Dragonfly

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This article is about the insect. For other uses, see Dragonfly (disambiguation).
"Anisoptera" redirects here. For other uses, see Anisoptera (disambiguation).
Dragonfly
Sympetrum flaveolum - side (aka).jpg
Yellow-winged darter
Scientific classification e
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Odonata
(unranked): Epiprocta
Suborder: Anisoptera
Selys, 1854
Families

A dragonfly is an insect belonging to the suborder Anisoptera (from Greek ανισος anisos, "uneven" + πτερος pteros, "wings", because the hindwing is broader than the forewing). It is 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 away from, and perpendicular to, the body when at rest, while damselflies hold the wings along the body. Dragonflies are agile fliers. Many dragonflies have brilliant iridescent or metallic colours produced by structural coloration, making them conspicuous in flight.

Fossils of very large dragonfly ancestors in the Protodonata are found from 325 million years ago in Upper Carboniferous rocks; these had wingspans of up to 750 mm. There are about 3000 species of Anisoptera in the world today. Most are tropical, with fewer species in temperate regions.

Dragonflies are predators, both as larvae and as adults. The larvae (nymphs) are aquatic, living in freshwater; the adults are fast agile fliers, sometimes migrating across oceans, and are often but not always found near water. They have a uniquely complex method of reproduction involving indirect insemination, delayed fertilisation 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 artefacts such as pottery, rock art and Art Nouveau jewels. They are used in traditional medicine in Japan and China, and caught for food and fried 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.

Phylogeny[edit]

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

Dragonflies are an ancient group. The oldest fossils are of the Protodonata from the 325 mya Upper Carboniferous of Europe, which had wingspans up to 750 mm; they became extinct in the Triassic. The Protoanisoptera, forerunners of the Anisoptera (true dragonflies), have also been found in Carboniferous rocks from Abilene, Kansas, USA and in Europe. Protoanisopteran fossils from Commentry, France and Bolsover, Derbyshire, England have wingspans up to 700 and 500 mm respectively. Today there are some 3000 species extant around the world.[1][2][3]

The relationships of Anisopteran families remained not fully resolved in 2013, but all the families are monophyletic except the Corduliidae; the Gomphidae are a sister taxon to all other Anisoptera, the Austropetaliidae are a sister to the Aeshnoidea, and the Chlorogomphidae are a sister to a clade that includes the Synthemistidae and Libellulidae.[4] 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







General description[edit]

Dragonfly anatomy (Orthetrum cancellatum shown). 1: Clypeus, 2: Antenna, 3,4 Frons, 5: Labrum, 6: Claw, 7: Tarsus 8: Tibia, 9: Spines, 10: Femur, 11: Coxa, 12: Sutures, 13: Thorax, 14: Abdomen, 15: Appendages (claspers), 16: Anal loop, 17: Triangles, 18: Rear wing, 19: Front wing, 20: Pterostigma, 21: Postnodal cross-veins, 22 : Node, 23: Antenodal cross-veins, 24: Arculus, 27: Prothorax, 28: Labium, 29: Mandibles, 30: Compound eye

Dragonflies (suborder Anisoptera) are heavy-bodied, strong-flying insects that hold their wings horizontally when at rest. By contrast, damselflies (suborder Zygoptera) are slender-bodied weak-fliers that fold their wings over the abdomen when stationary.[5]

An adult dragonfly has the same typical division into head, thorax and abdomen as do other 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, each consisting of nearly 28,000 ommatidia. There are also three simple eyes. The mouthparts are adapted for biting with a toothed jaw and the flap-like labrum, at the front of the mouth, can be shot rapidly forward to catch prey. The thorax bears two pairs of wings and three pairs of legs. The wings are long, veined and membranous, each pair being of similar size and shape, narrower at the tip and wider at the base. The veins contain hemolymph and both stiffen and strengthen the wings. 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 dragonfly, the wings of females are shorter and broader than those of males. The legs are seldom used for walking, but are used to catch and hold prey and for perching on plants. The abdomen is long and slender and bears the reproductive organs.[1][6]

Coloration[edit]

Iridescent structural coloration in a dragonfly's eyes

Many 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. Adults often change colour on reaching sexual maturity,[7] typically with a pale blue waxy powderiness called pruinosity; it wears off when scraped during mating, leaving darker areas.[8]

Non-iridescent structural blue in the green darner, Anax junius; the female (below) lacks blue.

