Hemiptera: Difference between revisions

Hemiptera; Temporal range: Moscovian-Holocene, 307–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N
	Species from each of the four suborders of Hemiptera, clockwise from top-left: Acanthosoma labiduroides (Heteroptera), Xenophyes forsteri (Coleorrhyncha), Magicicada septendecim (Auchenorrhyncha), and Aphids (Sternorrhyncha).
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
(unranked):	Paraneoptera
Superorder:	Condylognatha
Order:	Hemiptera; Linnaeus, 1758
Suborders
	Auchenorrhyncha; Coleorrhyncha; Heteroptera; Sternorrhyncha;

Browse history interactively

← Previous edit Next edit →

Content deleted Content added

VisualWikitext

Inline

Revision as of 09:48, 18 July 2024

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 cicadas, aphids, planthoppers, leafhoppers, assassin bugs, bed 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 ants, bees, beetles, 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 bug, Moreton Bay bug, mudbug) 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. pondskaters, water boatmen, giant 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 shellac. Cicadas have been used as food, and have appeared in literature since the Iliad in Ancient Greece.

Diversity

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 boatmen, backswimmers, pond skaters, and giant water bugs.^[12]

Taxonomy and phylogeny

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 Auchenorrhyncha. Fulgoromorpha and Cicadomorpha appear in the Upper Permian, as do Sternorrhyncha of the Psylloidea and Aleyrodoidea. Aphids 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]

Other insects

Paraneoptera

Psocodea

Trogiomorpha (granary booklice and relatives)

Psocomorpha (barklice)

Troctomorpha

Amphientometae

Sphaeropsocidae

	Liposcelididae (booklice)

	Phthiraptera (lice)

(paraphyletic with respect to Phthiraptera)

Condylognatha

Thysanoptera (thrips)

Hemiptera

Sternorrhyncha (aphids)

Heteroptera (shield bugs, assassin bugs, etc)

Coleorrhyncha (moss bugs)

Auchenorrhyncha

	Fulgoromorpha (planthoppers)

	Cicadomorpha (cicadas, leafhoppers, spittlebugs, etc)

(true bugs)

Hemiptera suborders
Suborder	No. of Species	First appearance	Examples	Characteristics
Auchenorrhyncha	over 42,000^[20]	Lower Permian	cicadas, leafhoppers, treehoppers, planthoppers, froghoppers	plant-sucking bugs; many can jump; many make calls, some loud
Coleorrhyncha	fewer than 30	Lower Jurassic	moss bugs (Peloridiidae)	small, rarely observed; found in/feed on moss; evolved before the splitting of Gondwana
Heteroptera	over 45,000^[21]	Triassic	shield bugs, seed bugs, assassin bugs, flower bugs, leaf-footed bugs, water bugs, plant bugs	larger bugs; some are predatory, others are plant-sucking
Sternorrhyncha	12,500	Upper Permian	aphids, whiteflies, scale insects	plant-sucking bugs, some major horticultural pests; most are small and sedentary or fully sessile;^[22]

Biology

Mouthparts

The defining feature of hemipterans is their "beak" in which the modified mandibles and maxillae form a "stylet" which is sheathed within a modified labium. The stylet is capable of piercing tissues and sucking liquids, while the labium supports it. The stylet contains a channel for the outward movement of saliva and another for the inward movement of liquid food. A salivary pump drives saliva into the prey; a cibarial pump extracts liquid from the prey. Both pumps are powered by substantial dilator muscles in the head. The beak is usually folded under the body when not in use. The diet is typically plant sap, but some hemipterans such as assassin bugs are predators.^[23]^[24]

Both herbivorous and predatory hemipterans inject enzymes to begin digestion extra-orally (before the food is taken into the body). These enzymes include amylase to hydrolyse starch, polygalacturonase to weaken the tough cell walls of plants, and proteinases to break down proteins.^[25]

Although the Hemiptera vary widely in their overall form, their mouthparts form a distinctive "rostrum". Other insect orders with mouthparts modified into anything like the rostrum and stylets of the Hemiptera include some Phthiraptera, but for other reasons they generally are easy to recognize as non-hemipteran. Similarly, the mouthparts of Siphonaptera, some Diptera and Thysanoptera superficially resemble the rostrum of the Hemiptera, but on closer inspection the differences are considerable. Aside from the mouthparts, various other insects can be confused with Hemiptera, but they all have biting mandibles and maxillae instead of the rostrum. Examples include cockroaches and psocids, both of which have longer, many-segmented antennae, and some beetles, but these have fully hardened forewings which do not overlap.^[26]

Wing structure

The forewings of Hemiptera are either entirely membranous, as in the Sternorrhyncha and Auchenorrhyncha, or partially hardened, as in most Heteroptera. The name "Hemiptera" is from the Greek ἡμι- (hemi; "half") and πτερόν (pteron; "wing"), referring to the forewings of many heteropterans which are hardened near the base, but membranous at the ends. Wings modified in this manner are termed hemelytra (singular: hemelytron), by analogy with the completely hardened elytra of beetles, and occur only in the suborder Heteroptera. In all suborders, the hindwings – if present at all – are entirely membranous and usually shorter than the forewings.^[12] The forewings may be held "roofwise" over the body (typical of Sternorrhyncha and Auchenorrhyncha),^[27] or held flat on the back, with the ends overlapping (typical of Heteroptera).^[12] The antennae in Hemiptera typically consist of four or five segments, although they can still be quite long, and the tarsi of the legs have two or three segments.^[28]

Sound production

Many hemipterans can produce sound for communication.^[29] The "song" of male cicadas, the loudest of any insect, is produced by tymbal organs on the underside of the abdomen, and is used to attract mates. The tymbals are drumlike disks of cuticle, which are clicked in and out repeatedly, making a sound in the same way as popping the metal lid of a jam jar in and out.^[30]

Stridulatory sounds are produced among the aquatic Corixidae and Notonectidae (backswimmers) using tibial combs rubbed across rostral ridges.^[31]^[32]

Life cycle

Hemipterans are hemimetabolous, meaning that they do not undergo metamorphosis, the complete change of form between a larval phase and an adult phase. Instead, their young are called nymphs, and resemble the adults to a greater or lesser degree. The nymphs moult several times as they grow, and each instar resembles the adult more than the previous one. Wing buds grow in later stage nymphs; the final transformation involves little more than the development of functional wings (if they are present at all) and functioning sexual organs, with no intervening pupal stage as in holometabolous insects.^[33]

Parthenogenesis and viviparity

Many aphids are parthenogenetic during part of the life cycle, such that females can produce unfertilized eggs, which are clones of their mother. All such young are females (thelytoky), so 100% of the population at these times can produce more offspring. Many species of aphid are also viviparous: the young are born live rather than laid as eggs. These adaptations enable aphids to reproduce extremely rapidly when conditions are suitable.^[34]

Locomotion

Hemipterans make use of a variety of modes of locomotion including swimming, skating on a water surface and jumping, as well as walking and flying like other insects.

