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39-766: For extinct groups, see text Odonata is an order of predatory flying insects that includes the dragonflies and damselflies (as well as the Epiophlebia damsel-dragonflies). The two major groups are distinguished with dragonflies (Anisoptera) usually being bulkier with large compound eyes together and wings spread up or out at rest, while damselflies (suborder Zygoptera ) are usually more slender with eyes placed apart and wings folded together along body at rest. Adult odonates can land and perch, but rarely walk. All odonates have aquatic larvae called naiads or nymphs , and all of them, larvae and adults, are carnivorous and are almost entirely insectivorous , although at

78-456: A cohors (plural cohortes ). Some of the plant families still retain the names of Linnaean "natural orders" or even the names of pre-Linnaean natural groups recognized by Linnaeus as orders in his natural classification (e.g. Palmae or Labiatae ). Such names are known as descriptive family names. In the field of zoology , the Linnaean orders were used more consistently. That is,

117-547: A copper -based protein that turns blue when oxygenated, instead of the iron -based hemoglobin in red blood cells found in vertebrates, giving hemolymph a blue-green color rather than the red color of vertebrate blood. When not oxygenated, hemolymph quickly loses its color and appears grey. The hemolymph of lower arthropods, including most insects , is not used for oxygen transport because these animals respirate through other means, such as tracheas, but it does contain nutrients such as proteins and sugars. Muscular movements by

156-460: 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). The smallest living dragonfly is Nannophya pygmaea (Anisoptera: Libellulidae) from east Asia, with

195-632: 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). 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

234-505: A capital letter. For some groups of organisms, their orders may follow consistent naming schemes . Orders of plants , fungi , and algae use the suffix -ales (e.g. Dictyotales ). Orders of birds and fishes use the Latin suffix -iformes meaning 'having the form of' (e.g. Passeriformes ), but orders of mammals and invertebrates are not so consistent (e.g. Artiodactyla , Actiniaria , Primates ). For some clades covered by

273-566: A distinct rank of biological classification having its own distinctive name (and not just called a higher genus ( genus summum )) was first introduced by the German botanist Augustus Quirinus Rivinus in his classification of plants that appeared in a series of treatises in the 1690s. Carl Linnaeus was the first to apply it consistently to the division of all three kingdoms of nature (then minerals , plants , and animals ) in his Systema Naturae (1735, 1st. Ed.). For plants, Linnaeus' orders in

312-418: A few insects living in low-oxygen environments are there hemoglobin-like molecules that bind oxygen and transport it to the tissues. Therefore, the demands placed upon the system are much lower. Some arthropods and most molluscs possess the copper -containing hemocyanin , however, for oxygen transport. In some species, hemolymph has other uses than just being a blood analogue. As the insect or arachnid grows,

351-875: A suborder along with Zygoptera and Anisozygoptera as well-understood and widely preferred terms. Cladogram of Epiprocta after Rehn et al. 2003: Zygoptera (damselflies) Epiophlebioptera † Isophlebioptera † Heterophlebioptera † Stenophlebioptera † Aeschnidioptera Anisoptera (dragonflies) Cladogram of Odonatoptera including Odonata by Deregnaucourt et al. 2023. † Meganeuridae † Polytaxineuridae † Permaeschnidae † Callimokaltaniidae † Hemizygopteridae † Ditaxineuridae † Iverya † Pirouteliidae † Kargalotypidae † Zygophlebiidae † Moltenophlebiidae † Permophlebiidae † Xamenophlebiidae † Triassologus † Sinotriadophlebiidae † Mitophlebiidae † Paurophlebiidae † Triadophlebiidae † Permagrionidae Order (biology) What does and does not belong to each order

390-470: 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

429-485: 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 ). World Odonata List considers Anisoptera as

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468-471: 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 . Tarsophlebiidae is a prehistoric family of Odonatoptera that can be considered either a basal lineage of Odonata or their immediate sister taxon . The phylogenetic tree of

507-419: Is determined by a taxonomist , as is whether a particular order should be recognized at all. Often there is no exact agreement, with different taxonomists each taking a different position. There are no hard rules that a taxonomist needs to follow in describing or recognizing an order. Some taxa are accepted almost universally, while others are recognized only rarely. The name of an order is usually written with

