Endopterygota

Endopterygota (from Ancient Greek endon “inner” + pterón, “wing” + New Latin -ota “having”), also known as Holometabola, is a superorder of insects within the infraclass Neoptera that go through distinctive larval, pupal, and adult stages. They undergo a radical metamorphosis, with the larval and adult stages differing considerably in their structure and behaviour. This is called holometabolism, or complete metamorphism.

The Endopterygota are among the most diverse insect superorders, with over 1 million living species divided between 11 orders, containing insects such as butterflies, flies, fleas, bees, ants, and beetles.[1]

They are distinguished from the Exopterygota (or Hemipterodea) by the way in which their wings develop. Endopterygota (meaning literally "internal winged forms") develop wings inside the body and undergo an elaborate metamorphosis involving a pupal stage. Exopterygota ("external winged forms") develop wings on the outside their bodies and do not go through a pupal stage. The latter trait is plesiomorphic, however, and not exclusively found in the exopterygotes, but also in groups such as Odonata (dragonflies and damselflies), which are not Neoptera, but more basal among insects.

The earliest endopterygote fossils date from the Carboniferous.[2]

Endopterygota
Temporal range: Pennsylvanian–Recent
Panorpa communis V
Panorpa communis, a scorpionfly (order Mecoptera)
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
(unranked): Eumetabola
(unranked): Endopterygota
Sharp, 1898
Orders

Systematics

Ontwikkelstadia wespenpoppen
Holometabolism stages in Hymenoptera.

ITIS considers any subdivision of the Neoptera beyond the orders invalid, but this is almost universally rejected.

The Endopterygota are sometimes divided into three assemblages: Neuropterida (Neuroptera, Megaloptera, Raphidioptera, Strepsiptera and Coleoptera), Hymenopteroida (Hymenoptera), and Panorpida (Siphonaptera, Diptera, Trichoptera, Lepidoptera and Mecoptera). The hymenopterans, with their highly developed social systems, were believed to have constituted the most advanced insects, despite their rather "primitive" anatomy compared to flies and beetles, for example.

More recently, this has increasingly been rejected and DNA sequence data seem to verify that the hymenopterans are indeed among the most basal endopterygotes, whereas flies and fleas are often considered the most radically advanced insects. This calls the previous subdivision into question, and consequently several new taxa have been proposed, splitting up the Endopterygota. While some groups (such as the "sucking-stinging" fly-flea assemblage or the caddisfly-butterfly group) seem indeed to be good clades, it is not likely that the relationships of the endopterygotes, or the neopteran insects in general, will be resolved in detail soon.

Endopterygota sensu stricto
Neuropterida

Megaloptera (alderflies and allies)

Raphidioptera (snakeflies)

Neuroptera (lacewings and allies)

Coleopterida

Coleoptera (beetles)

Strepsiptera (twisted-wing parasites)

Hymenoptera (sawflies, wasps, ants, bees)

Panorpida

Diptera (true flies)

Mecoptera (scorpionflies)

Siphonaptera (fleas)

Trichoptera (caddisflies)

Lepidoptera (butterflies and moths)

Incertae sedis

See also

References

  1. ^ Rolf G. Beutel; Hans Pohl (2006). "Endopterygote systematics – where do we stand and what is the goal (Hexapoda, Arthropoda)?". Systematic Entomology. 31 (2): 202–219. doi:10.1111/j.1365-3113.2006.00341.x.
  2. ^ A. Nel; P. Roques; P. Nel; J. Prokop; J. S. Steyer (2007). "The earliest holometabolous insect from the Carboniferous: a "crucial" innovation with delayed success (Insecta Protomeropina Protomeropidae)". Annales de la Société Entomologique de France. 43 (3): 349–355.
Chrysopoidea

Chrysopoidea is a lacewing superfamily in the suborder Hemerobiiformia.

Coniopterygoidea

Coniopterygoidea is a lacewing superfamily in the suborder Hemerobiiformia. In some classifications, Coniopterygoidea is an expansion used to signify that the spongillaflies (Sisyridae) and the dustywings (Coniopterygidae) are each other's closest relatives.

