Odonata

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Odonata
Temporal range: Triassic–Recent
Aeshna juncea hovering.jpg
Aeshna juncea hovering over a pond.
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
(unranked): Dicondylia
Subclass: Pterygota
Division: Palaeoptera
Superorder: Odonatoptera
Order: Odonata
Suborders

Odonata is an order of flying insects that includes the dragonflies and damselflies.

Contents

The two common groups are distinguished with dragonflies, placed in the suborder Epiprocta, usually being larger, with eyes together and wings up or out at rest, while damselflies, suborder Zygoptera, are usually smaller with eyes placed apart and wings along body at rest.

All Odonata have aquatic larvae called naiads (nymphs), and all of them, larvae and adults, are carnivorous. The adults can land, but rarely walk. Their legs are specialised for catching prey. They are almost entirely insectivorous.

Etymology and terminology

Johan Christian Fabricius coined the term Odonata in 1793 from the Ancient Greek ὀδώνodṓn (Ionic form of ὀδούςodoús) 'tooth'. One hypothesis is that it was because their maxillae are notably toothed. [1]

The word dragonfly usually denotes only Anisoptera, but is sometimes used to mean all Odonata. [2] Odonata enthusiasts avoid ambiguity by using the term true dragonfly, [3] or simply anisopteran, [4] when they mean just the Anisoptera. An alternative term warriorfly has been proposed. [5]

External morphology

Size

The largest living odonate is the giant Central American helicopter damselfly Megaloprepus coerulatus (Zygoptera: Pseudostigmatidae) with a wing span of 191 mm (7.5 in). The heaviest living odonates are Tetracanthagyna plagiata (Anisoptera: Aeshnidae) with a wing span of 165 mm (6.5 in), and Petalura ingentissima (Anisoptera: Petaluridae) with a body length of 117 mm (4.6 in) (some sources 125 mm (4.9 in)) and wing span of 160 mm (6.3 in). The longest extant odonate is the Neotropical helicopter damselfly Mecistogaster linearis (Zygoptera: Pseudostigmatidae) with a body length of 135 mm (5.3 in). Sometimes the giant Hawaiian darner Anax strenuus (Anisoptera: Aeshnidae) is claimed to be the largest living odonate with an alleged wing span of 190 mm (7.5 in), but this seems to be a myth as only 152 mm (6.0 in) wing spans are scientifically documented.[ citation needed ]

The smallest living dragonfly is Nannophya pygmaea (Anisoptera: Libellulidae) from east Asia, with a body length of 15 mm (0.59 in) and a wing span of 20 mm (0.79 in). The smallest damselflies (and also the smallest odonates) are species of the genus Agriocnemis (Zygoptera: Coenagrionidae) with a wing span of only 17–18 mm (0.67–0.71 in).[ citation needed ]

Description

Male blue ringtail (Austrolestes annulosus), a damselfly (Zygoptera: Lestidae) Common blue damselfly02.jpg
Male blue ringtail (Austrolestes annulosus), a damselfly (Zygoptera: Lestidae)
Dragonfly (top) and damselfly (bottom) wing shape and venation OdonataWings.jpg
Dragonfly (top) and damselfly (bottom) wing shape and venation

These insects characteristically have large rounded heads covered mostly by well-developed, compound eyes, which provide good vision, legs that facilitate catching prey (other insects) in flight, two pairs of long, transparent wings that move independently, and elongated abdomens. They have three ocelli and short antennae. The mouthparts are on the underside of the head and include simple chewing mandibles in the adult. [6]

Flight in the Odonata is direct, with flight muscles attaching directly to the wings; rather than indirect, with flight muscles attaching to the thorax, as is found in the Neoptera. This allows active control of the amplitude, frequency, angle of attack, camber and twist of each of the four wings entirely independently. [7]

