues and Art Nouveau jewellery. They are Chlorogomphidae
used in traditional medicine in Japan and China, and caught for food
in Indonesia. They are symbols of courage, strength, and happiness Cordulegastridae

in Japan, but seen as sinister in European folklore. Their bright
(spiketails)
colours and agile flight are admired in the poetry of Lord Tennyson
and the prose of H. E. Bates. Neopetaliidae
Libelluloidea
Taxonomy
Libbelulidae (skimmers,
The infraorder Anisoptera comes from Greek ἄνισος anisos etc)
"unequal"[3] and πτερόν pteron "wing"[4] because dragonflies' Corduliidae (skimmers,
hindwings are broader than their forewings.[5]
etc)
Macromiidae (cruisers)
Evolution
Synthemistidae (tigertails)
Dragonflies and their relatives are similar in structure to an ancient $Not a clade
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);.[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 The giant Upper Carboniferous
both of those groups are sometimes referred to as "giant dragonflies", in dragonfly relative, Meganeura
fact true dragonflies/odonata are more modern insects that had not monyi, attained a wingspan around
evolved yet. 680 mm (27 in).[6] Museum of
Toulouse
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 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

Mesurupetala, Late Jurassic
2013, but all the families are monophyletic except the Corduliidae; the
(Tithonian), Solnhofen limestone,
Gomphidae are a sister taxon to all other Anisoptera, the
Germany
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):
Dragonflies in Oze National Park
⁠
⁠Gomphidae (clubtails)
⁠
⁠Austropetaliidae
⁠ ⁠Aeshnoidea (hawkers)
⁠⁠
⁠
⁠Petaluridae (petaltails)
⁠
⁠Macromiidae (cruisers)
⁠
⁠Anisoptera ⁠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).[12]
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;[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 An aggregation of globe
example in the Mojave Desert, where they are active in shade temperatures skimmers, Pantala
between 18 and 45 °C (64 and 113 °F); these insects were able to survive body flavescens, during migration
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
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
Damselflies, like this Ischnura
ommatidia of just one size. These facets provide complete vision in the
senegalensis, are more slender in
frontal hemisphere of the dragonfly.[24] The compound eyes meet at the
build than dragonflies, and most top of the head (except in the Petaluridae and Gomphidae, as also in the
hold their wings closed over their genus Epiophlebia). Also, they have three simple eyes or ocelli. The
bodies. 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]
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.[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 Anatomy of a dragonfly
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]
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
Migrant hawker, Aeshna labium, armed with hooks and spines, which is used for catching prey. This
mixta, has the long labium is folded under the body at rest and struck out at great speed by hydraulic
slender abdomen of pressure created by the abdominal muscles.[12] Both damselsfly and dragonfly
aeshnid dragonflies. 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
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
Iridescent structural colouration in a
blue, waxy powderiness called pruinosity; it wears off when scraped
dragonfly's eyes during mating, leaving darker areas.[36]
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
Male green darner, Anax junius has
patches at the wing bases. Some aeshnids such as the brown hawker
noniridescent structural blue; the
(Aeshna grandis) have translucent, pale yellow wings.[38]
female (below) lacks the colour.
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]
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 Habitat preference: A four-spotted
from the absence of predators there.[41] Vegetation and its chaser, Libellula quadrimaculata on
characteristics including submerged, floating, emergent, or waterside an emergent plant, the water violet
are also important. Adults may require emergent or waterside plants to Hottonia palustris, with submerged
use as perches; others may need specific submerged or floating plants vegetation in the background
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 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]
Mating pair of marsh
the pair may also be described as being "in cop".[52] skimmers, Orthetrum
luzonicum, forming a
Egg-laying (ovipositing) involves not only the female darting over floating or
"heart"
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
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 Illustration of a naiad with mask
tissues are usually shaped like grains of extended
rice, while other eggs are the size of a
pinhead, ellipsoidal, or nearly
Nymph of emperor dragonfly, Anax spherical. A clutch may have as many as 1500 eggs, and they take about
imperator 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]
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 Ecdysis: Emperor dragonfly,
species" emerge within a few days of each other. The springtime darner Anax imperator, newly
(Basiaeschna janata), for example, is suddenly very common in the spring, but emerged and still soft,
disappears a few weeks later and is not seen again until the following year. By holding on to its dry exuvia,
contrast, a "summer species" emerges over a period of weeks or months, later and expanding its wings
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]
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.[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 Brown hawker, Aeshna grandis in
and generates a large amount of lift. flight: The hindwings are about 90°
out of phase with the forewings at
Phased-stroking, with the hindwings beating 90° ahead of the
this instant, suggesting fast flight.
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]
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]
Southern hawker, Aeshna cyanea: Dragonflies generate lift in at least four

its wings at this instant are ways at different times, including classical Red-veined darters
synchronised for agile flight. lift like an aircraft wing; supercritical lift (Sympetrum fonscolombii)
with the wing above the critical angle, flying "in cop" (male ahead)
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
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] 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.[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
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 Common clubtail, Gomphus
also some of the insect world's most efficient hunters, catching up to vulgatissimus, with prey
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 we 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
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
Southern red-billed hornbill with a
dragonfly nymphs[76] and they are also preyed on by newts, frogs, fish,
captured dragonfly in its bill
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 Akitsukuni (秋津国), Akitsushima (秋津島), 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]
Woodcut on paper, after

Dragonfly symbol on a Hopi Kitagawa Utamaro, 1788
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,
Art Museum 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 (https://archive.org/details/dragonflieswildg00cynt). Stackpole

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2/1/23, 8:40 AM Dragonfly - Wikipedia

3Distribution and diversity

8Dragonflies and humans

8.3In poetry and literature

10.2General and cited sources

in Indonesia. They are symbols of courage, strength, and happiness Cordulegastridae

The relationships of anisopteran families are not fully resolved as of

Dragonflies in Oze National Park

Distribution and diversity

Family Oriental Neotropical Australasian Afrotropical Palaearctic Nearctic Pacific World

Aeshnidae 149 129 78 44 58 40 13 456

Libellulidae 192 354 184 251 120 105 31 1037

Dragonflies live on every continent except Antarctica. In contrast to the

The thorax consists of three segments as

Many adult dragonflies have brilliant iridescent or metallic colours

Some dragonflies, such as the green

Dragonflies as a group occupy a considerable variety of habitats, but

Mating in dragonflies is a complex, precisely choreographed process. First, the

Dragonflies are hemimetabolous

Dragonflies in temperate areas can be categorized into two groups, an early

Dragonflies are powerful and agile fliers, capable of migrating across

Counter-stroking, with forewings beating 180° out of phase with the

The wings are powered directly, unlike

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

In high-speed territorial battles between male Australian emperors

Adult dragonflies hunt on the wing using their exceptionally acute

Dragonflies and humans

Woodcut on paper, after

Tiffany dragonfly pendant

In poetry and literature

5. Gordh, G.; Headrick, D. (2011). A dictionary of entomology (https://books.google.com/books?id=9IcmC

43. Dijkstra 2006, p. 246.

80. Mead, Kurt. "Dragonfly Biology 101" (https://web.archive.org/web/20150227190348/http://www.mndrag

General and cited sources

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