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Hemiptera (/hɛˈmɪptərə/; from Ancient Greek hemipterus 'half-winged') is an order of insects, commonly called true bugs, comprising over 80,000 species within groups such as the cicadasaphidsplanthoppersleafhoppersassassin bugsbed bugs, and shield bugs. They range in size from 1 mm (0.04 in) to around 15 cm (6 in), and share a common arrangement of piercing-sucking mouthparts.[3] The name "true bugs" is often limited to the suborder Heteroptera.[4]

Entomologists reserve the term bug for Hemiptera or Heteroptera,[5] which does not include other arthropods or insects of other orders such as antsbeesbeetles, or butterflies. In some variations of English, all terrestrial arthropods (including non-insect arachnids, and myriapods) also fall under the colloquial understanding of bug.[a]

Many insects with "bug" in their common name, especially in American English, belong to other orders; for example, the lovebug is a fly[9] and the Maybug and ladybug are beetles.[10] The term is occasionally extended to colloquial names for freshwater or marine crustaceans (e.g. Balmain bugMoreton Bay bugmudbug) and used by physicians and bacteriologists for disease-causing germs (e.g. superbugs).[5]

Most hemipterans feed on plants, using their sucking and piercing mouthparts to extract plant sap. Some are bloodsucking, or hematophagous, while others are predators that feed on other insects or small invertebrates. They live in a wide variety of habitats, generally terrestrial, though some are adapted to life in or on the surface of fresh water (e.g. pondskaterswater boatmengiant water bugs). Hemipterans are hemimetabolous, with young nymphs that somewhat resemble adults. Many aphids are capable of parthenogenesis, producing young from unfertilised eggs; this helps them to reproduce extremely rapidly in favourable conditions.

Humans have interacted with the Hemiptera for millennia. Some species, including many aphids, are significant agricultural pests, damaging crops by sucking the sap. Others harm humans more directly as vectors of serious viral diseases. The bed bug is a persistent parasite of humans, and some kissing bugs can transmit Chagas disease. Some species have been used for biological control of insect pests or of invasive plants. A few hemipterans, have been cultivated for the extraction of dyestuffs such as cochineal and carmine, and for shellacCicadas have been used as food, and have appeared in literature since the Iliad in Ancient Greece.

Diversity[edit]

Hemiptera is the largest order of hemimetabolous insects (not undergoing complete metamorphosis; though some examples such as male scale insects do undergo a form of complete metamorphosis [11]), containing over 95,000 named species. Other insect orders with more species are all holometabolous, meaning they have a pupal stage and undergo complete metamorphosis. The majority of species are terrestrial, including a number of important agricultural pests, but some are found in freshwater habitats. These include the water boatmenbackswimmerspond skaters, and giant water bugs.[12]

Taxonomy and phylogeny[edit]

Hemiptera belong to the insect superorder Paraneoptera, which includes lice (Psocodea), thrips (Thysanoptera), and the true bugs of Hemiptera. Within Paraneoptera, Hemiptera is most closely related to the sister clade Thysanoptera.[13]

The fossil record of hemipterans goes back to the Carboniferous (Moscovian).[14] The oldest fossils are of the Archescytinidae from the Lower Permian and are thought to be basal to the AuchenorrhynchaFulgoromorpha and Cicadomorpha appear in the Upper Permian, as do Sternorrhyncha of the Psylloidea and AleyrodoideaAphids and Coccoids appear in the Triassic. The Coleorrhyncha extend back to the Lower Jurassic.[15] The Heteroptera first appeared in the Triassic.[16]

The present members of the order Hemiptera (sometimes referred to as Rhynchota) were historically placed into two orders, the so-called Homoptera and Heteroptera/Hemiptera, based on differences in wing structure and the position of the rostrum. The order is now divided into four suborders, after the "Homoptera" were established as paraphyletic (now the Auchenorrhyncha and the Sternorrhyncha).[17][18][19]

The below cladogram shows Hemiptera's placement within Paraneoptera, as well as how Hemiptera's four suborders are related. English names are given in parentheses where possible.[13] 

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