In most families, there is a structure on the leading edge near the tip of the wing called the pterostigma. This is a thickened, hemolymph–filled and often colorful area bounded by veins. The functions of the pterostigma are not fully known, but it most probably has an aerodynamic effect and may also have a visual function. More mass at the end of the wing may also reduce the energy needed to move the wings up and down. The right combination of wing stiffness and wing mass could reduce the energy consumption of flying. A pterostigma is also found among other insects, such as bees. [8]

The nymphs have stockier, shorter, bodies than the adults. In addition to lacking wings, their eyes are smaller, their antennae longer, and their heads are less mobile than in the adult. Their mouthparts are modified, with the labium being adapted into a unique prehensile organ called a labial mask for grasping prey. [9] Damselfly nymphs breathe through external gills on the abdomen, while dragonfly nymphs respire through an organ in their rectum. [6]

Evolution

Fossil history

The giant Upper Carboniferous dragonfly ancestor, Meganeura monyi, had a wingspan of about 680 mm (27 in), Meganeura monyi au Museum de Toulouse.jpg
The giant Upper Carboniferous dragonfly ancestor, Meganeura monyi, had a wingspan of about 680 mm (27 in),

Members of the Odonata first appeared during the Triassic, [11] though members of their total group, Odonatoptera, first appeared in the Late Carboniferous, making them one of the earliest groups of winged insects. [11] The fossils of odonates and their cousins, including Paleozoic "giant dragonflies" like Meganeuropsis permiana from the Permian of North America, reached wing spans of up to 71 cm (28 in) and a body length of 43 cm (17 in), making it the largest insect of all time. [12] This insect belonged to the order Meganisoptera, the griffinflies, related to odonates but not part of the modern order Odonata in the restricted sense. They have one of the most complete fossil records going back 319 million years. [13]

The Odonata is closely related to mayflies and several extinct orders in a group called the Palaeoptera, but this grouping might be paraphyletic. What they do share with mayflies is the nature of how the wings are articulated and held in rest.[ citation needed ]

Tarsophlebiidae is a prehistoric family of Odonatoptera that can be considered either a basal lineage of Odonata or their immediate sister taxon.[ citation needed ]

Phylogeny

The phylogenetic tree of the orders and suborders of odonates according to Bybee et al 2021: [14]

Odonata

Zygoptera (damselflies) Ischnura heterosticta02.jpg

Epiprocta

Anisozygoptera Epiophlebia superstes (cropped).jpg

Anisoptera (dragonflies) Emperor-Dragonfly-(8) (cropped).JPG

Taxonomy

In some treatments, [15] the Odonata are understood in an expanded sense, essentially synonymous with the superorder Odonatoptera, but not including the prehistoric Protodonata. In this approach, instead of Odonatoptera, the term Odonatoidea is used. The systematics of the "Palaeoptera" are by no means resolved; what can be said however is that regardless of whether they are called "Odonatoidea" or "Odonatoptera", the Odonata and their extinct relatives do form a clade. [16]

The Anisoptera was long treated as a suborder, with a third suborder, the Anisozygoptera (ancient dragonflies). However, the combined suborder Epiprocta (in which Anisoptera is an infraorder) was proposed when it was thought that the "Anisozygoptera" was paraphyletic, composed of mostly extinct offshoots of dragonfly evolution. The four living species placed in that group are (in this treatment) in the infraorder Epiophlebioptera, whereas the fossil taxa that were formerly there are now dispersed about the Odonatoptera (or Odonata sensu lato). [17] World Odonata List considers Anisoptera as a suborder along with Zygoptera and Anisozygoptera as well-understood and widely preferred terms. [18] [19] [20]

Ecology and life cycle

Ovipositing flight of two pairs of azure damselflies (Coenagrion puella) 070526 142326 Libellen cr.jpg
Ovipositing flight of two pairs of azure damselflies ( Coenagrion puella )

Odonates are aquatic or semi-aquatic as juveniles. Thus, adults are most often seen near bodies of water and are frequently described as aquatic insects. However, many species range far from water. They are carnivorous (or more specifically insectivorous) throughout their life, mostly feeding on smaller insects.[ citation needed ]

