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Insect

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Insect
Temporal range: 396–0 Ma
Early Devonian[1] (but see text) – Present
Insect collage.png
Clockwise from top left: dance fly (Empis livida), long-nosed weevil (Rhinotia hemistictus), mole cricket (Gryllotalpa brachyptera), German wasp (Vespula germanica), emperor gum moth (Opodiphthera eucalypti), assassin bug (Harpactorinae)
A chorus of several Magicicada species
Scientific classification e
Kingdom: Animalia
Clade: Euarthropoda
Clade: Pancrustacea
Subphylum: Hexapoda
Class: Insecta
Linnaeus, 1758
Subgroups

See text.

Synonyms
  • Ectognatha
  • Entomida

Insects or Insecta (from Latin insectum, a calque of Greek ἔντομον [éntomon], "cut into sections") are by far the largest group of hexapod invertebrates within the arthropod phylum. Definitions and circumscriptions vary; in one approach insects comprise a class within the Phylum Arthropoda. As the term is used here, it is synonymous with Ectognatha.

Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs of jointed legs, compound eyes and one pair of antennae. They are the most diverse group of animals on the planet, including more than a million described species and representing more than half of all known living organisms.[2][3] The number of extant species is estimated at between six and ten million,[2][4][5] and potentially represent over 90% of the differing animal life forms on Earth.[6] Insects may be found in nearly all environments, although only a small number of species reside in the oceans, a habitat dominated by another arthropod group, crustaceans.

The life cycles of insects vary but most hatch from eggs. Insect growth is constrained by the inelastic exoskeleton and development involves a series of molts. The immature stages can differ from the adults in structure, habit and habitat, and can include a passive pupal stage in those groups that undergo four-stage metamorphosis (see holometabolism). Insects that undergo three-stage metamorphosis lack a pupal stage and adults develop through a series of nymphal stages.[7] The higher level relationship of the Hexapoda is unclear. Fossilized insects of enormous size have been found from the Paleozoic Era, including giant dragonflies with wingspans of 55 to 70 cm (22–28 in). The most diverse insect groups appear to have coevolved with flowering plants.

Adult insects typically move about by walking, flying or sometimes swimming (see § Locomotion below). As it allows for rapid yet stable movement, many insects adopt a tripedal gait in which they walk with their legs touching the ground in alternating triangles. Insects are the only invertebrates to have evolved flight. Many insects spend at least part of their lives under water, with larval adaptations that include gills, and some adult insects are aquatic and have adaptations for swimming. Some species, such as water striders, are capable of walking on the surface of water. Insects are mostly solitary, but some, such as certain bees, ants and termites, are social and live in large, well-organized colonies. Some insects, such as earwigs, show maternal care, guarding their eggs and young. Insects can communicate with each other in a variety of ways. Male moths can sense the pheromones of female moths over great distances. Other species communicate with sounds: crickets stridulate, or rub their wings together, to attract a mate and repel other males. Lampyridae in the beetle order communicate with light.

Humans regard certain insects as pests, and attempt to control them using insecticides and a host of other techniques. Some insects damage crops by feeding on sap, leaves or fruits. A few parasitic species are pathogenic. Some insects perform complex ecological roles; blow-flies, for example, help consume carrion but also spread diseases. Insect pollinators are essential to the life cycle of many flowering plant species on which most organisms, including humans, are at least partly dependent; without them, the terrestrial portion of the biosphere (including humans) would be devastated.[8] Many other insects are considered ecologically beneficial as predators and a few provide direct economic benefit. Silkworms and bees have been used extensively by humans for the production of silk and honey, respectively. In some cultures, people eat the larvae or adults of certain insects.