Some dragonflies such as the green darner, Anax junius, have a non-iridescent blue which is produced structurally by scatter from arrays of tiny spheres in the endoplasmic reticulum of epidermal cells underneath the cuticle.[9]

Distribution[edit]

Dragonflies, and indeed most families within the Anisoptera, are found on every continent of the world except Antarctica. In contrast to the damselflies (Zygoptera) which tend to have restricted distributions, some genera and individual species are found across whole continents. For example, the blue-eyed darner Rhionaeschna multicolor is found all across North America, and in Central America;[10] emperors Anax are found 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] Most Anisoptera species however are tropical, with far fewer species in temperate regions.[13]

Biology[edit]

Ecology[edit]

Dragonflies as a group have three distinctive patterns of behaviour:

  1. Firstly, they are amphibious: while the adults fly, breathe air and live on land, the larvae live in fresh water.
  2. Secondly, they are predatory: in every species, both the adults and the larvae hunt other animals. Adults generally catch flying insects; the larvae hunt a range of freshwater invertebrates including insects, but the prey of the larger dragonfly larvae ranges up to amphibian tadpoles and small fish.
  3. Thirdly, their reproduction, unlike that of any other group, involves indirect insemination and delayed fertilisation.[14]

Together, these three features explain many aspects of dragonfly behaviour. Adult males vigorously defend territories beside freshwater; these areas provide suitable habitat for the larvae to develop, and for females to lay their eggs. Swarms of feeding adults are readily explained through predation on swarming prey, such as emerging flying ants. Highly characteristic behaviours are associated with reproduction.[14]

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) will jostle other dragonflies to maintain the right to alight here.[15]

Defending a breeding territory is fairly common among male dragonflies, especially among species that congregate around ponds in large numbers. The territory will contain desirable features such as a sunlit stretch of shallow water, a special plant species or a particular substrate that is necessary for egg-laying. It 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 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. Any female will need to mate with the territory holder before laying her eggs.[15]

Reproduction[edit]

Pair of yellow striped hunters forming the mating posture known as the "heart" or "wheel"

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.[16] The pair fly in tandem with the male in front, typically to 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";[14][17] the pair may also be described as being "in cop".[18]

Egg-laying (ovipositing) involves not only the female, darting over floating or waterside vegetation to deposit eggs on a suitable substrate, but the male hovering above her or continuing to clasp her and flying in tandem, so as to prevent rival males from attempting to remove his sperm and insert their own, something made possible by delayed fertilisation[14][17] and driven by Sexual selection.[16] To do that, the male uses his penis to compress or to scrape out the sperm inserted by the previous male. This activity takes up much of the time that the copulating pair remain in the heart posture.[18] Egg-laying takes two different forms in different species of dragonfly. The females in some families have 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 females lay eggs by tapping the surface of the water repeatedly, by shaking the eggs out of her abdomen as she flies along, or in some cases by placing their eggs on vegetation.[18]

Life cycle[edit]

Dragonfly nymph eating tadpole

Female dragonflies lay eggs in or near water, often on floating or emergent plants. When laying eggs, some species will submerge themselves completely in order to lay their eggs on a good surface. The eggs then hatch into naiads (nymphs). Most of a dragonfly's life is spent in the naiad form, beneath the water's surface, using extendable jaws to catch other invertebrates (often mosquito larvae) or even vertebrates such as tadpoles and fish.[19] They breathe through gills in their rectum, and can rapidly propel themselves by suddenly expelling water through the anus.[20] Some naiads hunt on land.[21]

Adult emerging from larval skin

The larval stage of large 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. 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, 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.[22]

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" will 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" will emerge 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.[23]

Sex ratios[edit]

The sex ratio between males and females dragonfly vary 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 the 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.[24]

Brown Hawker, Aeshna grandis in flight. The hind wings are about 90 degrees 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,[25] 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.[26] They have four different styles of flight:[27]

  • Counter-stroking, with forewings beating 180 degrees out of phase with the hindwings, is used for hovering and slow flight. This style is efficient and generates a large amount of lift.[27]
  • Phased-stroking, with the hindwings beating 90 degrees ahead of the forewings, is used for fast flight. This style creates more thrust but less lift than counter-stroking.[27]
  • Synchronised-stroking, with forewings and hindwings beating together, is used when changing direction rapidly, as it maximises thrust.[27]
  • 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 simply glides while the male pulls the pair along by beating his wings.[27]
Southern Hawker, Aeshna cyanea: its wings at this instant are synchronised for agile flight.