Exopterygota

The Exopterygota, also known as Hemipterodea, are a superorder of insects of the subclass Pterygota in the infraclass Neoptera, in which the young resemble adults but have externally developing wings. They undergo a modest change between immature and adult, without going through a pupal stage. The nymphs develop gradually into adults through a process of moulting.

The Exopterygota are a highly diverse insect superorder, with at least 130,000 living species divided between 15 orders. They include cockroaches, termites, grasshoppers, thrips, lice and stick insects, among many other types of insects.

They are distinguished from the Endopterygota (or Holometabola) by the way in which their wings develop. Endopterygota (meaning literally "internal winged forms") develop wings inside the body and undergo an elaborate metamorphosis involving a pupal stage. Exopterygota ("external winged forms") develop wings on the outside of their bodies without going through a true pupal stage, though a few have something resembling a pupa (e.g., Aleyrodidae).

Ephemeroptera (mayflies) and Odonata (dragonflies and damselflies) also have gradual wing development, this being a plesiomorphic trait. These two orders belong to the infraclass Palaeoptera however, which is not included in Neoptera. As opposed to Neoptera, they cannot fold their wings over their back in the horizontal plane, only vertically (as damselflies do) if at all.

Flea

Flea, the common name for the order Siphonaptera, includes 2,500 species of small flightless insects that survive as external parasites of mammals and birds. Fleas live by consuming blood or hematophagy, from their hosts. Adult fleas grow to about 3 mm or 0.12 in long, are usually brown, and have bodies that are "flattened" sideways or narrow, enabling them to move through their host's fur or feathers. They lack wings, but have strong claws preventing them from being dislodged, mouthparts adapted for piercing skin and sucking blood, and hind legs extremely well adapted for jumping. They are able to leap a distance of some 50 times their body length, a feat second only to jumps made by another group of insects, the superfamily of froghoppers. Fleas' larvae are worm-like with no limbs; they have chewing mouthparts and feed on organic debris left on their host's skin.

The Siphonaptera are most closely related to the snow scorpionflies, or snow fleas in the UK, formally the Boreidae, placing them within the Endopterygote insect order Mecoptera. Fleas arose in the early Cretaceous, most likely as ectoparasites of mammals, before moving on to other groups, including birds. Each species of flea is more or less a specialist with respect to its host animal species: many species never breed on any other host, though some are less selective. Some families of fleas are exclusive to a single host group; for example, the Malacopsyllidae are found only on armadillos, the Ischnopsyllidae only on bats, and the Chimaeropsyllidae only on elephant shrews.

The oriental rat flea, Xenopsylla cheopis, is a vector of Yersinia pestis, the bacterium which causes bubonic plague. The disease was spread to humans by rodents such as the black rat, which were bitten by infected fleas. Major outbreaks included the Plague of Justinian, c. 540 and the Black Death, c. 1350, both of which killed a sizeable fraction of the world's population.

Fleas appear in human culture in such diverse forms as flea circuses, poems like John Donne's erotic The Flea, works of music such as by Modest Mussorgsky, and a film by Charlie Chaplin.

Glossata

Glossata (Fabricius, 1775) is the suborder of the insect order Lepidoptera that contains most lepidopteran species and includes all the superfamilies of moths and butterflies that have a coilable proboscis. (See also the suborders Zeugloptera, Aglossata, and Heterobathmiina).

Heteroneura

Heteroneura is a natural group (or clade) in the insect order Lepidoptera that comprises over 99% of all butterflies and moths. This is the sister group of the infraorder Exoporia (swift moths and their relatives), and is characterised by wing venation which is not similar or homoneurous in both pairs of wings.

