Wikipedia

Damselfly

Damselfliesareflying insectsof the suborderZygopterain the orderOdonata.They are similar todragonflies(which constitute the other odonatan suborder,Epiprocta) but are usually smaller and have slimmer bodies. Most species fold the wings along the body when at rest, unlike dragonflies which hold the wings flat and away from the body. Damselflies have existed since theLate Jurassic,[2]and are found on every continent exceptAntarctica.

Damselfly
Temporal range:Kimmeridgian–Recent
Male beautiful demoiselle
(Calopteryx virgo)
Female bluetail damselfly
(Ischnura heterosticta)
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Odonata
Suborder: Zygoptera
Selys,1854[1]
Superfamilies
$indicatesparaphyleticgroups

All damselflies arepredatory insects:bothnymphsand adults actively hunt and eat other insects. The nymphs are aquatic, with different species living in a variety offreshwater habitatsincluding acidicbogs,ponds,lakesandrivers.The nymphsmoultrepeatedly, at the last moult climbing out of the water to undergometamorphosis.The skin splits down the back, they emerge and inflate their wings and abdomen to gain their adult form. Their presence on a body of water indicates that it is relatively unpolluted, but their dependence on freshwater makes them vulnerable to damage to theirwetlandhabitats.

Some species of damselfly have elaboratecourtshipbehaviours. Many species aresexually dimorphic,the males often being more brightly coloured than the females. Like dragonflies, they reproduce using indirectinseminationand delayedfertilisation.A mating pair form a shape known as a "heart" or "wheel", the male clasping the female at the back of the head, the female curling her abdomen down to pick up sperm from secondary genitalia at the base of the male's abdomen. The pair often remain together with the male still clasping the female while she lays eggs within the tissue of plants in or near water using a robustovipositor.

Artificial fishing fliesthat mimic damselfly nymphs are used in wet-fly fishing.Damselflies are sometimes represented in personal jewellery such as brooches.

Classification

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The Zygoptera are an ancient group, with the earliest fossils dating to theKimmeridgianage of theLate Jurassic,around 152 million years ago.[2]Well-preservedEocenedamselfly larvae and exuviae are known from fossils preserved inamberin the Baltic region.[3]

Molecular analysis in 2021 confirms that most of the traditional families aremonophyletic,but shows that theAmphipterygidae,MegapodagrionidaeandProtoneuridaeareparaphyleticand will need to be reorganised. The Protoneuridae in particular is shown to be composed of six clades from five families. The result so far is 27 damselfly families, with 7 more likely to be created. The discoveredcladesdid not agree well with traditional characteristics used to classify living and fossil Zygoptera such as wing venation, so fossil taxa will need to be revisited. The 18 extant traditional families are provisionally rearranged as follows (the 3 paraphyletic families disappearing, and many details not resolved):[4]

Zygoptera
Lestoidea

Hemiphlebiidae(ancient greenling)

Perilestidae(shortwings)

Synlestidae(sylphs)

Lestidae(spreadwings)

Platystictoidea

Platystictidae(shadowdamsels)

"Calopterygoidea", revised

Calopterygidae(demoiselles)

Chlorocyphidae(jewels)

Dicteriadidae(barelegs)

Polythoridae(bannerwings)

13 more families

Euphaeidae(odalisques)

Lestoideidae(bluestreaks)

8 possible familiesincertae sedis

Coenagrionoidea

Isostictidae(narrow-wings)

Platycnemididae(white-legged damselflies)

Coenagrionidae(inc.Pseudostigmatidae) (pond damselflies)

General description

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Illustration of damselfly wing. Both fore and hind wings are similar.

The general body plan of a damselfly is similar to that of a dragonfly. Thecompound eyesare large but are more widely separated and relatively smaller than those of a dragonfly. Above the eyes is the frons or forehead, below this the clypeus, and on the lower lip or labium, an extensible organ used in the capture ofprey.The top of the head bears three simple eyes (ocelli), which may measure light intensity, and a tiny pair ofantennaethat serve no olfactory function but may measure air speed.[5]Many species aresexually dimorphic;the males are often brightly coloured and distinctive, while the females are plainer,crypticallycoloured, and harder to identify to species. For example, inCoenagrion,the Eurasian bluets, the males are bright blue with black markings, while the females are usually predominantly green or brown with black.[6]A few dimorphic species show female-limited polymorphism, the females being in two forms, one form distinct and the other with the patterning as in males. The ones that look like males, andromorphs, are usually under a third of the female population but the proportion can rise significantly and a theory that explains this response suggests that it helps overcome harassment by males.[7]Some Coenagrionid damselflies show male-limited polymorphism, an even less understood phenomenon.[8]