The wings are powered directly, with the flight muscles attached to the wing bases. Dragonflies have a high power/weight ratio, and have been documented accelerating at 4g linearly and 9g in sharp turns while pursuing prey.[27]

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.[27]

Flight speed[edit]

There are old and unreliable claims that dragonflies such as the southern giant darner can fly at up to 60 miles per hour (97 km/h).[28] However, the greatest reliable flight speed records are for other types of insects.[29] 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).[30] Dragonflies can fly at 100 body-lengths per second, and 3 lengths per second backwards.[1]

Motion camouflage[edit]

The principle of motion camouflage
Further information: motion camouflage

In territorial battles between male Australian emperors Hemianax papuensis (Aeshnidae), the fighting dragonflies adjust their flight paths to appear stationary to their rivals in "breakneck territorial dogfight[s]", minimizing the chance of being detected as they approach.[a][31][32] To achieve the effect, the attacking dragonfly flies towards its rival, choosing its path to remain on a line between the rival and the start of its attack path. The attacker thus looms larger as it closes on its rival, but does not otherwise appear to move. Researchers found that 6 of 15 encounters involved motion camouflage.[33]

Feeding[edit]

Common clubtail, Gomphus vulgatissimus, with prey

Dragonfly adults hunt on the wing using their exceptionally acute eyesight and strong agile flight.[17] They are almost exclusively carnivorous, eating a wide variety of insects from small midges to butterflies and damselflies. A dragonfly may consume as much as a fifth of its body weight in prey per day.[34]

Predators and parasites[edit]

Although dragonflies are swift and agile fliers, there are predators fast enough to catch them. These include falcons such as the American kestrel, the merlin,[35] and the hobby;[36] nighthawks, swifts, flycatchers and swallows also take some adults; some species of wasp, too, prey on dragonflies, using them to provision their nests, laying an egg on each captured insect. In the water, various species of duck and heron eat dragonfly larvae.[35]

Dragonflies are affected by three major types of parasite: water mites, gregarine protozoa, and trematode flatworms (flukes). Water mites, Hydracarina, can kill smaller dragonfly larvae, and may also be seen on adults.[37] Gregarines infect the gut and may cause blockage and secondary infection.[38] 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, it becomes infected by the adult or fluke stage of the trematode.[39]

Dragonflies and humans[edit]

Conservation[edit]

Habitat degradation has reduced dragonfly populations across the world, for example in Japan.[40] Over 60% of Japan's wetlands were lost in the twentieth 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.[41][42][43][44] Dragonflies are steadily declining in Africa, and represent a conservation priority.[45]

The dragonfly's long lifespan and low population density makes them 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.[46][47]

In culture[edit]

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.[48]: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.[49]

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.[50]

Tiffany & Co. Japonism vase with dragonfly handles, c. 1879. Walters Art Museum.

Images of dragonflies are common in Art Nouveau, especially in jewelry designs.[51] They have also been used as a decorative motif on fabrics and home furnishings.[52] Douglas, a British motorcycle manufacturer based in Bristol, named its innovatively designed postwar 350cc flat twin model the Dragonfly.[53]

As a seasonal symbol in Japan, the dragonfly is associated with autumn.[54] 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.[48]:38 Beyond this, one of Japan's historical names – Akitsushima (Kanji: Hiragana: あきつしま) – is an archaic form meaning "Dragonfly Islands".[55] 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.[56][57]

In Europe, dragonflies have often been seen as sinister. Some English vernacular names, such as "horse-stinger",[58] "devil's darning needle" and "ear cutter", link them with evil or injury.[49] Swedish folklore holds that the devil uses dragonflies to weigh people's souls.[48]:25–27 The Norwegian name for dragonflies is Øyenstikker ("eye-poker"), and in Portugal they are sometimes called tira-olhos ("eye-snatcher"). They are often associated with snakes, as in the Welsh name gwas-y-neidr, "adder's servant".[49] 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.[59]

In poetry and literature[edit]

Japanese Tsuba with a Dragonfly, 1931. Shibuichi with 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."[60] The poets Matsuo Bashō (1644-1694) and Hori Bakusui (1718-1783) wrote the following haiku relating the autumn season to the dragonfly:

Crimson pepper pod
add two pairs of wings, and look
darting dragonfly.