Swimming and skating

Several families of Heteroptera are water bugs, adapted to an aquatic lifestyle, such as the water boatmen (Corixidae), water scorpions (Nepidae), and backswimmers (Notonectidae). They are mostly predatory, and have legs adapted as paddles to help the animal move through the water.^[35] The pondskaters or water striders (Gerridae) are also associated with water, but use the surface tension of standing water to keep them above the surface;^[36] they include the sea skaters in the genus Halobates, the only truly marine group of insects.^[35]

Marangoni propulsion

Marangoni effect propulsion exploits the change in surface tension when a soap-like surfactant is released on to a water surface, in the same way that a toy soap boat propels itself. Water bugs in the genus Microvelia (Veliidae) can travel at up to 17 cm/s, twice as fast as they can walk, by this means.^[36]

Flight

Flight is well developed in the Hemiptera although mostly used for short distance movement and dispersal. Wing development is sometimes related to environmental conditions. In some groups of Hemiptera, there are variations of winged, short-winged, and wingless forms within a single species. This kind of polymorphism tends to be helpful when habitats are temporary with more energy put into reproduction when food is available and into dispersal through flight when food becomes scarce. In aphids, both winged and wingless forms occur with winged forms produced in greater numbers when food resources are depleted. Aphids and whiteflies can sometimes be transported very long distances by atmospheric updrafts and high altitude winds.^[37] Wing-length polymorphism is notably rare in tree-living Hemiptera.^[38]

Jumping

Many Auchenorrhyncha including representatives of the cicadas, leafhoppers, treehoppers, planthoppers, and froghoppers are adapted for jumping (saltation). Treehoppers, for example, jump by rapidly depressing their hind legs. Before jumping, the hind legs are raised and the femora are pressed tightly into curved indentations in the coxae. Treehoppers can attain a take-off velocity of up to 2.7 metres per second and an acceleration of up to 250 g. The instantaneous power output is much greater than that of normal muscle, implying that energy is stored and released to catapult the insect into the air.^[39] Cicadas, which are much larger, extend their hind legs for a jump in under a millisecond, again implying elastic storage of energy for sudden release.^[40]

Sedentary

Instead of relying on any form of locomotion, most Sternorrhyncha females are sedentary or completely sessile, attached to their host plants by their thin feeding stylets which cannot be taken out of the plant quickly.^[41]

Ecological roles

Feeding modes

Herbivores

Most hemipterans are phytophagous, using their sucking and piercing mouthparts to feed on plant sap. These include cicadas, leafhoppers, treehoppers, planthoppers, froghoppers, aphids, whiteflies, scale insects, and some other groups. Some are monophages, being host specific and only found on one plant taxon, others are oligophages, feeding on a few plant groups, while others again are less discriminating polyphages and feed on many species of plant.^[24] The relationship between hemipterans and plants appears to be ancient, with piercing and sucking of plants evident in the Early Devonian period.^[42]

Hemipterans can dramatically cut the mass of affected plants, especially in major outbreaks. They sometimes also change the mix of plants by predation on seeds or feeding on roots of certain species.^[43] Some sap-suckers move from one host to another at different times of year. Many aphids spend the winter as eggs on a woody host plant and the summer as parthenogenetically reproducing females on a herbaceous plant.^[44]

Phloem sap, which has a higher concentration of sugars and nitrogen, is under positive pressure unlike the more dilute xylem sap. Most of the Sternorrhyncha and a number of Auchenorrhynchan groups feed on phloem. Phloem feeding is common in the Fulgoromorpha, most Cicadellidae and in the Heteroptera.

The Typhlocybine Cicadellids specialize in feeding on non-vascular mesophyll tissue of leaves, which is more nutritious than the leaf epidermis. Most Heteroptera also feed on mesophyll tissue where they are more likely to encounter defensive secondary plant metabolites which often leads to the evolution of host specificity.^[45]

Obligate xylem feeding is a special habit that is found in the Auchenorrhyncha among Cicadoidea, Cercopoidea and in Cicadelline Cicadellids. Some phloem feeders may take to xylem sap facultatively, especially when facing dehydration.^[46] Xylem feeders tend to be polyphagous;^[47] to overcome the negative pressure of xylem requires a special cibarial pump.^[48]

Phloem feeding hemiptera typically have symbiotic micro-organisms in their gut that help to convert amino acids. Phloem feeders produce honeydew from their anus. A variety of organisms that feed on honeydew form symbiotic associations with phloem-feeders.^[49]^[50] Phloem sap is a sugary liquid low in amino acids, so insects have to process large quantities to meet their nutritional requirements. Xylem sap is even lower in amino acids and contains monosaccharides rather than sucrose, as well as organic acids and minerals. No digestion is required (except for the hydrolysis of sucrose) and 90% of the nutrients in the xylem sap can be utilised.^[24]^[51] Some phloem sap feeders selectively mix phloem and xylem sap to control the osmotic potential of the liquid consumed.^[52]

A striking adaptation to a very dilute diet is found in many hemipterans: a filter chamber, a part of the gut looped back on itself as a countercurrent exchanger, which permits nutrients to be separated from excess water.^[53] The residue, mostly water with sugars and amino acids, is quickly excreted as sticky "honey dew", notably from aphids but also from other Auchenorrhycha and Sternorrhyncha.^[54]

Some Sternorrhyncha including Psyllids and some aphids are gall formers. These sap-sucking hemipterans inject fluids containing plant hormones into the plant tissues inducing the production of tissue that covers to protects the insect and also act as sinks for nutrition that they feed on. The hackleberry gall psyllid for example, causes a woody gall on the leaf petioles of the hackleberry tree it infests,^[55] and the nymph of another psyllid produces a protective lerp out of hardened honeydew.^[24]

Predators

Most other hemipterans are predatory, feeding on other insects, or even small vertebrates. This is true of many aquatic species which are predatory, either as nymphs or adults.^[28] The predatory shield bug for example stabs caterpillars with its beak and sucks out the body fluids.^[56] The saliva of predatory heteropterans contains digestive enzymes such as proteinase and phospholipase, and in some species also amylase. The mouthparts of these insects are adapted for predation. There are toothed stylets on the mandibles able to cut into and abrade tissues of their prey. There are further stylets on the maxillae, adapted as tubular canals to inject saliva and to extract the pre-digested and liquified contents of the prey.^[57]

Haematophagic ectoparasites

A few hemipterans are haematophagic ectoparasites^[58]), feeding on the blood of larger animals. These include bedbugs and the triatomine kissing bugs of the assassin bug family Reduviidae, which can transmit the dangerous Chagas disease.^[35] The first known hemipteran to feed in this way on vertebrates was the extinct assassin bug Triatoma dominicana found fossilized in amber and dating back about twenty million years. Faecal pellets fossilised beside it show that it transmitted a disease-causing Trypanosoma and the amber included hairs of the likely host, a bat.^[59]

As symbionts

Some species of ant protect and farm aphids (Sternorrhyncha) and other sap-sucking hemipterans, gathering and eating the honeydew that these hemipterans secrete. The relationship is mutualistic, as both ant and aphid benefit. Ants such as the yellow anthill ant, Lasius flavus, breed aphids of at least four species, Geoica utricularia, Tetraneura ulmi, Forda marginata and Forda formicaria, taking eggs with them when they found a new colony; in return, these aphids are obligately associated with the ant, breeding mainly or wholly asexually inside anthills.^[60] Ants may also protect the plant bugs from their natural enemies, removing the eggs of predatory beetles and preventing access by parasitic wasps.^[24]