Holometabolism

Holometabolism, also called complete metamorphosis, is a form of insect development which includes four life stages: egg, larva, pupa, and imago or adult. Holometabolism is a synapomorphic trait of all insects in the superorder Endopterygota. Immature stages of holometabolous insects are very different from the mature stage. In some species the holometabolous life cycle prevents larvae from competing with adults because they inhabit different ecological niches. The morphology and behavior of each stage are adapted for different activities. For example, larval traits maximize feeding, growth, and development, while adult traits enable dispersal, mating, and egg laying. Some species of holometabolous insects protect and feed their offspring. Other insect developmental strategies include ametabolism and hemimetabolism.

Hymenoptera

Hymenoptera is a large order of insects, comprising the sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones.Females typically have a special ovipositor for inserting eggs into hosts or places that are otherwise inaccessible. The ovipositor is often modified into a stinger. The young develop through holometabolism (complete metamorphosis)—that is, they have a worm-like larval stage and an inactive pupal stage before they mature.

Mantispoidea

The Mantispoidea are a superfamily of lacewing insects in the suborder Hemerobiiformia. It has two families.

Mecoptera

Mecoptera (from the Greek: mecos = "long", ptera = "wings") are an order of insects in the superorder Endopterygota with about six hundred species in nine families worldwide. Mecopterans are sometimes called scorpionflies after their largest family, Panorpidae, in which the males have enlarged genitals that look similar to the stingers of scorpions, and long beaklike rostra. The Bittacidae, or hangingflies, are another prominent family and are known for their elaborate mating rituals, in which females choose mates based on the quality of gift prey offered to them by the males. A smaller group is the snow scorpionflies, family Boreidae, adults of which are sometimes seen walking on snowfields. In contrast, the majority of species in the order inhabit moist environments in tropical locations.

The Mecoptera are closely related to the Siphonaptera (fleas), and a little more distantly to the Diptera (true flies). They are somewhat fly-like in appearance, being small to medium-sized insects with long slender bodies and narrow membranous wings. Most breed in moist environments such as leaf litter or moss, and the eggs may not hatch until the wet season arrives. The larvae are caterpillar-like and mostly feed on vegetable matter, and the non-feeding pupae may pass through a diapause until weather conditions are favorable.

Early Mecoptera may have played an important role in pollinating extinct species of gymnosperms before the evolution of other insect pollinators such as bees. Adults of modern species are overwhelmingly predators or consumers of dead organisms; they are the first insects to arrive at a cadaver, making them useful in forensic entomology.

Megaloptera

Megaloptera is an order of insects. It contains the alderflies, dobsonflies and fishflies, and there are about 300 known species.

The order's name comes from Ancient Greek, from mega- (μέγα-) "large" + pteryx (πτέρυξ) "wing", in reference to the large, clumsy wings of these insects. Megaloptera are relatively unknown insects across much of their range, due to the adults' short lives, the aquatic larvae's often-high tolerance of pollution (so they are not often encountered by swimmers etc.), and the generally crepuscular or nocturnal habits. However, in the Americas the dobsonflies are rather well-known, as their males have tusk-like mandibles. These, while formidable in appearance, are relatively harmless to humans and other animals; much like a peacock's feathers, they serve mainly to impress females. However, the mandibles are also used to hold females during mating, and some male dobsonflies spar with each other in courtship displays, trying to flip each other over with their long mandibles. Dobsonfly larvae, commonly called hellgrammites, are often used for angling bait in North America.

The Megaloptera were formerly considered part of a group then called Neuroptera, together with lacewings and snakeflies, but these are now generally considered to be separate orders, with Neuroptera referring to the lacewings and relatives (which were formerly called Planipennia). The former Neuroptera, particularly the lacewing group, are nonetheless very closely related to each other, and the new name for this group is Neuropterida. This is either placed at superorder rank, with the Endopterygota—of which they are part—becoming an unranked clade above it, or the Endopterygota are maintained as a superorder, with an unranked Neuropterida being a part of them. Within the endopterygotes, the closest living relatives of the neuropteridan clade are the beetles.

The Asian dobsonfly Acanthacorydalis fruhstorferi can have a wingspan of up to 21.6 cm (8.5 in), making it the largest aquatic insect in the world by this measurement.