Etymology[edit]

The word "insect" comes from the Latin word insectum, meaning "with a notched or divided body", or literally "cut into", from the neuter singular perfect passive participle of insectare, "to cut into, to cut up", from in- "into" and secare "to cut";[9] because insects appear "cut into" three sections. Pliny the Elder introduced the Latin designation as a loan-translation of the Greek word ἔντομος (éntomos) or "insect" (as in entomology), which was Aristotle's term for this class of life, also in reference to their "notched" bodies. "Insect" first appears documented in English in 1601 in Holland's translation of Pliny. Translations of Aristotle's term also form the usual word for "insect" in Welsh (trychfil, from trychu "to cut" and mil, "animal"), Serbo-Croatian (zareznik, from rezati, "to cut"), Russian (насекомое nasekomoje, from seč'/-sekat', "to cut"), etc.[9][10]

Definitions[edit]

The precise definition of the taxon Insecta and the equivalent English name "insect" varies; three alternative definitions are shown in the table.

Definition of Insecta
Group Alternative definitions
Collembola (springtails) Insecta sensu lato
=Hexapoda
Protura (coneheads)
Diplura (two-pronged bristletails)
Archaeognatha (jumping bristletails) Insecta sensu stricto
=Ectognatha
Zygentoma (silverfish)
Pterygota (winged insects) Insecta sensu strictissimo

In the broadest circumscription, Insecta sensu lato consists of all hexapods.[11][12] Traditionally, insects defined in this way were divided into "Apterygota" (the first five groups in the table) – the wingless insects – and Pterygota – the winged insects.[13] However, modern phylogenetic studies have shown that "Apterygota" is not monophyletic,[14] and so does not form a good taxon. A narrower circumscription restricts insects to those hexapods with external mouthparts, and comprises only the last three groups in the table. In this sense, Insecta sensu stricto is equivalent to Ectognatha.[11][14] In the narrowest circumscription, insects are restricted to hexapods that are either winged or descended from winged ancestors. Insecta sensu strictissimo is then equivalent to Pterygota.[15] For the purposes of this article, the middle definition is used; insects consist of two wingless taxa, Archaeognatha (jumping bristletails) and Zygentoma (silverfish), plus the winged or secondarily wingless Pterygota.

Phylogeny and evolution[edit]




Hexapoda (Insecta, Collembola, Diplura, Protura)



Crustacea (crabs, shrimp, isopods, etc.)



Myriapoda

Pauropoda



Diplopoda (millipedes)



Chilopoda (centipedes)



Symphyla



Chelicerata

Arachnida (spiders, scorpions, mites, ticks, etc.)



Eurypterida (sea scorpions: extinct)



Xiphosura (horseshoe crabs)



Pycnogonida (sea spiders)




Trilobites (extinct)



A phylogenetic tree of the arthropods and related groups[16]

Evolution has produced enormous variety in insects. Pictured are some of the possible shapes of antennae.

The evolutionary relationship of insects to other animal groups remains unclear.

Although traditionally grouped with millipedes and centipedes—possibly on the basis of convergent adaptations to terrestrialisation[17]—evidence has emerged favoring closer evolutionary ties with crustaceans. In the Pancrustacea theory, insects, together with Entognatha, Remipedia, and Cephalocarida, make up a natural clade labeled Miracrustacea.[18]

A report in November 2014 unambiguously places the insects in one clade, with the crustaceans and myriapods, as the nearest sister clades.[19] This study resolved insect phylogeny of all extant insect orders, and provides "a robust phylogenetic backbone tree and reliable time estimates of insect evolution".[19]

Other terrestrial arthropods, such as centipedes, millipedes, scorpions, and spiders, are sometimes confused with insects since their body plans can appear similar, sharing (as do all arthropods) a jointed exoskeleton. However, upon closer examination, their features differ significantly; most noticeably, they do not have the six-legged characteristic of adult insects.[20]