Some leafhoppers (Auchenorrhyncha) are similarly "milked" by ants. In the Corcovado rain forest of Costa Rica, wasps compete with ants to protect and milk leafhoppers; the leafhoppers preferentially give more honeydew, more often, to the wasps, which are larger and may offer better protection.^[61]

As prey: defences against predators and parasites

Hemiptera form prey to predators including vertebrates, such as birds, and other invertebrates such as ladybirds.^[62]^[63] In response, hemipterans have evolved antipredator adaptations. Ranatra may feign death (thanatosis). Others such as Carpocoris purpureipennis secrete toxic fluids to ward off arthropod predators; some Pentatomidae such as Dolycoris are able to direct these fluids at an attacker. Toxic cardenolide compounds are accumulated by the heteropteran Oncopeltus fasciatus when it consumes milkweeds, while the coreid stinkbug Amorbus rubiginosus acquires 2-hexenal from its food plant, Eucalyptus. Some long-legged bugs mimic twigs, rocking to and fro to simulate the motion of a plant part in the wind.^[63] The nymph of the Masked hunter bug camouflages itself with sand grains, using its hind legs and tarsal fan to form a double layer of grains, coarser on the outside.^[64] The Amazon rain forest cicada Hemisciera maculipennis displays bright red deimatic flash coloration on its hindwings when threatened; the sudden contrast helps to startle predators, giving the cicada time to escape. The coloured patch on the hindwing is concealed at rest by an olive green patch of the same size on the forewing, enabling the insect to switch rapidly from cryptic to deimatic behaviour.^[65]^[b]

Some hemipterans such as firebugs have bold aposematic warning coloration, often red and black, which appear to deter passerine birds.^[67]^[68] Many hemipterans including aphids, scale insects and especially the planthoppers secrete wax to protect themselves from threats such as fungi, parasitoidal insects and predators, as well as abiotic factors like desiccation.^[69] Hard waxy coverings are especially important in the sedentary Sternorrhyncha such as scale insects, which have no means of escaping from predators; other Sternorrhyncha evade detection and attack by creating and living inside plant galls.^[41] Nymphal Cicadoidea and Cercopoidea have glands attached to the Malpighian tubules in their proximal segment that produce mucopolysaccharides, which form the froth around spittlebugs, offering a measure of protection.^[70]

Parental care is found in many species of Hemiptera especially in members of the Membracidae and numerous Heteroptera. In many species of shield bug, females stand guard over their egg clusters to protect them from egg parasitoids and predators.^[71] In the aquatic Belostomatidae, females lay their eggs on the back of the male which guards the eggs.^[72] Protection provided by ants is common in the Auchenorrhyncha.^[24]

Interaction with humans

As pests

Although many species of Hemiptera are significant pests of crops and garden plants, including many species of aphid and scale insects, other species are harmless. The damage done is often not so much the deprivation of the plant of its sap, but the fact that they transmit serious viral diseases between plants.^[73] They often produce copious amounts of honeydew which encourages the growth of sooty mould.^[74] Significant pests include the cottony cushion scale, a pest of citrus fruit trees,^[75] the green peach aphid and other aphids which attack crops worldwide and transmit diseases,^[76] and jumping plant lice which are often host plant-specific and transmit diseases.^{[citation needed]}

For pest control

genus Podisus Spined soldier bug eggs and then the recently hatched first instar bugs

Members of the families Reduviidae, Phymatidae and Nabidae are obligate predators. Some predatory species are used in biological pest control; these include various nabids,^[77] and even some members of families that are primarily phytophagous, such as the genus Geocoris in the family Lygaeidae.^[78] Other hemipterans are omnivores, alternating between a plant-based and an animal-based diet. For example, Dicyphus hesperus is used to control whitefly on tomatoes but also sucks sap, and if deprived of plant tissues will die even if in the presence of whiteflies.^[79]

The spined soldier bug (Podisus maculiventris) sucks body fluids from several pests including the larvae of the Colorado beetle and the Mexican bean beetle.^[80]

Insect products

Other hemipterans have positive uses for humans, such as in the production of the dyestuff carmine (cochineal). The FDA has created guidelines for how to declare when it has been added to a product.^[81] The scale insect Dactylopius coccus produces the brilliant red-coloured carminic acid to deter predators. Up to 100,000 scale insects need to be collected and processed to make a kilogram (2.2 lbs) of cochineal dye.^[82] A similar number of lac bugs are needed to make a kilogram of shellac, a brush-on colourant and wood finish.^[83] Additional uses of this traditional product include the waxing of citrus fruits to extend their shelf-life, and the coating of pills to moisture-proof them, provide slow-release or mask the taste of bitter ingredients.^[84]

As human parasites and disease vectors

Chagas disease is a modern-day tropical disease caused by Trypanosoma cruzi and transmitted by kissing bugs, so-called because they suck human blood from around the lips while a person sleeps.^[85]

The bed bug, Cimex lectularius, is an external parasite of humans. It lives in bedding and is mainly active at night, feeding on human blood, generally without being noticed.^[86]^[87] Bed bugs mate by traumatic insemination; the male pierces the female's abdomen and injects his sperm into a secondary genital structure, the spermalege. The sperm travel in the female's blood (haemolymph) to sperm storage structures (seminal conceptacles); they are released from there to fertilise her eggs inside her ovaries.^[87]^[88]

As food

Some larger hemipterans such as cicadas are used as food in Asian countries such as China,^[89] and they are much esteemed in Malawi and other African countries. Insects have a high protein content and good food conversion ratios, but most hemipterans are too small to be a useful component of the human diet.^[90] At least nine species of Hemiptera are eaten worldwide.^[91]

In art and literature

Cicadas have featured in literature since the time of Homer's Iliad, and as motifs in decorative art from the Chinese Shang dynasty (1766–1122 B.C.). They are described by Aristotle in his History of Animals and by Pliny the Elder in his Natural History; their mechanism of sound production is mentioned by Hesiod in his poem Works and Days "when the Skolymus flowers, and the tuneful Tettix sitting on his tree in the weary summer season pours forth from under his wings his shrill song".^[92]

In mythology and folklore

Among the bugs, cicadas in particular have been used as money, in folk medicine, to forecast the weather, to provide song (in China), and in folklore and myths around the world.^[93]

Threats

Large-scale cultivation of the oil palm Elaeis guineensis in the Amazon basin damages freshwater habitats and reduces the diversity of aquatic and semi-aquatic Heteroptera.^[94] Climate change may be affecting the global migration of hemipterans including the potato leafhopper, Empoasca fabae. Warming is correlated with the severity of potato leafhopper infestation, so increased warming may worsen infestations in future.^[95]

Notes

^ The Museum of New Zealand notes that "in everyday conversation", bug "refers to land arthropods with at least six legs, such as insects, spiders, and centipedes".^[6] In a chapter on "Bugs That Are Not Insects", the entomologist Gilbert Walbauer specifies centipedes, millipedes, arachnids (spiders, daddy longlegs, scorpions, mites, chiggers and ticks) as well as the few terrestrial crustaceans (sowbugs and pillbugs),^[7] but argues that "including legless creatures such as worms, slugs, and snails among the bugs stretches the word too much".^[5] "Bug" is used for a harmful microorganism, typically a bacterium.^[8]
^ The green/red flash coloration earns the insect the name of stop and go cicada.^[66]