546-480: Is the major tissue type of the open circulatory system characteristic of arthropods (for example, arachnids , crustaceans and insects ). In addition, some non-arthropods such as mollusks possess a hemolymphatic circulatory system. Oxygen-transport systems were long thought unnecessary in insects, but ancestral and functional hemocyanin has been found in the hemolymph. Insect "blood" generally does not carry hemoglobin , although hemoglobin may be present in

585-520: The International Code of Zoological Nomenclature , several additional classifications are sometimes used, although not all of these are officially recognized. In their 1997 classification of mammals , McKenna and Bell used two extra levels between superorder and order: grandorder and mirorder . Michael Novacek (1986) inserted them at the same position. Michael Benton (2005) inserted them between superorder and magnorder instead. This position

624-469: 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. 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. The Odonata

663-769: The Systema Naturae and the Species Plantarum were strictly artificial, introduced to subdivide the artificial classes into more comprehensible smaller groups. When the word ordo was first consistently used for natural units of plants, in 19th-century works such as the Prodromus Systematis Naturalis Regni Vegetabilis of Augustin Pyramus de Candolle and the Genera Plantarum of Bentham & Hooker, it indicated taxa that are now given

702-399: The hemocytes , within the hemolymph. They play a role in the arthropod immune system . The immune system resides in the hemolymph. This open system might appear to be inefficient compared to the closed circulatory systems of the vertebrates , but the two systems have very different demands placed on them. In vertebrates, the circulatory system is responsible for transporting oxygen to all

741-502: The Anisoptera. An alternative term warriorfly has been proposed. 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

780-555: The abdomen, while dragonfly nymphs respire through an organ in their rectum . Members of the crown group Odonata first appeared during the Late Triassic , though members of their total group , Odonatoptera , first appeared in the Late Carboniferous , making them one of the earliest groups of winged insects. The fossils of odonates and their cousins, including Paleozoic "giant dragonflies" like Meganeuropsis permiana from

819-505: The animal during locomotion can facilitate hemolymph movement, but diverting flow from one area to another is limited. When the heart relaxes, hemolymph is drawn back toward the heart through open-ended pores called ostia. Note that the term " ostia " is not specific to insect circulation; it literally means "doors" or "openings", and must be understood in context. Hemolymph can contain nucleating agents that confer extracellular freezing protection. Such nucleating agents have been found in

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858-479: The arthropod stage of development. An example of this is the silkworm and its need for glycine in the production of silk. Proteins present in the hemolymph vary in quantity during the course of development. These proteins are classified by their functions: chroma proteins, protease inhibitors, storage, lipid transport, enzymes, the vitellogenins, and those involved in the immune responses of arthropods. Some hemolymphic proteins incorporate carbohydrates and lipids into

897-413: The body cavity. The hemocoel is divided into chambers called sinuses. Coordinated movements of the body muscles gradually bring the hemolymph back to the dorsal sinus surrounding the hearts. Between contractions, tiny valves in the wall of the hearts open and allow hemolymph to enter. Hemolymph fills all of the interior (the hemocoel ) of the animal's body and surrounds all cells. It contains hemocyanin ,

936-472: The energy consumption of flying. A pterostigma is also found among other insects, such as bees. 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. Damselfly nymphs breathe through external gills on

975-588: The hemolymph of insects of several orders, i.e., Coleoptera (beetles), Diptera (flies), and Hymenoptera . Hemolymph is composed of water , inorganic salts (mostly sodium , chlorine , potassium , magnesium , and calcium ), and organic compounds (mostly carbohydrates , proteins , and lipids ). The primary oxygen transporter molecule is hemocyanin . Arthropod hemolymph contains high levels of free amino acids. Most amino acids are present but their relative concentrations vary from species to species. Concentrations of amino acids also vary according to

1014-415: The hemolymph works something like a hydraulic system, enabling the insect or arachnid to expand segments before they are sclerotized . It can also be used hydraulically as a means of assisting movement, such as in arachnid locomotion . Some species of insect or arachnid are able to autohaemorrhage when they are attacked by predators. Queens of the ant genus Leptanilla are fed with hemolymph produced by

1053-490: The larval stage they will eat anything that they can overpower, including small fish , tadpoles , and even adult newts . The adults are superb aerial hunters and their legs are specialised for catching prey in flight. Odonata in its narrow sense forms a subgroup of the broader Odonatoptera , which contains other dragonfly-like insects. The scientific study of the Odonata is called odonatology. Johan Christian Fabricius coined