In general, damselflies are smaller than dragonflies, the smallest being members of the genusAgriocnemis(wisps).[9]However, members of thePseudostigmatidae(helicopter damselflies or forest giants) are exceptionally large for the group, with wingspans as much as 19 cm (7.5 in) inMegaloprepus[10]and body length up to 13 cm (5.1 in) inPseudostigma aberrans.[11]

Terminal segments of the abdomen of a male damselfly (Pseudagrion caffrum) showing segments 8–10 (S8, S9, S10), the upper or superior appendages or cerci (c) and the inferior appendages or paraprocts (p)
Damselfly head displayingcompound eyes,ocelli,antennae,andmouth structures

The firstthoracicsegment is the prothorax, bearing the front pair of legs. The joint between head and prothorax is slender and flexible, which enables the damselfly to swivel its head and to manoeuvre more freely when flying. The remaining thoracic segments are the fused mesothorax and metathorax (together termed thesynthorax), each with a pair of wings and a pair of legs. A dark stripe known as the humeral stripe runs from the base of the front wings to the second pair of legs, and just in front of this is the pale-coloured, antehumeral stripe.[5]

Male (left) and female of the banded demoiselle,Calopteryx splendens,showing theirdifferently colouredwings

The forewings and hindwings are similar in appearance and are membranous, being strengthened and supported by longitudinal veins that are linked by many cross-veins and that are filled withhaemolymph.[12]Species markers include quadrangular markings on the wings known as thepterostigmaor stigma, and in almost all species, there is anodusnear the leading edge. The thorax houses the flight muscles.[5]Many damselflies (e.g. Lestidae, Platycnemidae, Coenagrionidae) have clear wings, but some (Calopterygidae, Euphaeidae) have coloured wings, whether uniformly suffused with colour or boldly marked with a coloured patch. In species such as the banded demoiselle,Calopteryx splendensthe males have both a darker green body and large dark violet-blue patches on all four wings, which flicker conspicuously in their aerial courtship dances; the females have pale translucent greenish wings.[13]

The abdomen is long and slender and consists of ten segments. The secondary genitalia in males are on the undersides of segments two and three and are conspicuous, making it easy to tell the sex of the damselfly when viewed from the side. The female genital opening is on the underside between segments eight and nine. It may be covered by a subgenital plate, or extended into a complex ovipositor that helps them lay eggs within plant tissue. The tenth segment in both sexes bearscerciand in males, its underside bears a pair of paraprocts.[5]

Damselflies (except spreadwings,Lestidae) rest their wings together, above their bodies, whereas dragonflies rest with their wings spread diametrically apart; the spreadwings rest with their wings slightly apart. Damselflies have slenderer bodies than dragonflies, and their eyes do not overlap. Damselfly nymphs differ from dragonflies nymphs in that theepiproctand pair ofparaproctsat the tip of their abdomen has been modified into caudal gills, in addition to being able to absorb oxygen through the wall of their rectum, whereas dragonflies breathe through internal rectal gills only.[14][15]Damselfly nymphs swim by fish-like undulations, the gills functioning like a tail. Dragonfly nymphs can forcibly expel water in their rectum for rapid escape.[16]

Distribution and diversity

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Odonates are found on all the continents except Antarctica.[17]Although some species of dragonfly have wide distributions, damselflies tend to have smaller ranges. Most odonates breed in fresh-water; a few damselflies in the family Caenagrionidae breed in brackish water (and a single dragonfly species breeds in seawater).[18][19]Dragonflies are more affected by pollution than are damselflies. The presence of odonates indicates that an ecosystem is of good quality. The most species-rich environments have a range of suitable microhabitats, providing suitable water bodies for breeding.[20][21]

Although most damselflies live out their lives within a short distance of where they were hatched, some species, and some individuals within species, disperse more widely.Forktailsin the familyCoenagrionidaeseem particularly prone to do this, large maleboreal bluets(Enallagma boreale) in British Columbia often migrating, while smaller ones do not.[22]These are known to leave their waterside habitats, flying upwards till lost from view, and presumably being dispersed to far off places by the stronger winds found at high altitudes.[22]In this way they may appear in a locality where no damselflies were to be seen the day before.Rambur's forktail(Ischnura ramburii) has been found, for example, onoil rigsfar out in theGulf of Mexico.[5]

The distribution and diversity of damselfly species in thebiogeographicalregions is summarized here. (There are no damselflies in theAntarctic.) Note that some species are widespread and occur in multiple regions.[21]

Overall, there are about 2942 extant species of damselflies placed in 309 genera.[21]

Biology

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Blue damselflies swarming over pond inKew Gardens.Some pairs (blue male in front) are seen.
Western forktail damselflyattempting on-the-fly catches. Each repeated in slow motion. The second prey landed, escaping capture. Finally closeup devouring medium sized fly.
Damselfly eating a Crane fly