Male Odonata have complex genitalia, different from those found in other insects. These include grasping cerci at the tip of the abdomen for holding the female, and a secondary set of copulatory organs located between the second and third abdominal segment in which the spermatozoa are stored after being produced by the primary genitals— whose external opening is known as the genital pore, on the ninth abdominal segment. This process is called intra-male sperm translocation (ST). [21] [22] Because the male copulatory organ has evolved independently from that in other insects, it has been suggested the stem-group dragonflies had external sperm transfer. [23] To mate, the male claspers grasps the female by the thorax (Zygoptera) or head (Anisoptera) while the female bends her abdomen so that her own genitalia can be grasped by the copulatory organ holding the sperm. This is known as the "wheel" position. [6]

Eggs are laid in water or on vegetation near water or wet places, and hatch to produce pronymphs which live off the nutrients that were in the egg. They then develop into instars with approximately 9–14 molts that are (in most species) voracious predators on other aquatic organisms, including small fishes. The nymphs grow and molt, usually in dusk or dawn, into the flying teneral immature adults, whose color is not yet developed. These transform into reproductive adults.[ citation needed ]

Odonates can act as bioindicators of water quality in rivers because they rely on high quality water for proper development in early life. Since their diet consists entirely of insects, odonate density is directly proportional to the population of prey, and their abundance indicates the abundance of prey in the examined ecosystem. [24] Species richness of vascular plants has also been positively correlated with the species richness of dragonflies in a given habitat. This means that in a location such as a lake, if one finds a wide variety of odonates, then a similarly wide variety of plants should also be present. This correlation is not common to all bioindicators, as some may act as indicators for a different environmental factor, such as the pool frog acting as a bioindicator of water quality due to its high quantity of time spent in and around water. [25]

In addition, odonates are very sensitive to changes to average temperature. Many species have moved to higher elevations and latitudes as global temperature rises and habitats dry out. Changes to the life cycle have been recorded with increased development of the instar stages and smaller adult body size as the average temperature increases. As the territory of many species starts to overlap, the rate hybridization of species that normally do not come in contact is increasing. [13]  If global climate change continues many members of Odonata will start to disappear. Because odonates are such an old order and have such a complete fossil record they are an ideal species to study insect evolution and adaptation. For example, they are one of the first insects to develop flight and it is likely that this trait only evolved once in insects, looking at how flight works in odonates, the rest of flight can be mapped out. [13]

See also

Lists of Odonata species of Australia - Britain - Canada (dragonflies | damselflies) - Estonia - Finland - France - India - Ireland - Slovenia - South Africa - Sri Lanka - Taiwan

Related Research Articles

<span class="mw-page-title-main">Dragonfly</span> Predatory winged insects

A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of true dragonflies are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threatens dragonfly populations around the world. Adult dragonflies are characterized by a pair of large, multifaceted, compound eyes, two pairs of strong, transparent wings, sometimes with coloured patches, and an elongated body. Many dragonflies have brilliant iridescent or metallic colours produced by structural coloration, making them conspicuous in flight. An adult dragonfly's compound eyes have nearly 24,000 ommatidia each.

<span class="mw-page-title-main">Damselfly</span> Suborder of insects

Damselflies are flying insects of the suborder Zygoptera in the order Odonata. They are similar to dragonflies but are 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 the Jurassic, and are found on every continent except Antarctica.

<span class="mw-page-title-main">Nymph (biology)</span> Immature form of some invertebrates

In biology, a nymph is the immature form of some invertebrates, particularly insects, which undergoes gradual metamorphosis (hemimetabolism) before reaching its adult stage. Unlike a typical larva, a nymph's overall form already resembles that of the adult, except for a lack of wings. In addition, while a nymph moults, it never enters a pupal stage. Instead, the final moult results in an adult insect. Nymphs undergo multiple stages of development called instars.