References

^ Wang, Yan-hui; Engel, Michael S.; Rafael, José A.; Wu, Hao-yang; Rédei, Dávid; et al. (2016). "Fossil record of stem groups employed in evaluating the chronogram of insects (Arthropoda: Hexapoda)". Scientific Reports. 6: 38939. Bibcode:2016NatSR...638939W. doi:10.1038/srep38939. PMC 5154178. PMID 27958352.
^ "Hemiptera". Integrated Taxonomic Information System.
^ "Hemiptera: bugs, aphids and cicadas". Commonwealth Scientific and Industrial Research Organisation. Retrieved 8 May 2007.
^ "Suborder Heteroptera – True Bugs". Bug guide. Iowa State University Entomology. n.d.
^ ^a ^b ^c Gilbert Waldbauer. The Handy Bug Answer Book. Visible Ink, 1998. p. 1. ISBN 9781578590490
^ "What is a bug? Insects, arachnids, and myriapods" at Museum of New Zealand Te Papa Tongarewa website. Accessed 10 March 2022.
^ Gilbert Waldbauer. The Handy Bug Answer Book. Visible Ink, 1998. pp. 5–26. ISBN 9781578590490
^ "BUG | Meaning & Definition for UK English". Lexico.com. Archived from the original on 11 December 2019. Retrieved 3 August 2022.
^ Denmark, Harold; Mead, Frank; Fasulo, Thomas (April 2010). "Lovebug, Plecia nearctica Hardy". Featured Creatures. University of Florida/IFAS. Retrieved 10 March 2022.
^ "Melolontha melolontha (cockchafer or May bug)". Natural History Museum. Archived from the original on 3 July 2015. Retrieved 12 July 2015.
^ Gullan, Penny; Kosztarab, Michael (1997). "Adaptations in scale insects". Annual Review of Entomology. 42: 23–50. doi:10.1146/annurev.ento.42.1.23. PMID 15012306.
^ ^a ^b ^c Coulson, Robert N.; Witter, John A. (1984). Forest Entomology: Ecology and Management. John Wiley & Sons. pp. 71–72. ISBN 978-0-471-02573-3.
^ ^a ^b Kevin P. Johnson; Christopher H. Dietrich; Frank Friedrich; Rolf G. Beutel; Benjamin Wipfler; et al. (December 2018). "Phylogenomics and the evolution of hemipteroid insects". Proceedings of the National Academy of Sciences. 115 (50): 12775–12780. Bibcode:2018PNAS..11512775J. doi:10.1073/pnas.1815820115. PMC 6294958. PMID 30478043.
^ André Nel; Patrick Roques; Patricia Nel; Alexander A. Prokin; Thierry Bourgoin; et al. (2013). "The earliest known holometabolous insects". Nature. 503 (7475): 257–261. Bibcode:2013Natur.503..257N. doi:10.1038/nature12629. PMID 24132233. S2CID 205235859.
^ Gillott, Cedric (2005). Entomology (3 ed.). Springer. p. 213.
^ Shcherbakov, D. E. (2000). "Permian faunas of Homoptera (Hemiptera) in relation to phytogeography and the Permo-Triassic crisis" (PDF). Paleontological Journal. 34 (3): S251–S267.
^ Song, Nan; Liang, Ai-Ping; Bu, Cui-Ping (2012). "A Molecular Phylogeny of Hemiptera Inferred from Mitochondrial Genome Sequences". PLOS ONE. 7 (11): e48778. Bibcode:2012PLoSO...748778S. doi:10.1371/journal.pone.0048778. PMC 3493603. PMID 23144967.
^ Forero, Dimitri (2008). "The Systematics of Hemiptera" (PDF). Revista Colombiana de Entomología. 34 (1): 1–21. doi:10.25100/socolen.v34i1.9244. S2CID 85830606. Archived from the original (PDF) on 13 January 2015.
^ Cryan, Jason C.; Urban, Julie M. (January 2012). "Higher-level phylogeny of the insect order Hemiptera: is Auchenorrhyncha really paraphyletic?". Systematic Entomology. 37 (1): 7–21. Bibcode:2012SysEn..37....7C. doi:10.1111/j.1365-3113.2011.00611.x. S2CID 86564882.
^ "Suborder Auchenorrhyncha". NCSU. Retrieved 12 July 2015.
^ Cassis, Gerasimos (2019). "True Bugs (Insecta: Hemiptera: Heteroptera): Evolution, Classification, Biodiversity and Biology". Reference Module in Life Sciences. doi:10.1016/B978-0-12-809633-8.20710-3. ISBN 9780128096338. S2CID 214379746.
^ "Sternorrhyncha". Amateur Entomologists' Society. Retrieved 13 July 2015.
^ Ruppert, Edward E.; Fox, Richard, S.; Barnes, Robert D. (2004). Invertebrate Zoology, 7th edition. Cengage Learning. pp. 728, 748. ISBN 978-81-315-0104-7.{{cite book}}: CS1 maint: multiple names: authors list (link)
^ ^a ^b ^c ^d ^e ^f Gullan, P.J.; Cranston, P.S. (2014). The Insects: An Outline of Entomology, 5th Edition. Wiley. pp. 80–81, 790–. ISBN 978-1-118-84616-2.
^ Wheeler, Alfred George (2001). Biology of the Plant Bugs (Hemiptera: Miridae): Pests, Predators, Opportunists. Cornell University Press. pp. 105–135. ISBN 978-0-8014-3827-1.
^ Chinery, Michael (1993). Insects of Britain and Northern Europe (3rd ed.). Collins. ISBN 978-0-00-219918-6.
^ Alford, David V. (2012). Pests of Ornamental Trees, Shrubs and Flowers: A Color Handbook. Academic Press. p. 12. ISBN 978-0-12-398515-6.
^ ^a ^b "Hemiptera". Discover Life. Retrieved 13 July 2015.
^ Virant-Doberlet, Meta; Cokl, Andrej (April 2004). "Vibrational Communication in Insects". Neotropical Entomology. 33 (2): 121–134. doi:10.1590/S1519-566X2004000200001.
^ Simmons, Peter; Young, David (1978). "The Tymbal Mechanism and Song Patterns of the Bladder Cicada, Cystosoma saundersii" (PDF). Journal of Experimental Biology. 76: 27–45. doi:10.1242/jeb.76.1.27. Archived from the original (PDF) on 24 December 2015. Retrieved 22 August 2015.
^ Wilcox, R. Stimson (1975). "Sound-producing Mechanisms of Bueno macrotibialis Hungerford (Hemiptera: Notonectidae)" (PDF). International Journal of Insect Morphology & Embryology. 4 (21): 169–182. doi:10.1016/0020-7322(75)90015-x. hdl:2027.42/22083.
^ Gooderham, John; Tsyrlin, Edward (2002). The Waterbug Book: A Guide to the Freshwater Macroinvertebrates of Temperate Australia. Csiro Publishing. p. 146. ISBN 978-0-643-06668-7.
^ Britton, David (9 July 2009). "Metamorphosis: a remarkable change". Australian Museum. Retrieved 13 July 2015.
^ Mackean, D.G. "Aphids, an Introduction". Biology Teaching Resources. Retrieved 13 July 2015.
^ ^a ^b ^c Jon Martin; Mick Webb. "Hemiptera...It's a Bug's Life" (PDF). Natural History Museum. Retrieved 26 July 2010.
^ ^a ^b Bush, John W. M.; Hu, David L. (2006). "Walking on Water: Biolocomotion at the Interface". Annual Review of Fluid Mechanics. 38 (1): 339–369. Bibcode:2006AnRFM..38..339B. doi:10.1146/annurev.fluid.38.050304.092157.