The higher-level phylogeny of the arthropods continues to be a matter of debate and research. In 2008, researchers at Tufts University uncovered what they believe is the world's oldest known full-body impression of a primitive flying insect, a 300-million-year-old specimen from the Carboniferous period.[21] The oldest definitive insect fossil is the Devonian Rhyniognatha hirsti, from the 396-million-year-old Rhynie chert. It may have superficially resembled a modern-day silverfish insect. This species already possessed dicondylic mandibles (two articulations in the mandible), a feature associated with winged insects, suggesting that wings may already have evolved at this time. Thus, the first insects probably appeared earlier, in the Silurian period.[1][22]

Four super radiations of insects have occurred: beetles (evolved about 300 million years ago), flies (evolved about 250 million years ago), and moths and wasps (evolved about 150 million years ago).[23] These four groups account for the majority of described species. The flies and moths along with the fleas evolved from the Mecoptera.

The origins of insect flight remain obscure, since the earliest winged insects currently known appear to have been capable fliers. Some extinct insects had an additional pair of winglets attaching to the first segment of the thorax, for a total of three pairs. As of 2009, no evidence suggests the insects were a particularly successful group of animals before they evolved to have wings.[24]

Late Carboniferous and Early Permian insect orders include both extant groups, their stem groups,[25] and a number of Paleozoic groups, now extinct. During this era, some giant dragonfly-like forms reached wingspans of 55 to 70 cm (22 to 28 in), making them far larger than any living insect. This gigantism may have been due to higher atmospheric oxygen levels that allowed increased respiratory efficiency relative to today. The lack of flying vertebrates could have been another factor. Most extinct orders of insects developed during the Permian period that began around 270 million years ago. Many of the early groups became extinct during the Permian-Triassic extinction event, the largest mass extinction in the history of the Earth, around 252 million years ago.[26]

The remarkably successful Hymenoptera appeared as long as 146 million years ago in the Cretaceous period, but achieved their wide diversity more recently in the Cenozoic era, which began 66 million years ago. A number of highly successful insect groups evolved in conjunction with flowering plants, a powerful illustration of coevolution.[27]

Many modern insect genera developed during the Cenozoic. Insects from this period on are often found preserved in amber, often in perfect condition. The body plan, or morphology, of such specimens is thus easily compared with modern species. The study of fossilized insects is called paleoentomology.

Evolutionary relationships[edit]

Insects are prey for a variety of organisms, including terrestrial vertebrates. The earliest vertebrates on land existed 400 million years ago and were large amphibious piscivores. Through gradual evolutionary change, insectivory was the next diet type to evolve.[28]

Insects were among the earliest terrestrial herbivores and acted as major selection agents on plants.[27] Plants evolved chemical defenses against this herbivory and the insects, in turn, evolved mechanisms to deal with plant toxins. Many insects make use of these toxins to protect themselves from their predators. Such insects often advertise their toxicity using warning colors.[29] This successful evolutionary pattern has also been used by mimics. Over time, this has led to complex groups of coevolved species. Conversely, some interactions between plants and insects, like pollination, are beneficial to both organisms. Coevolution has led to the development of very specific mutualisms in such systems.

Taxonomy[edit]

Classification
Insecta
Dicondylia
Pterygota

Cladogram of living insect groups,[30] with numbers of species in each group.[5] The Apterygota, Palaeoptera, and Exopterygota are possibly paraphyletic groups.

Traditional morphology-based or appearance-based systematics have usually given the Hexapoda the rank of superclass,[31]:180 and identified four groups within it: insects (Ectognatha), springtails (Collembola), Protura, and Diplura, the latter three being grouped together as the Entognatha on the basis of internalized mouth parts. Supraordinal relationships have undergone numerous changes with the advent of methods based on evolutionary history and genetic data. A recent theory is that the Hexapoda are polyphyletic (where the last common ancestor was not a member of the group), with the entognath classes having separate evolutionary histories from the Insecta.[32] Many of the traditional appearance-based taxa have been shown to be paraphyletic, so rather than using ranks like subclass, superorder, and infraorder, it has proved better to use monophyletic groupings (in which the last common ancestor is a member of the group). The following represents the best-supported monophyletic groupings for the Insecta.