<span class="mw-page-title-main">Green darner</span> Species of dragonfly

The green darner or common green darner, after its resemblance to a darning needle, is a species of dragonfly in the family Aeshnidae. One of the most common and abundant species throughout North America, it also ranges south to Panama. It is well known for its great migration distance from the northern United States south into Texas and Mexico. It also occurs in the Caribbean, Tahiti, and Asia from Japan to mainland China. It is the official insect for the state of Washington in the United States.

<span class="mw-page-title-main">Blue-tailed damselfly</span> Species of damselfly

The blue-tailed damselfly or common bluetail is a damselfly, belonging to the family Coenagrionidae.

<span class="mw-page-title-main">Calopterygidae</span> Family of damselflies

The Calopterygidae are a family of damselflies, in the suborder Zygoptera. They are commonly known as the broad-winged damselflies, demoiselles, or jewelwings. These rather large damselflies have wingspans of 50–80 mm, are often metallic-coloured, and can be differentiated from other damselflies by the broader connection between the wings and the body, as opposed to the abrupt narrowing seen in other damselfly families. The family contains some 150 species.

<span class="mw-page-title-main">Cordulegastridae</span> Family of dragonflies

The Cordulegastridae are a family of Odonata (dragonflies) from the suborder Anisoptera. They are commonly known as spiketails. Some vernacular names for the species of this family are biddie and flying adder. They have large, brown or black bodies with yellow markings, and narrow unpatterned wings. Their bright eyes touch at a single point, and they can be found along small, clear, woodland streams, flying slowly 30 to 70 cm above the water. When disturbed, however, they can fly very rapidly. They usually hunt high in forest vegetation, and prefer to capture prey resting on leaves or branches.

<i>Epiophlebia laidlawi</i> Species of dragonfly

Epiophlebia laidlawi, the Himalayan relict dragonfly, is one of four species of Epiprocta in the family Epiophlebiidae. They have at one time been classified as a suborder Anisozygoptera, considered as intermediate between the dragonflies and the damselflies, partly because the hind wings and fore wings are very similar in size and shape, and partly because the insect at rest holds them back over the body as damselflies do. These attributes now are known to be misleading however; the genus Epiophlebia shares a more recent ancestor with dragonflies and became separated from other Anisoptera in and around the uplifting Himalayas.

<span class="mw-page-title-main">Pseudostigmatidae</span> Family of damselflies

The Pseudostigmatidae are a family of tropical damselflies, known as helicopter damselflies, giant damselflies, or forest giants. The family includes the largest of all damselfly species. They specialize in preying on web-building spiders, and breed in phytotelmata, the small bodies of water held by plants such as bromeliads.

<i>Megaloprepus caerulatus</i> Species of damselfly

Megaloprepus caerulatus, also known as the blue-winged helicopter, is a forest giant damselfly of the family Coenagrionidae. Forest giant damselflies were previously recognized as their own family, Pseudostigmatidae. M. caerulatus is found in wet and moist forests in Central and South America. It has the greatest wingspan of any living damselfly or dragonfly, up to 19 centimeters (7.5 inches) in the largest males. Its large size and the markings on its wings make it a conspicuous species; a hovering Megaloprepus has been described as a "pulsating blue-and-white beacon".

<i>Thaumatoneura inopinata</i> Species of damselfly

Thaumatoneura inopinata is a species of damselfly, sometimes called the cascade damselfly or giant waterfall damsel, and the only member of the genus Thaumatoneura. It is unusual in flying among the falling water and spray from waterfalls in moist tropical or subtropical forests in Costa Rica.