^ Loxdale, H. D.; Hardie, J.; Halbert, S.; Foottit, R.; Kidd, N. A. C.; Carter, C. I. (1993). "The relative importance of short- and long-range movement of flying aphids" (PDF). Biological Reviews. 68 (2): 291–311. doi:10.1111/j.1469-185X.1993.tb00998.x. S2CID 86569822.
^ Waloff, Nadia (1983). "Absence of wing polymorphism in the arboreal, phytophagous species of some taxa of temperate Hemiptera: An hypothesis". Ecological Entomology. 8 (2): 229–232. Bibcode:1983EcoEn...8..229W. doi:10.1111/j.1365-2311.1983.tb00502.x. S2CID 83833159.
^ Burrows, M. (March 2013). "Jumping mechanisms of treehopper insects (Hemiptera, Auchenorrhyncha, Membracidae)". Journal of Experimental Biology. 216 (5): 788–799. doi:10.1242/jeb.078741. PMID 23155084.
^ Gorb, S. N. (July 2004). "The jumping mechanism of cicada Cercopis vulnerata (Auchenorrhyncha, Cercopidae): skeleton-muscle organisation, frictional surfaces, and inverse-kinematic model of leg movements". Arthropod Structure & Development. 33 (3): 201–220. Bibcode:2004ArtSD..33..201G. doi:10.1016/j.asd.2004.05.008. PMID 18089035.
^ ^a ^b Grimaldi, David; Engel, Michael S. (16 May 2005). Evolution of the Insects. Cambridge University Press. p. 289. ISBN 978-0-521-82149-0.
^ Labandeira, Conrad C. (May 1998). "Early History of Arthropod and Vascular Plant Associations". Annual Review of Earth and Planetary Sciences. 26: 329–377. Bibcode:1998AREPS..26..329L. doi:10.1146/annurev.earth.26.1.329.
^ Verne, Nole C. (2007). Forest Ecology Research Horizons. Nova Publishers. p. 210. ISBN 978-1-60021-490-5.
^ Dixon, A.F.G. (2012). Aphid Ecology: An optimization approach. Springer Science & Business Media. p. 128. ISBN 978-94-011-5868-8.
^ Tonkyn, David W.; Whitcomb, Robert F. (1987). "Feeding Strategies and the Guild Concept Among Vascular Feeding Insects and Microorganisms". In Harris, Kerry F. (ed.). Current Topics in Vector Research. pp. 179–199. doi:10.1007/978-1-4612-4712-8_6. ISBN 978-1-4612-9126-8.
^ Pompon, Julien; Quiring, Dan; Goyer, Claudia; Giordanengo, Philippe; Pelletier, Yvan (2011). "A phloem-sap feeder mixes phloem and xylem sap to regulate osmotic potential" (PDF). Journal of Insect Physiology. 57 (9): 1317–1322. Bibcode:2011JInsP..57.1317P. doi:10.1016/j.jinsphys.2011.06.007. PMID 21726563. Archived from the original (PDF) on 23 September 2015. Retrieved 22 August 2015.
^ Den, Francesca F.; Stewart, Alan J. A.; Gibson, Amos; Weiblen, George D.; Novotny, Vojtech (2013). "Low host specificity in species-rich assemblages of xylem- and phloem-feeding herbivores (Auchenorrhyncha) in a New Guinea lowland rain forest" (PDF). Journal of Tropical Ecology. 29 (6): 467–476. doi:10.1017/S0266467413000540. S2CID 12101409.
^ Novotny, Vojtech; Wilson, Michael R. (1997). "Why are there no small species among xylem-sucking insects?". Evolutionary Ecology. 11 (4): 419–437. Bibcode:1997EvEco..11..419N. doi:10.1023/a:1018432807165. S2CID 39222473.
^ Douglas, A.E. (2006). "Phloem-sap feeding by animals: problems and solutions" (PDF). Journal of Experimental Botany. 57 (4): 747–754. doi:10.1093/jxb/erj067. PMID 16449374.
^ Baje, Leontine; Stewart, A.J.A.; Novotny, Vojtech (2014). "Mesophyll cell-sucking herbivores (Cicadellidae: Typhlocybinae) on rainforest trees in Papua New Guinea: local and regional diversity of a taxonomically unexplored guild Ecological Entomology". Ecological Entomology. 39 (3): 325–333. doi:10.1111/een.12104. S2CID 55083254.
^ Panizzi, Antônio Ricardo; Parra, José R.P. (2012). Insect Bioecology and Nutrition for Integrated Pest Management. CRC Press. p. 108. ISBN 978-1-4398-3708-5.
^ Pompon, Julien; Quiring, Dan; Goyer, Claudia; Giordanengo, Philippe; Pelletier, Yvan (2011). "A phloem-sap feeder mixes phloem and xylem sap to regulate osmotic potential" (PDF). Journal of Insect Physiology. 57 (9): 1317–1322. Bibcode:2011JInsP..57.1317P. doi:10.1016/j.jinsphys.2011.06.007. PMID 21726563. Archived from the original (PDF) on 23 September 2015. Retrieved 22 August 2015.
^ "Digestive System". Entomology. University of Sydney. Archived from the original on 7 October 2012. Retrieved 22 August 2015.
^ Underwood, D. L. A. "Hemipteroids" (PDF). California State University. Archived from the original (PDF) on 4 March 2016. Retrieved 22 August 2015.
^ Hall, Donald W.; Butler, Jerry F.; Cromroy, Harvey L. (1 September 2014). "Hackberry petiole gall psyllid". Featured Creatures. University of Florida/IFAS. Retrieved 12 August 2015.
^ "Glossy Shield Bug: Cermatulus nasalis". Brisbane Insects and Spiders. 2010. Retrieved 16 July 2015.
^ Cohen, Allen C. (1990). "Feeding Adaptations of Some Predaceous Hemiptera" (PDF). Annals of the Entomological Society of America. 83 (6): 1215–1223. doi:10.1093/aesa/83.6.1215.
^ Goddard, Jerome (2009). "Bed Bugs (Cimex lectularius) and Clinical Consequences of Their Bites". JAMA. 301 (13): 1358–1366. doi:10.1001/jama.2009.405. PMID 19336711.
^ Poinar, G. (2005). "Triatoma dominicana sp. n. (Hemiptera: Reduviidae: Triatominae), and Trypanosoma antiquus sp. n. (Stercoraria: Trypanosomatidae), the First Fossil Evidence of a Triatomine-Trypanosomatid Vector Association". Vector-Borne and Zoonotic Diseases. 5 (1): 72–81. doi:10.1089/vbz.2005.5.72. PMID 15815152.
^ Ivens, A. B. F.; Kronauer, D. J. C.; Pen, I.; Weissing, F. J.; Boomsma, J. J. (2012). "Reproduction and dispersal in an ant-associated root aphid community". Molecular Ecology. 21 (17): 4257–69. Bibcode:2012MolEc..21.4257I. doi:10.1111/j.1365-294X.2012.05701.x. PMID 22804757. S2CID 34429056.
^ Choe, Jae (14 February 2012). Secret Lives of Ants. JHU Press. pp. 62–64. ISBN 978-1-4214-0521-6.
^ Evans, Edward W. (November 2009). "Lady beetles as predators of insects other than Hemiptera". Biological Control. 51 (2): 255–267. Bibcode:2009BiolC..51..255E. doi:10.1016/j.biocontrol.2009.05.011.