Insects can be divided into two groups historically treated as subclasses: wingless insects, known as Apterygota, and winged insects, known as Pterygota. The Apterygota consist of the primitively wingless order of the silverfish (Zygentoma). Archaeognatha make up the Monocondylia based on the shape of their mandibles, while Zygentoma and Pterygota are grouped together as Dicondylia. The Zygentoma themselves possibly are not monophyletic, with the family Lepidotrichidae being a sister group to the Dicondylia (Pterygota and the remaining Zygentoma).[33][34]

Paleoptera and Neoptera are the winged orders of insects differentiated by the presence of hardened body parts called sclerites, and in the Neoptera, muscles that allow their wings to fold flatly over the abdomen. Neoptera can further be divided into incomplete metamorphosis-based (Polyneoptera and Paraneoptera) and complete metamorphosis-based groups. It has proved difficult to clarify the relationships between the orders in Polyneoptera because of constant new findings calling for revision of the taxa. For example, the Paraneoptera have turned out to be more closely related to the Endopterygota than to the rest of the Exopterygota. The recent molecular finding that the traditional louse orders Mallophaga and Anoplura are derived from within Psocoptera has led to the new taxon Psocodea.[35] Phasmatodea and Embiidina have been suggested to form the Eukinolabia.[36] Mantodea, Blattodea, and Isoptera are thought to form a monophyletic group termed Dictyoptera.[37]

The Exopterygota likely are paraphyletic in regard to the Endopterygota. Matters that have incurred controversy include Strepsiptera and Diptera grouped together as Halteria based on a reduction of one of the wing pairs – a position not well-supported in the entomological community.[38] The Neuropterida are often lumped or split on the whims of the taxonomist. Fleas are now thought to be closely related to boreid mecopterans.[39] Many questions remain in the basal relationships among endopterygote orders, particularly the Hymenoptera.

The study of the classification or taxonomy of any insect is called systematic entomology. If one works with a more specific order or even a family, the term may also be made specific to that order or family, for example systematic dipterology.

Diversity[edit]

Though the true dimensions of species diversity remain uncertain, estimates range from 2.6–7.8 million species with a mean of 5.5 million.[40]

A pie chart of described eukaryote species, showing just over half of these to be insects

Between 950,000–1,000,000 of all described species are insects, so over 50% of all described eukaryotes (1.8 million) are insects (see illustration). With only 950,000 known non-insects, if the actual number of insects is 5.5 million, they may represent over 80% of the total, and with only about 20,000 new species of all organisms being described each year, most insect species likely will remain undescribed, unless species descriptions greatly increase in rate. Of the 24 orders of insects, four dominate in terms of numbers of described species, with at least 670,000 species included in Coleoptera, Diptera, Hymenoptera and Lepidoptera.

Comparison of the estimated number of species in the four most speciose insect orders[citation needed]
Described species Average description rate
(species per year)
Publication effort
Coleoptera 300,000–400,000 2308 0.01
Lepidoptera[41] 180,000 642 0.03
Diptera 90,000–150,000 1048 0.04
Hymenoptera 100,000–150,000 1196 0.02

A 2015 study estimated the number of beetle species at 0.9–2.1 million with a mean of 1.5 million.[40]

Morphology and physiology[edit]

External[edit]

Insect morphology
A- Head B- Thorax C- Abdomen
1. antenna
2. ocelli (lower)
3. ocelli (upper)
4. compound eye
5. brain (cerebral ganglia)
6. prothorax
7. dorsal blood vessel
8. tracheal tubes (trunk with spiracle)
9. mesothorax
10. metathorax
11. forewing
12. hindwing
13. mid-gut (stomach)
14. dorsal tube (Heart)
15. ovary
16. hind-gut (intestine, rectum & anus)
17. anus
18. oviduct
19. nerve chord (abdominal ganglia)
20. Malpighian tubes
21. tarsal pads
22. claws
23. tarsus
24. tibia
25. femur
26. trochanter
27. fore-gut (crop, gizzard)
28. thoracic ganglion
29. coxa
30. salivary gland
31. subesophageal ganglion
32. mouthparts