—Basho[61]

Dyed he is with the
Colour of autumnal days,
O red dragonfly.

—Bakusui[60]

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":[62]

Newly-emerged dragonfly with its larval skin (exuvia)

Today I saw the dragonfly
Come from the wells where he did lie.

An inner impulse rent the veil
Of his old husk: from head to tail
Came out clear plates of sapphire mail.
He dried his wings: like gauze they grew;
Through crofts and pastures wet with dew
A living flash of light he flew.

The novelist H. E. Bates described the rapid, agile flight of dragonflies in his 1937 non-fiction book[63] Down the River:[64]

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. It was like a crazy flight of almost invisible humming-birds. It never rested. Poised in that miraculous act of hovering, wings invisible, bodies like tiny fingers of blue steel, these small fragile creatures had exactly the same suspended power, the same dynamic and thrilling immobility, as the pike lying in wait in the water.[65]

Gallery[edit]

Notes[edit]

  1. ^ This is not to say that other species may not use the same technique, only that this species has been studied.

References[edit]

  1. ^ a b c "Introduction to the Odonata". UCMP Berkeley. Retrieved 24 February 2015. 
  2. ^ Zhang, Z.-Q. (2011). "Phylum Arthropoda von Siebold, 1848 In: Zhang, Z.-Q. (Ed.) Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness". Zootaxa 3148: 99–103. 
  3. ^ Dunkle, Sidney W. (2000). Dragonflies Through Binoculars: a field guide to the dragonflies of North America. Oxford University Press. ISBN 0-19-511268-7. 
  4. ^ Blanke, Alexander; Greve, Carola; Mokso, Rajmund; Beckmann, Felix; Misof, Bernhard (July 2013). "An updated phylogeny of Anisoptera including formal convergence analysis of morphological characters". Systematic Entomology 38 (3): 474–490. doi:10.1111/syen.12012. 
  5. ^ Ruppert, Edward E.; Fox, Richard, S.; Barnes, Robert D. (2004). Invertebrate Zoology, 7th edition. Cengage Learning. p. 745. ISBN 978-81-315-0104-7. 
  6. ^ Cynthia Berger (2004). Dragonflies. Stackpole Books. pp. 5–6. ISBN 978-0-8117-2971-0. 
  7. ^ Paulson, Dennis (19 December 2011). Dragonflies and Damselflies of the East. Princeton University Press. p. 32. ISBN 1-4008-3966-1. 
  8. ^ Powell 1999, p. 102.
  9. ^ Prum, Richard O.; Cole, Jeff A.; Torres, Rodolfo, H. (15 October 2004). "Blue integumentary structural colours in dragonflies (Odonata) are not produced by incoherent Tyndall scattering". Journal of Experimental Biology 207: 3999–4009. doi:10.1242/jeb.01240. 
  10. ^ Bybee, Seth (May 2012) [August 2005]. "Featured Creatures: dragonflies and damselflies". University of Florida. Retrieved 26 February 2015. 
  11. ^ Garrison, Rosser W.; Ellenrieder, Natalia von; Louton, Jerry A. (16 August 2006). Dragonfly Genera of the New World: An Illustrated and Annotated Key to the Anisoptera. JHU Press. p. 40. ISBN 978-0-8018-8446-7. 
  12. ^ "Emperor dragonfly (Anax imperator)". Arkive.org. Retrieved 26 February 2015. 
  13. ^ Powell 1999, p. 9.
  14. ^ a b c d Dijkstra 2006, pp. 8–9.
  15. ^ a b Cynthia Berger (2004). Dragonflies. Stackpole Books. pp. 32–34. ISBN 978-0-8117-2971-0. 
  16. ^ a b Cordero-Rivera, Adolfo; Cordoba-Aguilar, Alex (2010). 15. Selective Forces Propelling Genitalic Evolution in Odonata. p. 343. 
  17. ^ a b c Trueman & Rowe 2009, p. Life Cycle and Behavior.