1092-438: The orders and suborders of odonates according to Bybee et al. 2021:   Zygoptera (damselflies) [REDACTED]   Anisozygoptera [REDACTED]   Anisoptera (dragonflies) [REDACTED] In some treatments, 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,

1131-698: The orders in the zoology part of the Systema Naturae refer to natural groups. Some of his ordinal names are still in use, e.g. Lepidoptera (moths and butterflies) and Diptera (flies, mosquitoes, midges, and gnats). In virology , the International Committee on Taxonomy of Viruses 's virus classification includes fifteen taxomomic ranks to be applied for viruses , viroids and satellite nucleic acids : realm , subrealm , kingdom , subkingdom, phylum , subphylum , class, subclass, order, suborder, family, subfamily , genus, subgenus , and species. There are currently fourteen viral orders, each ending in

1170-547: The precursor of the currently used International Code of Nomenclature for algae, fungi, and plants . In the first international Rules of botanical nomenclature from the International Botanical Congress of 1905, the word family ( familia ) was assigned to the rank indicated by the French famille , while order ( ordo ) was reserved for a higher rank, for what in the 19th century had often been named

1209-494: The rank of family (see ordo naturalis , ' natural order '). In French botanical publications, from Michel Adanson 's Familles naturelles des plantes (1763) and until the end of the 19th century, the word famille (plural: familles ) was used as a French equivalent for this Latin ordo . This equivalence was explicitly stated in the Alphonse Pyramus de Candolle 's Lois de la nomenclature botanique (1868),

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1248-645: The structure. Nitrogen metabolism end products are present in the hemolymph in low concentrations. These include ammonia , allantoin , uric acid , and urea . Arthropod hormones are present, most notably the juvenile hormone. Trehalose can be present and sometimes in great amounts along with glucose . These sugar levels are maintained by the control of hormones. Other carbohydrates can be present. These include inositol , sugar alcohol , hexosamines, mannitol , glycerol and those components that are precursors to chitin . Free lipids are present and are used as fuel for flight. There are free-floating cells,

1287-419: The suffix -virales . Hemolymph Hemolymph , or haemolymph , is a fluid, analogous to the blood in vertebrates , that circulates in the interior of the arthropod (invertebrate) body, remaining in direct contact with the animal's tissues. It is composed of a fluid plasma in which hemolymph cells called hemocytes are suspended. In addition to hemocytes, the plasma also contains many chemicals. It

1326-465: The term Odonata in 1793 from the Ancient Greek ὀδών odṓn ( Ionic form of ὀδούς odoús ) "tooth". One hypothesis is that it was because their maxillae are notably toothed. The word dragonfly usually denotes only Anisoptera , but is sometimes used to mean all Odonata. Odonata enthusiasts avoid ambiguity by using the term true dragonfly , or simply anisopteran , when they mean just

1365-494: 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 . The Anisoptera was long treated as a suborder, with a third suborder, the Anisozygoptera (ancient dragonflies). However, the combined suborder Epiprocta (in which Anisoptera

1404-413: The tissues and removing carbon dioxide from them. It is this requirement that establishes the level of performance demanded of the system. The efficiency of the vertebrate system is far greater than is needed for transporting nutrients, hormones, and so on, whereas in insects, exchange of oxygen and carbon dioxide occurs in the tracheal system . Hemolymph plays no part in the process in most insects. Only in

1443-407: The tracheal system instead and play some role in respiration. In the grasshopper , the closed portion of the system consists of tubular hearts and an aorta running along the dorsal side of the insect. The hearts pump hemolymph into the sinuses of the hemocoel where exchanges of materials take place. The volume of hemolymph needed for such a system is kept to a minimum by a reduction in the size of

1482-482: The underside of the head and include simple chewing mandibles in the adult. 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. In most families, there

1521-418: Was adopted by Systema Naturae 2000 and others. In botany , the ranks of subclass and suborder are secondary ranks pre-defined as respectively above and below the rank of order. Any number of further ranks can be used as long as they are clearly defined. The superorder rank is commonly used, with the ending -anae that was initiated by Armen Takhtajan 's publications from 1966 onwards. The order as

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