Metapterygota

Metapterygota is a clade of winged insects containing order Odonata and Infraclass Neoptera.

Neuropterida

The Neuropterida are a clade, sometimes placed at superorder level, of holometabolous insects with over 5,700 described species, containing the orders Neuroptera (lacewings, antlions), Megaloptera (alderflies, dobsonflies), and Raphidioptera (snakeflies).

Historically, they were known as Neuroptera, but this name nowadays refers to lacewings and their relatives (e.g. antlions) only, which formerly were known as Planipennia. Part of the Endopterygota and related to beetles, they can be considered an unranked taxon. Arguably, the Endopterygota might be considered an unranked clade instead, and be split up in numerous superorders to signify the close relationships of certain endopterygote groups.The Mecoptera (scorpionflies) were formerly included here too by some authors, but they actually belong to the Mecopteroidea (or Antliophora), the endopterygote clade containing also true flies and fleas.

Neuropterida are fairly primitive-looking insects, with large wings but weak wing muscles, giving them a clumsy flight. Most are active at dusk or in the night as adults, and the larvae of many are aquatic, living in rivers. At least the larvae, but in many cases the adults too, are predators of small arthropods. Adult neuropteridans range in size from that of a midge to that of a large dragonfly (15 cm wingspan); the largest species tend to resemble drab, clumsily flying damselflies.

In addition to the three living orders, there is an entirely extinct family of Neuropterida, the monotypic Rafaelidae. These are of an indeterminate but probably rather basal position; thus the single genus Rafaelia from the Early Cretaceous Santana Formation's Crato Member in Brazil might for the time being be better placed in the Neuropterida directly, without assigning it to an order, until relatives are found and/or its systematic position gets resolved better. The extinct order Glosselytrodea may also be a member or close relative, though classification is unclear.

Panorpida

Panorpida or Mecopterida is a proposed superorder of Endopterygota. The conjectured monophyly of the Panorpida is historically based on morphological evidence, namely the reduction or loss of the ovipositor and several internal characteristics, including a muscle connecting a pleuron and the first axillary sclerite at the base of the wing, various features of the larval maxilla and labium, and basal fusion of CuP and A1 veins in the hind wings. The monophyly of the Panorpida is also supported by recent molecular data.

Pimplinae

Pimplinae are a worldwide subfamily of the parasitic wasp family Ichneumonidae.Pimplinae are parasitoids of Endopterygota, often the pupae of Lepidoptera. Various species parasitize the egg sacs and adults of spiders. There are 95 genera.

Pimplinae are generally sturdy black insects with orange markings. The first tergite is box-like with the spiracle anterior to the middle.

Pterogeniidae

Pterogeniidae is a family of beetles, in the large suborder Polyphaga.

Pterygota

The Pterygota are a subclass of insects that includes the winged insects. It also includes insect orders that are secondarily wingless (that is, insect groups whose ancestors once had wings but that have lost them as a result of subsequent evolution).The pterygotan group comprises almost all insects. The insect orders not included are the Archaeognatha (jumping bristletails) and the Zygentoma (silverfishes and firebrats), two primitively wingless insect orders. Also not included are the three orders no longer considered to be insects: Protura, Collembola, and Diplura.

Snakefly

Snakeflies are a group of insects comprising the order Raphidioptera, which is divided into two families: Raphidiidae and Inocelliidae consisting of roughly 260 species. Together with the Megaloptera they were formerly placed within the Neuroptera, but now these two are generally regarded as separate orders. Members of this order have been considered living fossils, as the phenotype of a species from the early Jurassic period (140 million years ago) closely resembles modern-day species.

Tephritoidea

The Tephritoidea are a superfamily of flies. The following families are included:

Pallopteridae — flutter flies

Piophilidae — skippers

Platystomatidae — signal flies

Pyrgotidae

Richardiidae

Tephritidae — fruit flies

Ulidiidae (Otitidae) — picture-winged fliesThe Tachiniscinae, formerly ranked as the family Tachiniscidae, are now included in the Tephritidae.

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