Damselfly

Damselflies (suborder Zygoptera) are insects in the order Odonata. They are similar to dragonflies, but are smaller, have slimmer bodies, and most species close the wings along the body when at rest.

Damselflies have existed since at least the Lower Permian. Today they are found on every continent except Antarctica. Their presence on a body of water indicates that it is unpolluted.

All damselflies are predatory; both nymphs and adults eat other insects. The nymphs are aquatic, living in freshwater habitats including acid bogs, lakes and rivers. They moult repeatedly, at the last moult climbing out of the water and inflating their wings and abdomen to give them their adult shape.

Phylogeny
The Zygoptera are an ancient group, with fossils known from the lower Permian, at least 250 million years ago. All the fossils of that age are of adults, similar in structure to modern damselflies, so it is not known if their larvae were aquatic at that time. The earliest Odonata larval fossils are from the Mesozoic. Well-preserved Eocene damselfly larvae and exuviae are known from fossils preserved in amber in the Baltic region.

Molecular analysis in 2013 confirms that most of the traditional families are monophyletic, but shows that the Amphipterygidae, Megapodagrionidae and Protoneuridae are paraphyletic and 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 discovered clades did 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 rearranged as follows (the 3 paraphyletic families disappearing):

Dashed lines indicate unresolved relationships.

General description
The general body plan of a damselfly is similar to that of a dragonfly. The compound eyes are 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 upper lip the labrum, an extensible organ used in the capture of prey. The top of the head bears three simple eyes (ocelli), which may measure light intensity, and a tiny pair of antennae that serve no olfactory function but may measure air speed.

The first thoracic segment 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 the synthorax), 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. The forewings and hindwings are similar in appearance and are membranous, being supported by a network of veins into which haemolymph flows. Species markers include quadrangular markings on the wings known as the pterostigma or stigma, and in almost all species, there is a nodus near the leading edge. The thorax houses the flight muscles.

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. The tenth segment in both sexes bears cerci and in males, its underside bears a pair of paraprocts.

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 they possess caudal gills (on the abdomen) whereas dragonflies breathe through the rectum.

Distribution
Odonates are found on all the continents except Antarctica. Although some species of dragonfly have wide distributions, damselflies tend to have smaller ranges. Dragonflies are more affected by pollution than are damselflies and the presence of dragonflies and damselflies indicates that the ecosystem is of good quality. The most species rich environments have a range of suitable microhabitats and suitable water bodies for breeding.

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. Forktails in the family Coenagrionidae seem particularly prone to do this, large male boreal bluets (Enallagma boreale) in British Columbia often migrating, while smaller ones do not. There have also been observations in the literature of damselflies leaving their waterside habitats, flying upwards till lost from view, and presumably being dispersed to far places by the stronger winds found at high altitudes. 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, on oil rigs far out in the Gulf of Mexico.

Biology
The adults 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. They are less dependent than dragonflies on warmth and can be observed hunting during cold spells. In tropical South America, helicopter damselflies (Pseudostigmatidae) feed on spiders, hovering near an orb web and plucking the spider, or its entangled prey, from the web. There are few pools and lakes in these habitats, and these damselflies breed in temporary water bodies in holes in trees, the rosettes of bromeliads and even the hollow stems of bamboos.

Ecology
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, and odontates have been used for bio-indication purposes regarding the quality of the ecosystem. Different species have different requirements for their larvae with regard to water depth, water movement and pH. The European common blue damselfly (Enallagma cyathigerum) for example can occur at high densities in acid waters where fish are absent, such as in bog pools. The scarce blue-tailed damselfly (Ischnura pumilio) in contrast requires base-rich habitats 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.

Damselflies, both nymphs and adults, are eaten by a range of predators including birds, fish, frogs, dragonflies, other damselflies, water spiders, water beetles, backswimmers and giant water bugs.

Damselflies have a variety of internal and external parasites. Particularly prevalent are the gregarine protozoans found 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. Bright red water mites Hydracarina are often seen on the outside of both nymphs and adults, and can move from one to the other at metamorphosis. They suck the body fluids and may actually kill young nymphs, but adults are relatively unaffected.

Behaviour
The blue-fronted dancer (Argia apicalis) of North America is fairly typical of damselfly behaviour. The adults hatched over an extended period and first appeared at the study site in mid-summer. Males visited water on about 40% of their adult days while females merely did so on 20%, both spending the rest of the time elsewhere, mostly in woods. The males arrived at water earlier in the day than females and start patrolling areas, stationing themselves about 2 metres (6 ft) from adjoining males. They darted at invaders and successfully drove them off in most instances although no physical contact occurred. Peak activity was around midday. The arrival of a female in the patrolled territory was greeted with immediate copulation lasting about 16 minutes, with no preliminary display or courtship. There followed a phase of 25 minutes while they explored the pond, an ovipositing phase with the pair in tandem lasting about 60 minutes followed by a further 20 minute egg-laying period by the female alone. Females avoided further sexual activity by flying rapidly away. Marked males had an apparent life expectancy of about 8 days and females 7, though the oldest marked male lived for 33 days. By late afternoon, all the damselflies had departed.

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. Flights of attrition are engaged in by the ebony 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. Complex courtship behaviours are exhibited by some species. The male river 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. Swift forktail (Ischnura erratica) males display to each other with their blue-tipped abdomens raised.

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. The desert 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.

Reproduction
Mating in damselflies, as in dragonflies, is a complex, precisely choreographed process involving both indirect insemination and delayed fertilisation. 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. 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"; the pair may also be described as being "in cop".

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 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, something made possible by delayed fertilisation and driven by sexual selection. 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. 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.

Nearly 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. For example, the red-eyed damselfly Erythromma najas lays eggs, in tandem, into leaves or stems of floating or sometimes emergent plants; in contrast, the scarce bluetail Ischnura pumilio oviposits alone, the female choosing mostly emergent grasses and rushes, and laying her eggs in their stems either above or just below the waterline. The willow emerald Chalcolestes 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.

Life cycle
Damselflies are hemimetabolous insects which have no pupal stage in their development. The female inserts the eggs by means of her ovipositor into slits made in water plants or other underwater substrates and the larvae, known as naiads or nymphs, are completely aquatic. An exception to this is the spreadwings. Late in the year these lay eggs above the waterline 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.

The nymphs are voracious predators and feed by means of a flat labium (a toothed mouthpart on the lower jaw) which pierces and seizes the daphnia, mosquito larvae, and other small aquatic organisms on which they 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. 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. The exoskeleton hardens 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.

Conservation
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 for hydroelectric schemes and the general degradation of wetlands and marshes. 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 have unintended consequences. In Hawaii, the introduction of the mosquitofish (Gambusia affinis) was effective in controlling mosquitoes but nearly exterminated the island's endemic damselflies.

In human culture
Damselfly is a 2005 short film directed by Ben O'Connor. Damselfly is also the title of a 2012 novel in the Faeble series by S. L. Naeole, and of a 1994 poem by August Kleinzahler, which contains the lines "And that blue there, cobalt / a moment, then iridescent, / fragile as a lady's pin / hovering above the nasturtium?" The poet John Engels published Damselfly, Trout, Heron in his 1983 collection Weather-Fear: New and Selected Poems.

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

Damselflies have formed subjects for personal jewellery such as brooches since at least 1880.