Adult damselflies catch and eat flies, mosquitoes, and other small insects. Often they hover among grasses and low vegetation, picking prey off stems and leaves with their spiny legs (unlike dragonflies which prefer catching flying prey).[22][23]Although predominantly using vision to locate their prey, adults may also make use of olfactory cues.[24]No species are known to hunt at night, but some are crepuscular, perhaps taking advantage of newly hatched flies and other aquatic insects at a time when larger dragonflies are roosting.[25]In tropical South America, helicopter damselflies (Pseudostigmatidae) feed onspiders,hovering near an orb web and plucking the spider, or its entangled prey, from the web.[26]There are few pools and lakes in these habitats, and these damselflies breed in temporary water bodies in holes in trees, the rosettes ofbromeliadsand even the hollow stems ofbamboos.[27]

The nymphs of damselflies have been less researched than their dragonfly counterparts, and many have not even been identified. They choose their prey according to size and seem less able to overpower larger prey than can dragonfly nymphs. The major part of the diet of most species appears to becrustaceanssuch aswater fleas.[25]

Ecology

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Fine damselfly habitat: panorama ofThursley Common,looking over the acid bog pools

Damselflies exist in a range of habitats in and around the wetlands needed for their larval development; these include open spaces for finding mates, suitable perches, open aspect, roosting sites, suitable plant species for ovipositing and suitable water quality. Odonates have been used forbio-indicationpurposes regarding the quality of theecosystem.Different species have different requirements for their larvae with regard to water depth, water movement and pH.[28] The Europeancommon blue damselfly(Enallagma cyathigerum) for example can occur at high densities inacidwaters where fish are absent, such as in bog pools.[29] Thescarce blue-tailed damselfly(Ischnura pumilio) in contrast requiresbase-richhabitats and water with a slow flow-rate. It is found in ditches, quarries, seeps, flushes, marshes and pools. It tolerates high levels of zinc and copper in the sediment but requires suitable emergent plants for egg-laying without the water being choked by plants.[28]Damselflies' dependence on freshwater habitats makes them very vulnerable to damage to wetlands through drainage for agriculture or urban growth.[30]

In the tropics, the helicopter damselflyMecistogaster modesta(Pseudostigmatidae) breeds inphytotelmata,the small bodies of water trapped bybromeliads,epiphyticplants of therainforestof northwestCosta Rica,at the high density of some 6000 larvae per hectare in patches of secondary forest.[31]Another tropical species, the cascade damselflyThaumatoneura inopinata(Megapodagrionidae), inhabitswaterfallsin Costa Rica and Panama.[32][33]

Damselflies, both nymphs and adults, are eaten by a range ofpredatorsincluding birds, fish, frogs, dragonflies, other damselflies,water spiders,water beetles,backswimmersandgiant water bugs.[22]

Damselfly grooming
Water mites parasitizingPseudagrion decorum

Damselflies have a variety of internal and external parasites. Particularly prevalent are thegregarine protozoansfound in the gut. In a study of the European common blue damselfly, every adult insect was infected at the height of the flying season. When present in large numbers, these parasites can cause death by blocking the gut.[22]Water mitesHydracarinaare often seen on the outside of both nymphs and adults, and can move from one to the other at metamorphosis.[22]They suck the body fluids and may actually kill young nymphs, but adults are relatively unaffected, it being necessary for the completion of the mite's life cycle that it returns to water, a feat accomplished when the adult damselfly breeds.[34]

Behaviour

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Female blue-fronted dancer

Many damselflies have elaborate courtship behaviours. These are designed to show off the male's distinctive characteristics, bright colouring or flying abilities, thus demonstrating his fitness.Calopteryxmales will hover in front of a female with alternating fast and slow wingbeats; if she is receptive she will remain perched, otherwise she will fly off. The maleriver jewelwing(Calopteryx aequabilis) performs display flights in front of the female, fluttering his forewings while keeping his hindwings still, and raising his abdomen to reveal the white spots on his wings.[35]Platycyphamales will hover in front of a female, thrusting their bright white legs forward in front of their heads. Flattened tibia and bright leg colouring are seen inPlatycnemis phasmovolansand a few otherPlatycnemididae,including the extinctYijenplatycnemis huangi.[36]Rhinocyphabobs up and down, often low over fast-flowing forested and shaded streams, displaying its bright-coloured body and wings.[37]Some species (R. biceriata,R. humeralis) have a foot waggling behaviour: they thrust a leg forward and vibrate it towards ovipositing females while in flight.[38]Vibrating the tibia is seen inLibellago semiopacadespite it lacking bright colouration on the tibia suggesting that foot waggling is a generalized excitary signal in Chlorocyphidae damselflies.[38]Foot waggling has been observed inCalopteryxsp.,Platycypha fitzsimonsi,[39]andPlatycypha caligata.Male members of the familyProtoneuridaewith vividly coloured wings display these to visiting females.[37]Swift forktail(Ischnura erratica) males display to each other with their blue-tipped abdomens raised.[40]

Euphaea fraserimalesdefend their territory,with open wings and prominentlydisplaythe iridescent copper markings of the upper hind wing.

