Damselfly
Damselfly Temporal range:
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Male beautiful demoiselle ( Calopteryx virgo )
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Female bluetail damselfly (Ischnura heterosticta) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Odonata |
Suborder: | Zygoptera Selys, 1854[1]
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Superfamilies | |
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Damselflies are
All damselflies are
Some species of damselfly have elaborate
Artificial fishing flies that mimic damselfly nymphs are used in wet-fly fishing. Damselflies are sometimes represented in personal jewellery such as brooches.
Classification
The Zygoptera are an ancient group, with the earliest fossils dating to the Late Jurassic, around 153 million years ago.[2] Well-preserved Eocene damselfly larvae and exuviae are known from fossils preserved in amber in the Baltic region.[3]
Molecular analysis in 2021 confirms that most of the traditional families are
Zygoptera |
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General description
The general body plan of a damselfly is similar to that of a dragonfly. The
In general, damselflies are smaller than dragonflies, the smallest being members of the genus
The first
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 with
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 bears cerci and in males, its underside bears a pair of paraprocts.[5]
Damselflies (except spreadwings,
Distribution and diversity
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. 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.[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, on oil rigs far out in the Gulf of Mexico.[5]
The distribution and diversity of damselfly species in the
Family | Oriental
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Neotropical
|
Australasian | Afrotropical
|
Palaearctic
|
Nearctic
|
Pacific | World |
---|---|---|---|---|---|---|---|---|
Hemiphlebiidae | 1 | 1 | ||||||
Lestidae | 40 | 42 | 29 | 17 | 18 | 18 | 3 | 151 |
Perilestidae | 19 | 19 | ||||||
Synlestidae | 18 | 1 | 8 | 11 | 6 | 39 | ||
Platystictidae | 136 | 43 | 44 | 1 | 1 | 224 | ||
Amphipterygidae |
4 | 4 | ||||||
Argiolestidae | 10 | 73 | 19 | 6 | 108 | |||
Calopterygidae | 66 | 68 | 5 | 20 | 37 | 8 | 185 | |
Chlorocyphidae | 86 | 17 | 42 | 3 | 144 | |||
Devadattidae |
6 | 6 | ||||||
Dicteriadidae | 2 | 2 | ||||||
Euphaeidae | 65 | 1 | 11 | 68 | ||||
Heteragrionidae | 51 | 51 | ||||||
Hypolestidae | 2 | 2 | ||||||
Lestoideidae | 9 | 9 | ||||||
Megapodagrionidae | 29 | 29 | ||||||
Pentaphlebiidae | 2 | 2 | ||||||
Philogangidae |
4 | 2 | 4 | |||||
Philogeniidae | 39 | 39 | ||||||
Philosinidae | 12 | 12 | ||||||
Polythoridae | 59 | 59 | ||||||
Pseudolestidae |
1 | 1 | ||||||
Rimanellidae |
1 | 1 | ||||||
Thaumatoneuridae | 5 | 5 | ||||||
Incertae sedis | 25 | 11 | 19 | 9 | 61 | |||
Coenagrionidae | 193 | 554 | 152 | 202 | 96 | 103 | 91 | 1266 |
Isostictidae | 41 | 46 | ||||||
Platycnemididae | 199 | 122 | 70 | 22 | 404 |
Overall, there are about 2942 extant species of damselflies placed in 309 genera.[21]
Biology
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).
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 be
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. Odonates have been used for
In the tropics, the helicopter damselfly Mecistogaster modesta (
Damselflies, both nymphs and adults, are eaten by a range of
Damselflies have a variety of internal and external parasites. Particularly prevalent are the
Behaviour
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. Calopteryx males 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 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.[35] Platycypha males 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 in Platycnemis phasmovolans and a few other Platycnemididae, including the extinct Yijenplatycnemis huangi.[36] Rhinocypha bobs 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 in Libellago semiopaca despite 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 in Calopteryx sp., Platycypha fitzsimonsi,[39] and Platycypha caligata. Male members of the family Protoneuridae with 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]
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 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.[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] 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.[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
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 of sexual cannibalism exist where females (of Ischnura graellsii) eat males while in copula.[53]
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 of sperm 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 by sexual 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]
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Hetaerina americanamating: sperm removal
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.
Many damselflies are able to produce more than one brood per year (voltinism); this is negatively correlated with latitude, becoming more common towards the equator, except in the Lestidae.[61]
Life cycle
Damselflies are hemimetabolous insects 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 underwater substrates and the larvae, known as naiads or nymphs, are almost all completely aquatic.[5] Exceptions include the Hawaiian Megalagrion oahuense and an unidentified Megapodagrionid from New Caledonia,[63] which are terrestrial in their early stages.[55] The spreadwings lay 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]
The nymphs are voracious predators and feed by means of a flat labium (a toothed mouthpart on the lower jaw) that forms the so-called mask; it is rapidly extended to seize and pierce the Daphnia (water fleas), mosquito larvae, 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. 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.[5]
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.[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 have unintended consequences.[64] In Hawaii, the introduction of the mosquitofish (Gambusia affinis) was effective in controlling mosquitoes but nearly exterminated the island's endemic damselflies.[65] The ancient greenling Hemiphlebia mirabilis has been an important flagship species for conservation action in preserving its habitat in Australia.[66]
In culture
Damselflies have formed subjects for personal jewellery such as brooches since at least 1880.[68]
Damselfly is a 2005 short film directed by Ben O'Connor.[69]
Damselfly is the title of a 2012 novel in the Faeble series 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 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?"[72] The poet John Engels published “Damselfly, Trout, Heron” in his 1983 collection Weather-Fear: New and Selected Poems.[73]
See also
- List of damselflies of the world
- List of odonates of Sri Lanka
- Protozygoptera, a fossil group that somewhat resembled the damselflies
Citations
- .