<i>Epiophlebia</i> Genus of dragonflies

The genus Epiophlebia is the sole member of the family Epiophlebiidae, which is itself the sole living representative of the Epiproctan infraorder Epiophlebioptera, and it contains only three species. The first two species were historically placed in their own suborder Anisozygoptera, considered intermediate between dragonflies and damselflies, mainly because the hind wings are very similar in size and shape to the forewings and held back over the body at rest, as in damselflies. It has more recently been recognized that the genus Epiophlebia shares a more recent ancestor with dragonflies, and the group has accordingly been reclassified as an infraorder within the dragonflies. Very recently a third species, Epiophlebia sinensis, has been described from Heilongjiang province in northeast China, bridging Epiophlebia distribution gap between Nepal and Japan. A fourth species has been claimed from larval material from South China, but this is not universally accepted. Epiophlebia species are a freshwater indicator of a river ecosystem health. A study that has been conducted on the head anatomy of Epiophlebia has verified the presence of 41 muscles in the head of the larva. Like in true dragonflies (Anisoptera) the aquatic nymphs breathe through a rectal chamber, but jet propulsion has yet to be documented. Epiophlebia species are a representative of a dragonfly fauna which originated during the Jurassic period on the rising continent of Eurasia.

<i>Epiophlebia superstes</i> Species of dragonfly

Epiophlebia superstes, the Japanese relict dragonfly, is one of the four species of the genus Epiophlebia, belonging to the family Epiophlebiidae, which is itself the sole living representative of the epiproctan infraorder Epiophlebioptera.

<span class="mw-page-title-main">Stenophlebiidae</span> Extinct family of insects

The Stenophlebiidae is an extinct family of medium-sized to large fossil odonates from the Upper Jurassic and Cretaceous period that belongs to the damsel-dragonfly grade ("anisozygopteres") within the stem group of Anisoptera. They are characterized by their long and slender wings, and the transverse shape of the discoidal triangles in their wing venation.

<span class="mw-page-title-main">Tarsophlebiidae</span> Extinct family of flying insects

The Tarsophlebiidae is an extinct family of medium-sized fossil odonates from the Upper Jurassic and Lower Cretaceous period of Eurasia. They are either the most basal member of the damsel-dragonfly grade ("anisozygopteres") within the stem group of Anisoptera, or the sister group of all Recent odonates. They are characterized by the basally open discoidal cell in both pairs of wings, very long legs, paddle-shaped male cerci, and a hypertrophied ovipositor in females.

<i>Bradinopyga geminata</i> Species of dragonfly

Bradinopyga geminata is a species of dragonfly in the family Libellulidae known commonly as the granite ghost. It is native to India, Sri Lanka and Thailand, where it is a common and widespread species.

Odonata are insects with an incomplete metamorphosis (hemimetabolous). The aquatic larva or nymph hatches from an egg, and develops through eight to seventeen instars before leaving the water and emerging as the winged adult or imago.