Other behaviours observed in damselflies include wing-warning, wing-clapping, flights of attrition and abdominal bobbing. Wing-warning is a rapid opening and closing of the wings and is aggressive, while wing-clapping involves a slower opening of the wings followed by a rapid closure, up to eight times in quick succession, and often follows flight; it may serve a thermo-regulatory function.[41]Flights of attrition are engaged in by theebony jewelwing(Calopteryx maculata) and involve males bouncing around each other while flying laterally and continuing to do so, sometimes over a considerable distance, until one insect is presumably exhausted and gives up.[42]Characteristics of displays and coloration of males are suggested to be the common cues used by females to choose mates. In at least one species,Mnais costalis,males with more sunlight in their territories had higher wing-beat frequency and were more likely to mate. Females preferred "hotter" males because they would be on warmer territories for egg laying.[43]

At night, damselflies usually roost in dense vegetation, perching with the abdomen alongside a stem. If disturbed they will move around to the other side of the stem but will not fly off. Spreadwings fully fold their wings when roosting.[5]Thedesert shadowdamsel(Palaemnema domina) aggregates to roost in thick places near streams in the heat of the day. While there it engages in wing-clapping, the exact function of which is unknown.[44]Some species such as the rubyspot damselfly,Hetaerina americana,form night roosting aggregations, with a preponderance of males; this may have an anti-predator function or may be simply the outcome of choosing safe roosting sites.[45]

Reproduction

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First stage:Ceriagrion cerinorubellumpair with male transferring sperm from the primary to the secondary sex organs
Second stage: "heart" or "wheel" position, the female receiving sperm from the male secondary sex organ

Mating in damselflies, as in dragonflies, is a complex, precisely choreographed process involving both indirect insemination and delayed fertilisation.[46][47]The male first 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.[47][48]The pair fly 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";[46][49]the pair may also be described as being "in cop".[50]Males may transfer the sperm to their secondary genitalia either before a female is held, in the early stage when the female is held by the legs or after the female is held between the terminal claspers. This can lead to variations in the tandem postures.[51]The spermatophore may also have nutrition in addition to sperms as a "nuptial gift".[52]Some cases ofsexual cannibalismexist where females (ofIschnura graellsii) eat males while in copula.[53]

Parthenogenesis(reproduction from unfertilised eggs) is exceptional, and has only been recorded in nature in femaleIschnura hastataon theAzores Islands.[21][54]

Yellow-striped blue dart,Pseudagrion indicum,laying eggs; the male (above) continues to hold the female with his claspers.

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, mate-guarding, or in some species continuing to clasp her and flying in tandem. The male attempts to prevent rivals from removing his sperm and inserting their own,[55]a form ofsperm competition(the sperms of the last mated male have the greatest chance of fertilizing the eggs, also known as sperm precedence[56]) made possible by delayed fertilisation[46][49]and driven bysexual selection.[47][48]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 remain in the heart posture.[50]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.[55]

Willow emerald,Chalcolestes viridis,still in tandem, laying eggs into a series of slits in a twig

All damselflies lay their eggs inside plant tissues; those that lay eggs underwater may submerge themselves for 30 minutes at a time, climbing along the stems of aquatic plants and laying eggs at intervals.[57]For example, the red-eyed damselflyErythromma najaslays eggs, in tandem, into leaves or stems of floating or sometimes emergent plants; in contrast, the scarce bluetailIschnura pumiliooviposits alone, the female choosing mostly emergent grasses and rushes, and laying her eggs in their stems either above or just below the waterline.[58]The willow emeraldChalcolestes viridis(a spreadwing) is unusual in laying eggs only in woody plant tissue, choosing thin twigs of trees that hang over water, and scarring the bark in the process.[59]A possible exception is an apparent instance ofovo-viviparity,in whichHeliocypha perforatawas filmed in western China depositing young larvae (presumably hatched from eggs inside the female's body) onto a partly submerged branch of a tree.[60]

Many damselflies are able to produce more than one brood per year (voltinism); this is negatively correlated withlatitude,becoming more common towards the equator, except in the Lestidae.[61]

Life cycle

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Damselflies arehemimetabolousinsects that have no pupal stage in their development.[62]The female inserts the eggs by means of her ovipositor into slits made in water plants or other underwatersubstratesand the larvae, known as naiads or nymphs, are almost all completely aquatic.[5]Exceptions include the HawaiianMegalagrion oahuenseand an unidentified Megapodagrionid from New Caledonia,[63]which are terrestrial in their early stages.[55]Thespreadwingslay eggs above the waterline late in the year and the eggs overwinter, often covered by snow. In spring they hatch out in the meltwater pools and the nymphs complete their development before these temporary pools dry up.[22]