^ ^a ^b Edmunds, M. (1974). Defence in Animals. Longman. pp. 172, 191–194, 201, 261 and passim. ISBN 978-0-582-44132-3.
^ Wierauch C. (2006). "Anatomy of Disguise: Camouflaging Structures in Nymphs of Some Reduviidae (Heteroptera)". Am. Mus. Novit. 3542: 1–18. doi:10.1206/0003-0082(2006)3542[1:aodcsi]2.0.co;2. hdl:2246/5820. S2CID 7894145.
^ Cott, Hugh B. (1940). Adaptive Coloration in Animals. Methuen. pp. 375–376.
^ "Better IDs for E.A. Seguy Cicada Illustrations". Retrieved 24 August 2015.
^ Wheeler, Alfred George (2001). Biology of the Plant Bugs (Hemiptera: Miridae): Pests, Predators, Opportunists. Cornell University Press. pp. 100ff. ISBN 978-0-8014-3827-1.
^ Exnerova, Alice; et al. (April 2003). "Reactions of passerine birds to aposematic and non-aposematic firebugs (Pyrrhocoris apterus ; Heteroptera)". Biological Journal of the Linnean Society. 78 (4): 517–525. doi:10.1046/j.0024-4066.2002.00161.x.
^ Lucchi, Andrea; Mazzoni, Emanuele (2004). "Wax Production in Adults of Planthoppers (Homoptera: Fulgoroidea) with Particular Reference to Metcalfa pruinosa (Flatidae)" (PDF). Annals of the Entomological Society of America. 97 (6): 1294–1298. doi:10.1603/0013-8746(2004)097[1294:wpiaop]2.0.co;2.
^ Rakitov, Roman A. (2002). "Structure and Function of the Malpighian Tubules, and Related Behaviors in Juvenile Cicadas: Evidence of Homology with Spittlebugs (Hemiptera: Cicadoidea & Cercopoidea)". Zoologischer Anzeiger. 241 (2): 117–130. Bibcode:2002ZooAn.241..117R. doi:10.1078/0044-5231-00025.
^ Tallamy, Douglas W.; Schaefer, Carl (1997). "Maternal care in the Hemiptera: ancestry, alternatives, and current adaptive value". In Jae C. Choe; Bernard J. Crespi (eds.). The Evolution of Social Behaviour in Insects and Arachnids. Cambridge University Press. pp. 91–115.
^ Crowl, T.A.; Alexander Jr., J.E. (1989). "Parental care and foraging ability in male water bugs (Belostoma flumineum)". Canadian Journal of Zoology. 67 (2): 513–515. doi:10.1139/z89-074.
^ "Insect groups (Orders)". Amateur Entomologists' Society. Retrieved 16 July 2015.
^ "Sooty moulds". Royal Horticultural Society. Retrieved 16 July 2015.
^ David L. Green (10 August 2003). "Cottony cushion scale: The pest that launched a revolution in pest control methods". Retrieved 16 July 2015.
^ Capinera, John L. (October 2005). "Featured creatures". University of Florida website – Department of Entomology and Nematology. University of Florida. Retrieved 16 July 2015.
^ Mahr, Susan (1997). "Know Your Friends: Damsel Bugs". Biological Control News. IV (2). Archived from the original on 20 April 2018. Retrieved 16 July 2015.
^ James Hagler. Weeden, Catherine R.; Shelton, Anthony M.; Hoffman, Michael P. (eds.). "Geocoris spp. (Heteroptera: Lygaeidae): Bigeyed Bug". Biological Control: A Guide to Natural Enemies in North America. Cornell University. Retrieved 16 July 2015.
^ Torres, Jorge Braz; Boyd, David W. (2009). "Zoophytophagy in predatory Hemiptera". Brazilian Archives of Biology and Technology. 52 (5): 1199–1208. doi:10.1590/S1516-89132009000500018.
^ Coll, M.; Ruberson, J.R., eds. (1998). Predatory Heteroptera: their ecology and use in biological control. Entomological Society of America. ISBN 978-0-938522-62-1.^{[permanent dead link]}
^ FDA Color Additives,"Guidance for Industry: Cochineal Extract and Carmine: Declaration by Name on the Label of All Foods and Cosmetic Products That Contain These Color Additives; Small Entity Compliance Guide". www.fda.gov. Retrieved 2016-02-22.
^ "Cochineal and Carmine". Major colourants and dyestuffs, mainly produced in horticultural systems. FAO. Retrieved 16 June 2015.
^ "How Shellac Is Manufactured". The Mail (Adelaide, SA : 1912 – 1954). 18 December 1937. Retrieved 17 July 2015.
^ Pearnchob, N.; Siepmann, J.; Bodmeier, R. (2003). "Pharmaceutical applications of shellac: moisture-protective and taste-masking coatings and extended-release matrix tablets". Drug Development and Industrial Pharmacy. 29 (8): 925–938. doi:10.1081/ddc-120024188. PMID 14570313. S2CID 13150932.
^ "American Trypanosomiasis". Centers for Disease Control (CDC). Retrieved 17 July 2015.
^ Goddard, J; deShazo, R (1 April 2009). "Bed bugs (Cimex lectularius) and clinical consequences of their bites". JAMA: The Journal of the American Medical Association. 301 (13): 1358–66. doi:10.1001/jama.2009.405. PMID 19336711.
^ ^a ^b Reinhardt, Klaus; Siva-Jothy, Michael T. (January 2007). "Biology of the Bed Bugs (Cimicidae)" (PDF). Annual Review of Entomology. 52: 351–374. doi:10.1146/annurev.ento.52.040306.133913. PMID 16968204. Archived from the original (PDF) on 5 July 2010. Retrieved 26 May 2010.
^ Carayon, J. 1959 Insémination par "spermalège" et cordon conducteur de spermatozoids chez Stricticimex brevispinosus Usinger (Heteroptera, Cimicidae). Rev. Zool. Bot. Afr. 60, 81–104.
^ Greenaway, Twilight. "The Best Way to Handle the Coming Cicada Invasion? Heat Up the Deep Fryer". Smithsonian. Retrieved 12 July 2015.
^ Anthes, Emily (14 October 2014). "Could insects be the wonder food of the future?". BBC. Retrieved 17 July 2015.
^ "Future Prospects for Food and Feed Security". Archived from the original on 4 April 2016. Retrieved 13 April 2016.
^ Myers, J. G. (1929). Insect Singers (PDF). G. Routledge and Sons.
^ "Cicada". Britannica. Retrieved 12 July 2015.
^ Cunha, Erlane Jose; Montag, Luciano; Juen, Leandro (2015). "Oil palm crops effects on environmental integrity of Amazonian streams and Heteropteran (Hemiptera) species diversity". Ecological Indicators. 52: 422–429. Bibcode:2015EcInd..52..422C. doi:10.1016/j.ecolind.2014.12.024.
^ Baker, Mitchell B.; Venugopal, P. Dilip; Lamp, William O. (2015). "Climate Change and Phenology: Empoasca fabae (Hemiptera: Cicadellidae) Migration and Severity of Impact". PLOS ONE. 10 (5): e0124915. Bibcode:2015PLoSO..1024915B. doi:10.1371/journal.pone.0124915. PMC 4430490. PMID 25970705.