  18. ^ a b c Cynthia Berger (2004). Dragonflies. Stackpole Books. p. 39. ISBN 978-0-8117-2971-0. Romantic souls are pleased to note that at the climactic moment, the two slender bodies form a delicate heart shape. Experts say the pair is now "in cop". 
  19. ^ Dragon fly naiad labium extended to capture prey
  20. ^ Mill, P. J.; Pickard, R. S. (1975). "Jet-propulsion in anisopteran dragonfly larvae". Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology 97 (4): 329–338. doi:10.1007/BF00631969. 
  21. ^ Grzimeck, HC; Bernard (1975). Grzimek's Animal Life Encyclopedia Vol 22. Detroit: Visible Ink Press. p. 348. 
  22. ^ Cynthia Berger (2004). Dragonflies. Stackpole Books. pp. 10–11. ISBN 978-0-8117-2971-0. 
  23. ^ Cynthia Berger (2004). Dragonflies. Stackpole Books. p. 9. ISBN 978-0-8117-2971-0. 
  24. ^ Foster, S.E; Soluk, D.A (2006). "Protecting more than the wetland: The importance of biased sex ratios and habitat segregation for conservation of the Hine's emerald dragonfly, Somatochlora hineana Williamson". Biological Conservation: 158–166. 
  25. ^ "Dragonflies". BBC Nature: Wildlife. Retrieved 15 July 2014. 
  26. ^ Waldbauer, Gilbert (2006). A Walk Around the Pond: Insects in and Over the Water. Harvard University Press. p. 105. ISBN 9780674022119. 
  27. ^ a b c d e f g Rowe, Richard J. "Dragonfly Flight". Tree of Life. Retrieved 26 February 2015. 
  28. ^ Tillyard, Robert John (1917). The Biology of Dragonflies. pp. 322–323. Retrieved 15 December 2010. I doubt if any greater speed than this occurs amongst Odonata 
  29. ^ Dean, T. J. (2003-05-01). "Chapter 1 — Fastest Flyer". Book of Insect Records. University of Florida. 
  30. ^ s "Frequently Asked Questions about Dragonflies". British Dragonfly Society. Retrieved 5 July 2011. 
  31. ^ Hopkin, Michael (June 5, 2003). "Nature News". Dragonfly flight tricks the eye. Nature.com. Retrieved January 16, 2012. 
  32. ^ Mizutani, A. K., Chahl, J. S. & Srinivasan, M. V. (June 5, 2003). "Insect behaviour: Motion camouflage in dragonflies". Nature 65 (423): 604. doi:10.1038/423604a. 
  33. ^ Glendinning, Paul (27 January 2004). "Motion Camouflage". The mathematics of motion camouflage. The Royal Society. 
  34. ^ Powell 1999, p. 12.
  35. ^ a b Berger, Cynthia (2004). Dragonflies. Stackpole Books. pp. 48–49. ISBN 978-0-8117-2971-0. 
  36. ^ "Hobby". BBC Nature. Retrieved 26 February 2015. 
  37. ^ Mead, Kurt. "Dragonfly Biology 101". Minnesota Dragonfly Society. Retrieved 27 February 2015. 
  38. ^ Córdoba-Aguilar, Alex (28 August 2008). Dragonflies and Damselflies: Model Organisms for Ecological and Evolutionary Research. OUP Oxford. p. 176. ISBN 978-0-19-155223-6. 
  39. ^ "An Introduction To The Study of Invertebrate Zoology Animalia, SubKingdom: Metazoa,. Platyhelminthes". University of California, Riverside. Retrieved 27 February 2015. 
  40. ^ Taku, Kadoya; Shin-ichi, Suda; Izumi, Washitani (2009). "Dragonfly Crisis in Japan: A likely Consequence of Recent Agricultural Habitat Degradation". Biological Conservation 142 (9): 1889–1905. 
  41. ^ Bambaradeniya, Channa N. B.; Amarasinghe, Felix P. (2003). "Biodiversity Associated with the Rice Field Agro-ecosystem in Asian countries: A Brief Review". International Water Management Institute 63. 
  42. ^ Kobori, Hiromi; Primack, Richard B. (2003). "Participatory Conservation Approaches for Satoyama, the Traditional Forest and Agricultural Landscape of Japan". Ambio 4: 307–311. 
  43. ^ Washitani, Izumi (2008). "Restoration of Biologically-Diverse Floodplain Wetlands Including Paddy Fields". Global Environmental Research 12: 95–99. 