Blue-tailed damselfly (Ischnura elegans) nymph, showing the three tail appendages
Prodasineura verticalisfemale emerging from thesplit skin of the nymph

The nymphs are voraciouspredatorsand feed by means of a flatlabium(a toothed mouthpart on the lower jaw) that forms the so-called mask; it is rapidly extended to seize and pierce theDaphnia(water fleas),mosquitolarvae, and other small aquatic organisms on which damselfly nymphs feed. They breathe by means of three large external, fin-like gills on the tip of the abdomen, and these may also serve for locomotion in the same manner as a fish's tail.[5]Compared to dragonfly larvae, the nymphs show little variation in form. They tend to be slender and elongate, many having morphological adaptations for holding their position in fast flowing water. They are more sensitive than dragonfly nymphs to oxygen levels and suspended fine particulate matter, and do not bury themselves in the mud.[25]

The nymphs proceed through about a dozen moults as they grow. In the later stages, the wing pads become visible. When fully developed, the nymphs climb out of the water and take up a firm stance, the skin on the thorax splits and the adult form wriggles out. This has a soft body at first and hangs or stands on its empty larval case. It pumps haemolymph into its small limp wings, which expand to their full extent. The haemolymph is then pumped back into the abdomen, which also expands fully. Theexoskeletonhardens and the colours become more vivid over the course of the next few days. Most damselflies emerge in daytime, and in cool conditions the process takes several hours. On a hot day, the cuticle hardens rapidly and the adult can be flying away within half an hour.[5]

Conservation

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Conservation of Odonata has usually concentrated on the more iconic suborder Anisoptera, the dragonflies. However, the two suborders largely have the same needs, and what is good for dragonflies is also good for damselflies. The main threats experienced by odonates are the clearance of forests, the pollution of waterways, the lowering of groundwater levels, the damming of rivers forhydroelectric schemesand the general degradation of wetlands and marshes.[64]The clearance of tropical rainforests is of importance because the rate of erosion increases, streams and pools dry up and waterways become clogged with silt. The presence of alien species can also haveunintended consequences.[64]In Hawaii, the introduction of themosquitofish(Gambusia affinis) was effective in controlling mosquitoes but nearly exterminated the island's endemic damselflies.[65]The ancient greenlingHemiphlebia mirabilishas been an important flagship species for conservation action in preserving its habitat in Australia.[66]

In culture

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Fishing fliesthat mimic damselfly nymphs are sometimes used in wet-fly fishing, where the hook and line are allowed to sink below the surface.[67]

Damselflies have formed subjects for personaljewellerysuch as brooches since at least 1880.[68]

Damselflyis a 2005 short film directed by Ben O'Connor.[69]

Damselflyis the title of a 2012 novel in theFaebleseries by S. L. Naeole[70]and of a 2018 novel by Chandra Prasad.[71]

Modern poems with the damselfly as a subject include a 1994 poem byAugust Kleinzahler,which contains the lines "And that blue there, cobalt / a moment, then iridescent, / fragile as a lady's pin / hovering above the nasturtium?"[72]The poetJohn Engelspublished “Damselfly, Trout, Heron” in his 1983 collectionWeather-Fear: New and Selected Poems.[73]