References

  1. ↑ Mickel, Clarence E. (1934). "The significance of the dragonfly name "Odonata"". Annals of the Entomological Society of America. 27 (3): 411–414. doi:10.1093/aesa/27.3.411.
  2. ↑ "odonate". Merriam-Webster.com Dictionary .
  3. ↑ Field guide to lower aquarium animals. Cranbrook Institute of Science. 1939.
  4. ↑ Orr, A. G. (2005). Dragonflies of Peninsular Malaysia and Singapore. Natural History Publications (Borneo). ISBN   978-983-812-103-3.
  5. ↑ Philip S. Corbet; Stephen J. Brook (2008). Dragonflies. London: Collins. ISBN   978-0-00-715169-1.
  6. 1 2 3 Hoell, H.V., Doyen, J.T. & Purcell, A.H. (1998). Introduction to Insect Biology and Diversity (2nd ed.). Oxford University Press. pp. 355–358. ISBN   978-0-19-510033-4.{{cite book}}: CS1 maint: multiple names: authors list (link)
  7. ↑ Richard J. Bomphrey, Toshiyuki Nakata, Per Henningsson, Huai-Ti Lin (2016) Flight of the dragonflies and damselflies, Phil. Trans. R. Soc. B 371 20150389; DOI: 10.1098/rstb.2015.0389.
  8. ↑ Norberg, R. Åke (1972). "The pterostigma of insect wings an inertial regulator of wing pitch". Journal of Comparative Physiology A. 81 (1): 9–22. doi:10.1007/BF00693547. S2CID   23441098.
  9. ↑ The toolkit of a hunter – functional morphology of larval mouthparts in a dragonfly
  10. ↑ The Biology of Dragonflies. Cambridge University Press Archive. 2018. p. 324. GGKEY:0Z7A1R071DD. No Dragonfly at present existing can compare with the immense Meganeura monyi of the Upper Carboniferous, whose expanse of wing was somewhere about twenty-seven inches.
  11. 1 2 Kohli, Manpreet Kaur; Ware, Jessica L.; Bechly, Günter (2016). "How to date a dragonfly: Fossil calibrations for odonates". Palaeontologia Electronica. 19 (1). doi: 10.26879/576 .
  12. ↑ Mitchell, F.L.; Lasswell, J. (2005). A dazzle of dragonflies. Texas A&M University Press. p. 47.
  13. 1 2 3 Bybee, Seth; Córdoba-Aguilar, Alex; Duryea, M. Catherine; et al. (December 2016). "Odonata (dragonflies and damselflies) as a bridge between ecology and evolutionary genomics". Frontiers in Zoology. 13 (1): 46. doi: 10.1186/s12983-016-0176-7 . ISSN   1742-9994. PMC   5057408 . PMID   27766110.
  14. ↑ Bybee, Seth M.; Kalkman, Vincent J.; Erickson, Robert J.; et al. (2021). "Phylogeny and classification of Odonata using targeted genomics". Molecular Phylogenetics and Evolution. Elsevier BV. 160: 107115. doi: 10.1016/j.ympev.2021.107115 . hdl: 11093/2768 . ISSN   1055-7903. PMID   33609713.
  15. ↑ E.g. Trueman & Rowe (2008)
  16. ↑ Trueman [2008]
  17. ↑ Lohmann (1996), Rehn (2003)
  18. ↑ Paulson, D.; Schorr, M.; Abbott, J.; Bota-Sierra, C.; Deliry, C.; Dijkstra, K.-D.; Lozano, F. (2023). "World Odonata List". OdonataCentral, University of Alabama . Retrieved 14 Mar 2023.
  19. ↑ Dijkstra, K-D. B., G. Bechly, S. M. Bybee, R. A. Dow, H. J. Dumont, G. Fleck, R. W. Garrison, M. Hämäläinen, V. J. Kalkman, H. Karube, M. L. May, A. G. Orr, D. R. Paulson, A. C. Rehn, G. Theischinger, J. W. H. Trueman, J. van Tol, N. von Ellenrieder, & J. Ware. 2013. The classification and diversity of dragonflies and damselflies (Odonata). Zootaxa 3703(1): 36-45.
  20. ↑ 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: 107115. doi: 10.1016/j.ympev.2021.107115 . hdl: 11093/2768 . ISSN   1055-7903. PMID   33609713.
  21. ↑ Studies on the secondary male Genitalia of the type species of some dragonflies (Odonata: Anisoptera: Libellulidae)
  22. ↑ Evolution and diversity of intra-male sperm translocation in Odonata: a unique behaviour in animals [2019]
  23. ↑ The Phylogeny of Hexapoda (Arthropoda) and the Evolution of Megadiversity
  24. ↑ Golfieri, Bruno; Hardersen, Sönke; Maiolini, Bruno; Surian, Nicola (2016). "Odonates as indicators of the ecological integrity of the river corridor: Development and application of the Odonate River Index (ORI) in northern Italy". Ecological Indicators. 61: 234–247. doi:10.1016/j.ecolind.2015.09.022. ISSN   1470-160X.
  25. ↑ Sahlén, Göran; Ekestubbe, Katarina (16 May 2000). "Identification of dragonflies (Odonata) as indicators of general species richness in boreal forest lakes". Biodiversity and Conservation . 10 (5): 673–690. doi:10.1023/A:1016681524097. S2CID   40735036.
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