Insects have segmented bodies supported by exoskeletons, the hard outer covering made mostly of chitin. The segments of the body are organized into three distinctive but interconnected units, or tagmata: a head, a thorax and an abdomen.[42] The head supports a pair of sensory antennae, a pair of compound eyes, and, if present, one to three simple eyes (or ocelli) and three sets of variously modified appendages that form the mouthparts. The thorax has six segmented legs—one pair each for the prothorax, mesothorax and the metathorax segments making up the thorax—and, none, two or four wings. The abdomen consists of eleven segments, though in a few species of insects, these segments may be fused together or reduced in size. The abdomen also contains most of the digestive, respiratory, excretory and reproductive internal structures.[31]:22–48 Considerable variation and many adaptations in the body parts of insects occur, especially wings, legs, antenna and mouthparts.

Segmentation[edit]

The head is enclosed in a hard, heavily sclerotized, unsegmented, exoskeletal head capsule, or epicranium, which contains most of the sensing organs, including the antennae, ocellus or eyes, and the mouthparts. Of all the insect orders, Orthoptera displays the most features found in other insects, including the sutures and sclerites.[43] Here, the vertex, or the apex (dorsal region), is situated between the compound eyes for insects with a hypognathous and opisthognathous head. In prognathous insects, the vertex is not found between the compound eyes, but rather, where the ocelli are normally. This is because the primary axis of the head is rotated 90° to become parallel to the primary axis of the body. In some species, this region is modified and assumes a different name.[43]:13

The thorax is a tagma composed of three sections, the prothorax, mesothorax and the metathorax. The anterior segment, closest to the head, is the prothorax, with the major features being the first pair of legs and the pronotum. The middle segment is the mesothorax, with the major features being the second pair of legs and the anterior wings. The third and most posterior segment, abutting the abdomen, is the metathorax, which features the third pair of legs and the posterior wings. Each segment is dilineated by an intersegmental suture. Each segment has four basic regions. The dorsal surface is called the tergum (or notum) to distinguish it from the abdominal terga.[31] The two lateral regions are called the pleura (singular: pleuron) and the ventral aspect is called the sternum. In turn, the notum of the prothorax is called the pronotum, the notum for the mesothorax is called the mesonotum and the notum for the metathorax is called the metanotum. Continuing with this logic, the mesopleura and metapleura, as well as the mesosternum and metasternum, are used.[43]

The abdomen is the largest tagma of the insect, which typically consists of 11–12 segments and is less strongly sclerotized than the head or thorax. Each segment of the abdomen is represented by a sclerotized tergum and sternum. Terga are separated from each other and from the adjacent sterna or pleura by membranes. Spiracles are located in the pleural area. Variation of this ground plan includes the fusion of terga or terga and sterna to form continuous dorsal or ventral shields or a conical tube. Some insects bear a sclerite in the pleural area called a laterotergite. Ventral sclerites are sometimes called laterosternites. During the embryonic stage of many insects and the postembryonic stage of primitive insects, 11 abdominal segments are present. In modern insects there is a tendency toward reduction in the number of the abdominal segments, but the primitive number of 11 is maintained during embryogenesis. Variation in abdominal segment number is considerable. If the Apterygota are considered to be indicative of the ground plan for pterygotes, confusion reigns: adult Protura have 12 segments, Collembola have 6. The orthopteran family Acrididae has 11 segments, and a fossil specimen of Zoraptera has a 10-segmented abdomen.[43]

Exoskeleton[edit]

The insect outer skeleton, the cuticle, is made up of two layers: the epicuticle, which is a thin and waxy water resistant outer layer and contains no chitin, and a lower layer called the procuticle. The procuticle is chitinous and much thicker than the epicuticle and has two layers: an outer layer known as the exocuticle and an inner layer known as the endocuticle. The tough and flexible endocuticle is built from numerous layers of fibrous chitin and proteins, criss-crossing each other in a sandwich pattern, while the exocuticle is rigid and hardened.[31]:22–24 The exocuticle is greatly reduced in many soft-bodied insects (e.g., caterpillars), especially during their larval stages.