See also

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Citations

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  1. ^Selys-Longchamps, E. (1854).Monographie des caloptérygines(in French). Brussels and Leipzig: C. Muquardt. pp. 1–291 [2].doi:10.5962/bhl.title.60461.hdl:2027/hvd.32044107312183.
  2. ^abBechly, G. (2019). "New fossil Odonata from the Upper Jurassic of Bavaria with a new fossil calibration point for Zygoptera".Palaeoentomology.2(6): 618–632.doi:10.11646/palaeoentomology.2.6.13.S2CID210159976.
  3. ^Bechly, Günter; Wichard, Wilfried (30 December 2008)."Damselfly and dragonfly nymphs in Eocene Baltic amber (Insecta: Odonata), with aspects of their palaeobiology"(PDF).Palaeodiversity.1:37–73. Archived fromthe original(PDF)on 2 April 2015.Retrieved16 March2015.
  4. ^Bybee, Seth M.; Kalkman, Vincent J.; Erickson, Robert J.; et al. (2021)."Phylogeny and classification of Odonata using targeted genomics".Molecular Phylogenetics and Evolution.160.Elsevier BV: 107115.Bibcode:2021MolPE.16007115B.doi:10.1016/j.ympev.2021.107115.hdl:11093/2768.ISSN1055-7903.PMID33609713.
  5. ^abcdefghiPaulson, Dennis (2011).Dragonflies and Damselflies of the East.Princeton University Press. pp. 10–32.ISBN978-1-4008-3966-7.
  6. ^Dijkstra 2006,pp. 20, 104.
  7. ^Gossum, Hans Van; Sherratt, Thomas N. (2008). "A dynamical model of sexual harassment in damselflies and its implications for female-limited polymorphism".Ecological Modelling.210(1–2): 212–220.Bibcode:2008EcMod.210..212V.doi:10.1016/j.ecolmodel.2007.07.023.
  8. ^Gossum, Hans Van; Sherratt, Tom N.; Cordero-Rivera, Adolfo (2008)."The evolution of sex-limited colour polymorphism".In Cordoba-Aguilar, Alex (ed.).Dragonflies and Damselflies. Model organisms for ecological and evolutionary research.Oxford University Press. pp. 219–229.ISBN9780199230693.
  9. ^Kipping, Jens; Martens, Andreas; Suhling, Frank (2012). "Africa's smallest damselfly – a newAgriocnemisfrom Namibia ".Organisms Diversity & Evolution.12(3): 301–306.doi:10.1007/s13127-012-0084-4.S2CID13356379.
  10. ^Groenevelda, Linn F.; Viola Clausnitzerb; Heike Hadrysa (2007). "Convergent Evolution of Gigantism in Damselflies of Africa and South America? Evidence from Nuclear and Mitochondrial Sequence Data".Molecular Phylogenetics and Evolution.42(2): 339–46.Bibcode:2007MolPE..42..339G.doi:10.1016/j.ympev.2006.05.040.PMID16945555.
  11. ^Hedström, Ingemar; Göran Sahlén (2001)."A key to the adult Costa Rican" helicopter "damselflies (Odonata: Pseudostigmatidae) with notes on their phenology and life zone preferences".Rev. Biol. Trop.49(3–4): 1037–1056.PMID12189786.
  12. ^Silsby, Jill (2001).Dragonflies of the World.Csiro Publishing. p. 9.ISBN978-0-643-10249-1.
  13. ^Dijkstra 2006,pp. 23, 65–67.
  14. ^Dragonflies and Damselflies of Texas and the South-Central United States: Texas, Louisiana, Arkansas, Oklahoma, and New Mexico - Princeton University
  15. ^Physiology of Insects
  16. ^Borror, Donald J.; Triplehorn, Charles A.; Triplehorn, Norman F.Study of Insects(6 ed.). Philadelphia: Saunders College Publishing. pp. 187–201.
  17. ^Nilsson, Anders (1997).Aquatic insects of North Europe: A taxonomic handbook.Apollo Books. p. 14.ISBN978-87-88757-07-1.
  18. ^Osburn, Raymond C. (1906)."Observations and Experiments on Dragon-Flies in Brackish Water".The American Naturalist.40(474): 395–399.doi:10.1086/278632.
  19. ^Dunson, William A. (1980). "Adaptations of Nymphs of a Marine Dragonfly, Erythrodiplax berenice, to Wide Variations in Salinity".Physiological Zoology.53(4): 445–452.doi:10.1086/physzool.53.4.30157882.S2CID87698039.
  20. ^"Introduction to the Odonata".UCMP Berkeley. Archived fromthe originalon 27 August 2017.Retrieved13 March2015.
  21. ^abcdSuhling, F.; Sahlén, G.; Gorb, S.; Kalkman, V.J.; Dijkstra, K-D.B.; van Tol, J. (2015). "Order Odonata". In Thorp, James; D. Christopher Rogers (eds.).Ecology and general biology. Thorp and Covich's Freshwater Invertebrates(4 ed.). Academic Press. pp. 893–932.ISBN978-0-12-385026-3.
  22. ^abcdefgAcorn, John (2004).Damselflies of Alberta: Flying Neon Toothpicks in the Grass.University of Alberta. pp.9–15.ISBN978-0-88864-419-0.
  23. ^Dragonflies and Damselflies of Georgia and the Southeast