- doi:10.26879/576.
- ^ 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 from the original (PDF) on 2 April 2015. Retrieved 16 March 2015.
- PMID 33609713.
- ^ ISBN 978-1-4008-3966-7.
- ^ Dijkstra 2006, pp. 20, 104.
- .
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- PMID 16945555.
- PMID 12189786.
- ISBN 978-0-643-10249-1.
- ^ Dijkstra 2006, pp. 23, 65–67.
- ^ Dragonflies and Damselflies of Texas and the South-Central United States: Texas, Louisiana, Arkansas, Oklahoma, and New Mexico - Princeton University
- ^ Physiology of Insects
- ^ Borror, Donald J.; Triplehorn, Charles A.; Triplehorn, Norman F. Study of Insects (6 ed.). Philadelphia: Saunders College Publishing. pp. 187–201.
- ISBN 978-87-88757-07-1.
- doi:10.1086/278632.
- S2CID 87698039.
- ^ "Introduction to the Odonata". UCMP Berkeley. Archived from the original on 27 August 2017. Retrieved 13 March 2015.
- ^ ISBN 978-0-12-385026-3.
- ^ ISBN 978-0-88864-419-0.
- ^ Dragonflies and Damselflies of Georgia and the Southeast
- PMID 24486162.
- ^ ISBN 978-1-4020-8176-7.
- S2CID 82370392.
- ^ 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 from the original (PDF) on 2007-06-28.
- ^ ISBN 978-1-84911-093-8.
- ^ Dijkstra 2006, p. 102.
- .
- S2CID 53603745.
- ^ 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.
- ]
- ISBN 978-1-4020-2703-1.
- ^ Paulson 2009, p. 42.
- PMID 28317876.
- ^ ISBN 978-0-643-10249-1.
- ^ a b Orr, A. G. (1996). "Territorial and courtship displays in Bornean Chlorocyphidae (Zygoptera)" (PDF). Odonatologica. 25 (2): 119–141.
- ^ Robertson, H. M. (1982). "Courtship displays and mating behaviour of three species of Chlorocyphida (Zygoptera)" (PDF). Odonatologica. 11 (1): 53–58.
- ^ Paulson 2009, p. 108.
- JSTOR 1934877.
- ^ Paulson 2009, p. 44.
- ISSN 1432-0762.
- ^ Paulson 2009, p. 185.
- S2CID 85986224.
- ^ a b c Dijkstra 2006, pp. 8–9.
- ^ .
- ^ a b Cordero-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 from the original (PDF) on 2015-02-25. Retrieved 2015-03-16.
- ^ a b Trueman & Rowe 2009, p. Life Cycle and Behavior.
- ^ a b Berger 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'.
- PMID 5834678.
- ^ 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.
- doi:10.1127/entom.gen/17/1992/17. Archived from the original(PDF) on 2013-06-13. Retrieved 2017-04-28.
- PMID 19513091.
- ^ ISBN 978-0-674-04619-1.
- ^ 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.
- ISBN 978-0-8117-2731-0.
- ISBN 978-1-4008-5186-7.
- ^ Dijkstra 2006, p. 84.
- S2CID 83520585.
- S2CID 55299239.
- ISBN 978-1-84593-542-9.
- .
- ^ a b Moore, N.W. (1997). "Dragonflies: status survey and conservation action plan" (PDF). International Union for Conservation of Nature. Archived from the original (PDF) on 12 April 2016. Retrieved 17 March 2015.
- ^ Gagné, W.C. (1981). "Status of Hawaii endangered species: insects and land snails". ʻElepaio. 42: 31–36.
- ^ 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.
- ^ Wada, Wes (2012). "Fishing Tips for the Juicebug Damsel Nymph". Fly foundry. Archived from the original on 19 November 2016. Retrieved 13 March 2015.
- ^ "Antique "Damselfly" Brooch in Silver-topped Gold with Ruby Eyes". Macklowe Gallery. Retrieved 13 March 2015.[dead link]
- ^ "Ben O'Connor". British Council. Retrieved 13 March 2015.
- ^ Naeole, S. L. (2012). Damselfly. Crystal Quill.
- ^ "Damselfly". Marly Rusoff Literary Agency. Retrieved 21 December 2020.
- ^ Kleinzahler, August (August 1994). "The Damselfly". Poetry Magazine. Retrieved 13 March 2015.
- ^ Engels, John (1983). Damselfly, Trout, Heron. University of Georgia Press. Retrieved 13 March 2015.
General and cited sources
- Berger, Cynthia (2004). Dragonflies. Stackpole Books. p. 2. ISBN 978-0-8117-2971-0.
- Dijkstra, Klaas-Douwe B. (2006). Field Guide to the Dragonflies of Britain and Europe. British Wildlife Publishing. ISBN 978-0-9531399-4-1.
- Paulson, Dennis (2009). Dragonflies and Damselflies of the West. Princeton University Press. ISBN 978-1-4008-3294-1.
- Trueman, John W. H.; Rowe, Richard J. (2009). "Odonata". Tree of Life. Archived from the original on 21 November 2010. Retrieved 25 February 2015.
External links
- Tree of Life: Odonata
- Dragonflies and damselflies on the UF / IFAS Featured Creatures Web site
- Minnesota Dragonfly Society: Biology and Ecology