  44. ^ Kurechi, Masayuki (2007). "Restoring Rice Paddy Wetland Environments and the Local Sustainable Society-Project for Achieving Co-Existence of Rice Paddy Agriculture with Waterbirds at Kabukuri-numa, Miyagi Prefecture, Japan". Global Environmental Research 11 (2): 141. 
  45. ^ Simaika, John P.; Samways, Michael J.; Kipping, Jens; Suhling, Frank; Dijkstra, Klaas-Douwe B.; Clausnitzer, Viola; Boudot, Jean Pierre; Domisch, Sami (2013). "Continental-Scale Conservation Prioritization of African Dragonflies". Biological Conservation 157: 245–254. 
  46. ^ Soluk, Daniel; Zercher, Deanna; Worthington, Amy (2011). "Influence of roadways on patterns of mortality and flight behavior of adult dragonflies near wetland areas". Elsevier. 
  47. ^ Soluk, Daniel A. (2011). "Influence of roadways on patterns of mortality and flight behavior of adult dragonflies near wetland areas". Biological Conservation: 1638–43. 
  48. ^ a b c Mitchell, Forrest L.; Lasswell, James L. (2005). A Dazzle of Dragonflies. College Station, TX: Texas A&M University Press. ISBN 1-58544-459-6. 
  49. ^ a b c Corbet, Phillip S. (1999). Dragonflies: Behavior and Ecology of Odonata. Ithaca, NY: Cornell University Press. pp. 559–561. ISBN 0-8014-2592-1. 
  50. ^ Lehmann, Tracy Hobson (19 June 2008). "Dragonflying: the new birding". San Antonio Express-News. 
  51. ^ Moonan, Wendy (August 13, 1999). "Dragonflies Shimmering as Jewelry". New York Times. pp. E2:38. 
  52. ^ Large, Elizabeth (June 27, 1999). "The latest buzz; In the world of design, dragonflies are flying high". The Sun (Baltimore, MD). pp. 6N. 
  53. ^ Brown, Roland (November–December 2007). "1955 Douglas Dragonfly". Motorcycle Classics. Retrieved 2009-08-10. 
  54. ^ Baird, Merrily (2001). Symbols of Japan: Thematic Motifs in Art and Design. New York: Rizzoli. pp. 108–9. ISBN 0-8478-2361-X. 
  55. ^ Nussbaum, Louis Frédéric et al. (2005). "Akitsushima" in Japan encyclopedia, p. 20., p. 20, at Google Books
  56. ^ Nihonto
  57. ^ 杉浦洋一 (Youichi Sugiura); John K. Gillespie (ジョン・K・ギレスピー) (1999). 日本文化を英語で紹介する事典: A Bilingual Handbook on Japanese Culture (in Japanese & English). 日本国東京都千代田区 (Chiyoda, JP-13): 株式会社ナツメ社 (Kabushiki gaisha Natsume Group). p. 305. ISBN 4-8163-2646-4. Retrieved 2010-04-26. 
  58. ^ Trueman, John W. H.; Rowe, Richard J. "Odonata: Dragonflies and Damselflies". Tree of Life. Retrieved 27 February 2015. 
  59. ^ Hand, Wayland D. (1973). "From Idea to Word: Folk Beliefs and Customs Underlying Folk Speech". American Speech 48 (1/2): 67–76. doi:10.2307/3087894. JSTOR 3087894. 
  60. ^ a b Waldbauer, Gilbert; Waldbauer, Gilbert (30 June 2009). A Walk around the Pond: insects in and over the water. Harvard University Press. p. 247. ISBN 978-0-674-04477-7. 
  61. ^ Mitchell, Forrest Lee; Lasswell, James (2005). A Dazzle Of Dragonflies. Texas A&M University Press. p. 36. ISBN 978-1-58544-459-5. 
  62. ^ Tennyson, Alfred, Lord (17 November 2013). Delphi Complete Works of Alfred, Lord Tennyson (Illustrated). Delphi Classics. pp. 544–545. ISBN 978-1-909496-24-8. 
  63. ^ "Down the River". H. E. Bates Companion. Retrieved 27 February 2015. 
  64. ^ Powell 1999, p. 7.
  65. ^ Bates, H. E. (12 February 1937). "Country Life: Pike and Dragonflies". The Spectator (5668): 269 (online p. 17). 

Sources[edit]

  • Dijkstra, Klaas-Douwe B. (2006). Field Guide to the Dragonflies of Britain and Europe. British Wildlife Publishing. ISBN 0-9531399-4-8. 
  • 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]