  24. ^Piersanti, Silvana; Frati, Francesca; Conti, Eric; Gaino, Elda; Rebora, Manuela; Salerno, Gianandrea (2014). "First evidence of the use of olfaction in Odonata behaviour".Journal of Insect Physiology.62:26–31.Bibcode:2014JInsP..62...26P.doi:10.1016/j.jinsphys.2014.01.006.PMID24486162.
  25. ^abcHeckman, Charles W. (2008).Encyclopedia of South American Aquatic Insects: Odonata - Zygoptera: Illustrated Keys to Known Families, Genera, and Species in South America.Springer. pp. 17, 31–33.ISBN978-1-4020-8176-7.
  26. ^Ingley, Spencer J.; Bybee, Seth M.; Tennessen, Kenneth J.; Whiting, Michael F.; Branham, Marc A. (2012). "Life on the fly: phylogenetics and evolution of the helicopter damselflies (Odonata, Pseudostigmatidae)".Zoologica Scripta.41(6): 637–650.doi:10.1111/j.1463-6409.2012.00555.x.S2CID82370392.
  27. ^Fincke, Ola M. (2006)."Use of Forest and Tree Species, and Dispersal by Giant Damselflies (Pseudostigmatidae): Their Prospects in Fragmented Forests"(PDF).In Adolfo Cordero Rivera (ed.).Fourth WDA International Symposium of Odonatology, Pontevedra (Spain), July 2005.Sofia—Moscow: Pensoft Publishers. pp. 103–125. Archived fromthe original(PDF)on 2007-06-28.
  28. ^abAllen, Katherine (2009).The ecology and conservation of threatened damselflies.The Environment Agency. pp. 1–6.ISBN978-1-84911-093-8.
  29. ^Dijkstra 2006,p. 102.
  30. ^Corbet, P. S. (1980). "Biology of Odonata".Annual Review of Entomology.25:189–217.doi:10.1146/annurev.en.25.010180.001201.
  31. ^Srivastava, Diane S.; Melnychuk, Michael C.; Ngai, Jacqueline T. (2005). "Landscape variation in the larval density of a bromeliad-dwelling zygopteran, Mecistogaster modesta (Odonata: Pseudostigmatidae)".International Journal of Odonatology.8(1): 67–79.Bibcode:2005IJOdo...8...67S.doi:10.1080/13887890.2005.9748244.S2CID53603745.
  32. ^Calvert, Philip P. (1914)."Studies on Costa Rican Odonata. V. The waterfall-dwellers: Thaumatoneura imagos and possible male dimorphism".Entomological News and Proceedings of the Entomological Section.25(8): 337–348.
  33. ^Paulson, D. R. (2009)."Thaumatoneura inopinata".IUCN Red List of Threatened Species.2009:e.T21721A9313826.doi:10.2305/IUCN.UK.2009-2.RLTS.T21721A9313826.en.Retrieved13 November2021.[permanent dead link‍]
  34. ^Proctor, Heather (2004).Aquatic Mites from Genes to Communities: From Genes to Communities.Springer Science & Business Media. pp. 79–84.ISBN978-1-4020-2703-1.
  35. ^Paulson 2009,p. 42.
  36. ^Zheng, Daran; Nel, André; Jarzembowski, Edmund A.; Chang, Su-Chin; Zhang, Haichun; Xia, Fangyuan; Liu, Haoying; Wang, Bo (2017)."Extreme adaptations for probable visual courtship behaviour in a Cretaceous dancing damselfly".Scientific Reports.7(1): 44932.Bibcode:2017NatSR...744932Z.doi:10.1038/srep44932.ISSN2045-2322.PMC5357891.PMID28317876.
  37. ^abSilsby, Jill (2001).Dragonflies of the World.Csiro Publishing. pp. 58–59.ISBN978-0-643-10249-1.
  38. ^abOrr, A. G. (1996)."Territorial and courtship displays in Bornean Chlorocyphidae (Zygoptera)"(PDF).Odonatologica.25(2): 119–141.
  39. ^Robertson, H. M. (1982)."Courtship displays and mating behaviour of three species of Chlorocyphida (Zygoptera)"(PDF).Odonatologica.11(1): 53–58.
  40. ^Paulson 2009,p. 108.
  41. ^Bick, George H.; Bick, Juanda C. (1961). "Demography and Behavior of the Damselfly,Argia Apicalis(Say), (Odonata: Coenagriidae) ".Ecology.46(4): 461–472.doi:10.2307/1934877.JSTOR1934877.
  42. ^Paulson 2009,p. 44.
  43. ^Tsubaki, Yoshitaka; Samejima, Yuka; Siva-Jothy, Michael T. (2010)."Damselfly females prefer hot males: higher courtship success in males in sunspots".Behavioral Ecology and Sociobiology.64(10): 1547–1554.doi:10.1007/s00265-010-0968-2.ISSN1432-0762.
  44. ^Paulson 2009,p. 185.
  45. ^Switzer, Paul V.; Grether, Gregory F. (2000)."Characteristics and Possible Functions of Traditional Night Roosting Aggregations in Rubyspot Damselflies"(PDF).Behaviour.137(4): 401–416.doi:10.1163/156853900502141.S2CID85986224.
  46. ^abcDijkstra 2006,pp. 8–9.
  47. ^abcBattin, Tom (1993)."The odonate mating system, communication, and sexual selection: A review".Bolletino di Zoologia.60(4): 353–360.doi:10.1080/11250009309355839.