External links

Cladogram of Hemiptera at Tree of Life Project

[9] The Museum of New Zealand notes that "in everyday conversation", bug "refers to land arthropods with at least six legs, such as insects, spiders, and centipedes".^[6] In a chapter on "Bugs That Are Not Insects", the entomologist Gilbert Walbauer specifies centipedes, millipedes, arachnids (spiders, daddy longlegs, scorpions, mites, chiggers and ticks) as well as the few terrestrial crustaceans (sowbugs and pillbugs),^[7] but argues that "including legless creatures such as worms, slugs, and snails among the bugs stretches the word too much".^[5] "Bug" is used for a harmful microorganism, typically a bacterium.^[8]

[68] The green/red flash coloration earns the insect the name of stop and go cicada.^[66]

[Wang2016-1] Wang, Yan-hui; Engel, Michael S.; Rafael, José A.; Wu, Hao-yang; Rédei, Dávid; et al. (2016). "Fossil record of stem groups employed in evaluating the chronogram of insects (Arthropoda: Hexapoda)". Scientific Reports. 6: 38939. Bibcode:2016NatSR...638939W. doi:10.1038/srep38939. PMC 5154178. PMID 27958352.

[2] "Hemiptera". Integrated Taxonomic Information System.

[3] "Hemiptera: bugs, aphids and cicadas". Commonwealth Scientific and Industrial Research Organisation. Retrieved 8 May 2007.

[4] "Suborder Heteroptera – True Bugs". Bug guide. Iowa State University Entomology. n.d.

[Walbauer1-5] Gilbert Waldbauer. The Handy Bug Answer Book. Visible Ink, 1998. p. 1. ISBN 9781578590490

[6] "What is a bug? Insects, arachnids, and myriapods" at Museum of New Zealand Te Papa Tongarewa website. Accessed 10 March 2022.

[Walbauer5-7] Gilbert Waldbauer. The Handy Bug Answer Book. Visible Ink, 1998. pp. 5–26. ISBN 9781578590490

[8] "BUG | Meaning & Definition for UK English". Lexico.com. Archived from the original on 11 December 2019. Retrieved 3 August 2022.

[denmark-10] Denmark, Harold; Mead, Frank; Fasulo, Thomas (April 2010). "Lovebug, Plecia nearctica Hardy". Featured Creatures. University of Florida/IFAS. Retrieved 10 March 2022.

[11] "Melolontha melolontha (cockchafer or May bug)". Natural History Museum. Archived from the original on 3 July 2015. Retrieved 12 July 2015.

[Gullan1997-12] Gullan, Penny; Kosztarab, Michael (1997). "Adaptations in scale insects". Annual Review of Entomology. 42: 23–50. doi:10.1146/annurev.ento.42.1.23. PMID 15012306.

[Coulson-13] Coulson, Robert N.; Witter, John A. (1984). Forest Entomology: Ecology and Management. John Wiley & Sons. pp. 71–72. ISBN 978-0-471-02573-3.

[Johnson-14] Kevin P. Johnson; Christopher H. Dietrich; Frank Friedrich; Rolf G. Beutel; Benjamin Wipfler; et al. (December 2018). "Phylogenomics and the evolution of hemipteroid insects". Proceedings of the National Academy of Sciences. 115 (50): 12775–12780. Bibcode:2018PNAS..11512775J. doi:10.1073/pnas.1815820115. PMC 6294958. PMID 30478043.

[natureholometabolousinsects-15] André Nel; Patrick Roques; Patricia Nel; Alexander A. Prokin; Thierry Bourgoin; et al. (2013). "The earliest known holometabolous insects". Nature. 503 (7475): 257–261. Bibcode:2013Natur.503..257N. doi:10.1038/nature12629. PMID 24132233. S2CID 205235859.

[16] Gillott, Cedric (2005). Entomology (3 ed.). Springer. p. 213.

[17] Shcherbakov, D. E. (2000). "Permian faunas of Homoptera (Hemiptera) in relation to phytogeography and the Permo-Triassic crisis" (PDF). Paleontological Journal. 34 (3): S251–S267.

[Song2012-18] Song, Nan; Liang, Ai-Ping; Bu, Cui-Ping (2012). "A Molecular Phylogeny of Hemiptera Inferred from Mitochondrial Genome Sequences". PLOS ONE. 7 (11): e48778. Bibcode:2012PLoSO...748778S. doi:10.1371/journal.pone.0048778. PMC 3493603. PMID 23144967.

[Forero-19] Forero, Dimitri (2008). "The Systematics of Hemiptera" (PDF). Revista Colombiana de Entomología. 34 (1): 1–21. doi:10.25100/socolen.v34i1.9244. S2CID 85830606. Archived from the original (PDF) on 13 January 2015.

[20] Cryan, Jason C.; Urban, Julie M. (January 2012). "Higher-level phylogeny of the insect order Hemiptera: is Auchenorrhyncha really paraphyletic?". Systematic Entomology. 37 (1): 7–21. Bibcode:2012SysEn..37....7C. doi:10.1111/j.1365-3113.2011.00611.x. S2CID 86564882.

[21] "Suborder Auchenorrhyncha". NCSU. Retrieved 12 July 2015.

[Cassis2019-22] Cassis, Gerasimos (2019). "True Bugs (Insecta: Hemiptera: Heteroptera): Evolution, Classification, Biodiversity and Biology". Reference Module in Life Sciences. doi:10.1016/B978-0-12-809633-8.20710-3. ISBN 9780128096338. S2CID 214379746.

[23] "Sternorrhyncha". Amateur Entomologists' Society. Retrieved 13 July 2015.

[Ruppert-24] Ruppert, Edward E.; Fox, Richard, S.; Barnes, Robert D. (2004). Invertebrate Zoology, 7th edition. Cengage Learning. pp. 728, 748. ISBN 978-81-315-0104-7.{{cite book}}: CS1 maint: multiple names: authors list (link)

[Gullan-25] ^ ^a ^b ^c ^d ^e ^f Gullan, P.J.; Cranston, P.S. (2014). The Insects: An Outline of Entomology, 5th Edition. Wiley. pp. 80–81, 790–. ISBN 978-1-118-84616-2.

[Wheeler2001-26] Wheeler, Alfred George (2001). Biology of the Plant Bugs (Hemiptera: Miridae): Pests, Predators, Opportunists. Cornell University Press. pp. 105–135. ISBN 978-0-8014-3827-1.

[Chinery-27] Chinery, Michael (1993). Insects of Britain and Northern Europe (3rd ed.). Collins. ISBN 978-0-00-219918-6.

[Alford-28] Alford, David V. (2012). Pests of Ornamental Trees, Shrubs and Flowers: A Color Handbook. Academic Press. p. 12. ISBN 978-0-12-398515-6.