Insects are the only invertebrates to have developed active flight capability, and this has played an important role in their success.[31]:186 Their muscles are able to contract multiple times for each single nerve impulse, allowing the wings to beat faster than would ordinarily be possible. Having their muscles attached to their exoskeletons is more efficient and allows more muscle connections; crustaceans also use the same method, though all spiders use hydraulic pressure to extend their legs, a system inherited from their pre-arthropod ancestors. Unlike insects, though, most aquatic crustaceans are biomineralized with calcium carbonate extracted from the water.[44][45]

Internal[edit]

Nervous system[edit]

The nervous system of an insect can be divided into a brain and a ventral nerve cord. The head capsule is made up of six fused segments, each with either a pair of ganglia, or a cluster of nerve cells outside of the brain. The first three pairs of ganglia are fused into the brain, while the three following pairs are fused into a structure of three pairs of ganglia under the insect's esophagus, called the subesophageal ganglion.[31]:57

The thoracic segments have one ganglion on each side, which are connected into a pair, one pair per segment. This arrangement is also seen in the abdomen but only in the first eight segments. Many species of insects have reduced numbers of ganglia due to fusion or reduction.[46] Some cockroaches have just six ganglia in the abdomen, whereas the wasp Vespa crabro has only two in the thorax and three in the abdomen. Some insects, like the house fly Musca domestica, have all the body ganglia fused into a single large thoracic ganglion.

At least a few insects have nociceptors, cells that detect and transmit signals responsible for the sensation of pain.[47][not in citation given] This was discovered in 2003 by studying the variation in reactions of larvae of the common fruitfly Drosophila to the touch of a heated probe and an unheated one. The larvae reacted to the touch of the heated probe with a stereotypical rolling behavior that was not exhibited when the larvae were touched by the unheated probe.[48] Although nociception has been demonstrated in insects, there is no consensus that insects feel pain consciously[49]

Insects are capable of learning.[50]

Digestive system[edit]

An insect uses its digestive system to extract nutrients and other substances from the food it consumes.[51] Most of this food is ingested in the form of macromolecules and other complex substances like proteins, polysaccharides, fats and nucleic acids. These macromolecules must be broken down by catabolic reactions into smaller molecules like amino acids and simple sugars before being used by cells of the body for energy, growth, or reproduction. This break-down process is known as digestion.

The main structure of an insect's digestive system is a long enclosed tube called the alimentary canal, which runs lengthwise through the body. The alimentary canal directs food unidirectionally from the mouth to the anus. It has three sections, each of which performs a different process of digestion. In addition to the alimentary canal, insects also have paired salivary glands and salivary reservoirs. These structures usually reside in the thorax, adjacent to the foregut.[31]:70–77

The salivary glands (element 30 in numbered diagram) in an insect's mouth produce saliva. The salivary ducts lead from the glands to the reservoirs and then forward through the head to an opening called the salivarium, located behind the hypopharynx. By moving its mouthparts (element 32 in numbered diagram) the insect can mix its food with saliva. The mixture of saliva and food then travels through the salivary tubes into the mouth, where it begins to break down.[52][53] Some insects, like flies, have extra-oral digestion. Insects using extra-oral digestion expel digestive enzymes onto their food to break it down. This strategy allows insects to extract a significant proportion of the available nutrients from the food source.[54]:31 The gut is where almost all of insects' digestion takes place. It can be divided into the foregut, midgut and hindgut.

Foregut[