  48. ^abCordero-Rivera, Adolfo; Cordoba-Aguilar, Alex (2010)."Selective Forces Propelling Genitalic Evolution in Odonata"(PDF).In Leonard, Janet; Alex Córdoba-Aguilar (eds.).The Evolution of Primary Sexual Characters in Animals.Oxford University Press. p. 343. Archived fromthe original(PDF)on 2015-02-25.Retrieved2015-03-16.
  49. ^abTrueman & Rowe 2009,p. Life Cycle and Behavior.
  50. ^abBerger 2004,p. 39: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'.
  51. ^Bick, G. H.; Bick, J. C. (1965). "Sperm Transfer in Damselflies (Odonata: Zygoptera)".Annals of the Entomological Society of America.58(4): 592.doi:10.1093/aesa/58.4.592.PMID5834678.
  52. ^Cordero Rivera, A; Cordoba-Aguilar, A (2010). "Selective forces propelling genitalic evolution in Odonata". In Leonard J; Cordoba-Aguilar, A (eds.).The evolution of primary characters in animals.New York: Oxford University Press. pp. 332–352.
  53. ^Cordero, Adolfo (1992)."Sexual Cannibalism in the Damselfly Species Ischnura graellsii (Odonata: Coenagrionidae)"(PDF).Entomologia Generalis.17(1): 17–20.doi:10.1127/entom.gen/17/1992/17.Archived fromthe original(PDF)on 2013-06-13.Retrieved2017-04-28.
  54. ^Lorenzo-Carballa, M. O.; Cordero-Rivera, A. (2009)."Thelytokous parthenogenesis in the damselflyIschnura hastata(Odonata, Coenagrionidae): genetic mechanisms and lack of bacterial infection ".Heredity.103(5): 377–384.doi:10.1038/hdy.2009.65.PMID19513091.
  55. ^abcCardé, Ring T.; Resh, Vincent H. (2012).A World of Insects: The Harvard University Press Reader.Harvard University Press.pp. 195–197.ISBN978-0-674-04619-1.
  56. ^Tennessen, K.J. (2009). "Odonata (Dragonflies, Damselflies)". In Resh, Vincent H.; Ring T. Cardé (eds.).Encyclopedia of Insects(2 ed.). Academic Press. pp. 721–729.
  57. ^Lawlor, Elizabeth P. (1999).Discover Nature in Water & Wetlands: Things to Know and Things to Do.Stackpole Books. pp. 88, 94–96.ISBN978-0-8117-2731-0.
  58. ^Smallshire, Dave; Swash, Andy (2014).Britain's Dragonflies: A Field Guide to the Damselflies and Dragonflies of Britain and Ireland: A Field Guide to the Damselflies and Dragonflies of Britain and Ireland.Princeton University Press. pp. 94–96.ISBN978-1-4008-5186-7.
  59. ^Dijkstra 2006,p. 84.
  60. ^Salindra, H. G.; Dayananda, K.; Kitching, Roger L. (2014)."Ovo-viviparity in the Odonata? The case of Heliocypha perforata (Zygoptera: Chlorocyphidae)".International Journal of Odonatology.17(4): 181–185.Bibcode:2014IJOdo..17..181D.doi:10.1080/13887890.2014.959076.S2CID83520585.
  61. ^Corbet, Philip S.; Suhling, Frank; Soendgerath, Dagmar (2006)."Voltinism of Odonata: a review".International Journal of Odonatology.9(1): 1–44.Bibcode:2006IJOdo...9....1C.doi:10.1080/13887890.2006.9748261.S2CID55299239.
  62. ^A Dictionary of Entomology.CABI. 2011. p. 679.ISBN978-1-84593-542-9.
  63. ^Willey, Ruth Lippitt (1955)."A terrestrial damselfly nymph (Medapodarionidae) from New Caledonia"(PDF).Psyche: A Journal of Entomology.62(4): 137–144.doi:10.1155/1955/39831.
  64. ^abMoore, N.W. (1997)."Dragonflies: status survey and conservation action plan"(PDF).International Union for Conservation of Nature. Archived fromthe original(PDF)on 12 April 2016.Retrieved17 March2015.
  65. ^Gagné, W.C. (1981). "Status of Hawaii endangered species: insects and land snails".ʻElepaio.42:31–36.
  66. ^New, Timothy Richard."The Hemiphlebia damselfly Hemiphlebia mirabilis Selys (Odonata, Zygoptera) as a flagship species for aquatic insect conservation in south-eastern Australia".The Victorian Naturalist.124(4): 269–272.
  67. ^Wada, Wes (2012)."Fishing Tips for the Juicebug Damsel Nymph".Fly foundry. Archived fromthe originalon 19 November 2016.Retrieved13 March2015.
  68. ^"Antique" Damselfly "Brooch in Silver-topped Gold with Ruby Eyes".Macklowe Gallery.Retrieved13 March2015.[dead link‍]
  69. ^"Ben O'Connor".British Council.Retrieved13 March2015.
  70. ^Naeole, S. L. (2012).Damselfly.Crystal Quill.
  71. ^"Damselfly".Marly Rusoff Literary Agency.Retrieved21 December2020.
  72. ^Kleinzahler, August (August 1994)."The Damselfly".Poetry Magazine.Retrieved13 March2015.
  73. ^Engels, John (1983).Damselfly, Trout, Heron.University of Georgia Press.Retrieved13 March2015.

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