[Discover-29] "Hemiptera". Discover Life. Retrieved 13 July 2015.

[30] Virant-Doberlet, Meta; Cokl, Andrej (April 2004). "Vibrational Communication in Insects". Neotropical Entomology. 33 (2): 121–134. doi:10.1590/S1519-566X2004000200001.

[31] Simmons, Peter; Young, David (1978). "The Tymbal Mechanism and Song Patterns of the Bladder Cicada, Cystosoma saundersii" (PDF). Journal of Experimental Biology. 76: 27–45. doi:10.1242/jeb.76.1.27. Archived from the original (PDF) on 24 December 2015. Retrieved 22 August 2015.

[32] Wilcox, R. Stimson (1975). "Sound-producing Mechanisms of Bueno macrotibialis Hungerford (Hemiptera: Notonectidae)" (PDF). International Journal of Insect Morphology & Embryology. 4 (21): 169–182. doi:10.1016/0020-7322(75)90015-x. hdl:2027.42/22083.

[33] Gooderham, John; Tsyrlin, Edward (2002). The Waterbug Book: A Guide to the Freshwater Macroinvertebrates of Temperate Australia. Csiro Publishing. p. 146. ISBN 978-0-643-06668-7.

[34] Britton, David (9 July 2009). "Metamorphosis: a remarkable change". Australian Museum. Retrieved 13 July 2015.

[35] Mackean, D.G. "Aphids, an Introduction". Biology Teaching Resources. Retrieved 13 July 2015.

[NHM-36] Jon Martin; Mick Webb. "Hemiptera...It's a Bug's Life" (PDF). Natural History Museum. Retrieved 26 July 2010.

[Bush-37] Bush, John W. M.; Hu, David L. (2006). "Walking on Water: Biolocomotion at the Interface". Annual Review of Fluid Mechanics. 38 (1): 339–369. Bibcode:2006AnRFM..38..339B. doi:10.1146/annurev.fluid.38.050304.092157.

[38] Loxdale, H. D.; Hardie, J.; Halbert, S.; Foottit, R.; Kidd, N. A. C.; Carter, C. I. (1993). "The relative importance of short- and long-range movement of flying aphids" (PDF). Biological Reviews. 68 (2): 291–311. doi:10.1111/j.1469-185X.1993.tb00998.x. S2CID 86569822.

[39] Waloff, Nadia (1983). "Absence of wing polymorphism in the arboreal, phytophagous species of some taxa of temperate Hemiptera: An hypothesis". Ecological Entomology. 8 (2): 229–232. Bibcode:1983EcoEn...8..229W. doi:10.1111/j.1365-2311.1983.tb00502.x. S2CID 83833159.

[40] Burrows, M. (March 2013). "Jumping mechanisms of treehopper insects (Hemiptera, Auchenorrhyncha, Membracidae)". Journal of Experimental Biology. 216 (5): 788–799. doi:10.1242/jeb.078741. PMID 23155084.

[41] Gorb, S. N. (July 2004). "The jumping mechanism of cicada Cercopis vulnerata (Auchenorrhyncha, Cercopidae): skeleton-muscle organisation, frictional surfaces, and inverse-kinematic model of leg movements". Arthropod Structure & Development. 33 (3): 201–220. Bibcode:2004ArtSD..33..201G. doi:10.1016/j.asd.2004.05.008. PMID 18089035.

[GrimaldiEngel2005-42] Grimaldi, David; Engel, Michael S. (16 May 2005). Evolution of the Insects. Cambridge University Press. p. 289. ISBN 978-0-521-82149-0.

[43] Labandeira, Conrad C. (May 1998). "Early History of Arthropod and Vascular Plant Associations". Annual Review of Earth and Planetary Sciences. 26: 329–377. Bibcode:1998AREPS..26..329L. doi:10.1146/annurev.earth.26.1.329.

[Verne2007-44] Verne, Nole C. (2007). Forest Ecology Research Horizons. Nova Publishers. p. 210. ISBN 978-1-60021-490-5.

[Dixon-45] Dixon, A.F.G. (2012). Aphid Ecology: An optimization approach. Springer Science & Business Media. p. 128. ISBN 978-94-011-5868-8.

[46] Tonkyn, David W.; Whitcomb, Robert F. (1987). "Feeding Strategies and the Guild Concept Among Vascular Feeding Insects and Microorganisms". In Harris, Kerry F. (ed.). Current Topics in Vector Research. pp. 179–199. doi:10.1007/978-1-4612-4712-8_6. ISBN 978-1-4612-9126-8.

[47] Pompon, Julien; Quiring, Dan; Goyer, Claudia; Giordanengo, Philippe; Pelletier, Yvan (2011). "A phloem-sap feeder mixes phloem and xylem sap to regulate osmotic potential" (PDF). Journal of Insect Physiology. 57 (9): 1317–1322. Bibcode:2011JInsP..57.1317P. doi:10.1016/j.jinsphys.2011.06.007. PMID 21726563. Archived from the original (PDF) on 23 September 2015. Retrieved 22 August 2015.

[48] Den, Francesca F.; Stewart, Alan J. A.; Gibson, Amos; Weiblen, George D.; Novotny, Vojtech (2013). "Low host specificity in species-rich assemblages of xylem- and phloem-feeding herbivores (Auchenorrhyncha) in a New Guinea lowland rain forest" (PDF). Journal of Tropical Ecology. 29 (6): 467–476. doi:10.1017/S0266467413000540. S2CID 12101409.

[49] Novotny, Vojtech; Wilson, Michael R. (1997). "Why are there no small species among xylem-sucking insects?". Evolutionary Ecology. 11 (4): 419–437. Bibcode:1997EvEco..11..419N. doi:10.1023/a:1018432807165. S2CID 39222473.

[50] Douglas, A.E. (2006). "Phloem-sap feeding by animals: problems and solutions" (PDF). Journal of Experimental Botany. 57 (4): 747–754. doi:10.1093/jxb/erj067. PMID 16449374.

[51] Baje, Leontine; Stewart, A.J.A.; Novotny, Vojtech (2014). "Mesophyll cell-sucking herbivores (Cicadellidae: Typhlocybinae) on rainforest trees in Papua New Guinea: local and regional diversity of a taxonomically unexplored guild Ecological Entomology". Ecological Entomology. 39 (3): 325–333. doi:10.1111/een.12104. S2CID 55083254.

[Panizzi-52] Panizzi, Antônio Ricardo; Parra, José R.P. (2012). Insect Bioecology and Nutrition for Integrated Pest Management. CRC Press. p. 108. ISBN 978-1-4398-3708-5.

[53] Pompon, Julien; Quiring, Dan; Goyer, Claudia; Giordanengo, Philippe; Pelletier, Yvan (2011). "A phloem-sap feeder mixes phloem and xylem sap to regulate osmotic potential" (PDF). Journal of Insect Physiology. 57 (9): 1317–1322. Bibcode:2011JInsP..57.1317P. doi:10.1016/j.jinsphys.2011.06.007. PMID 21726563. Archived from the original (PDF) on 23 September 2015. Retrieved 22 August 2015.

[54] "Digestive System". Entomology. University of Sydney. Archived from the original on 7 October 2012. Retrieved 22 August 2015.

[55] Underwood, D. L. A. "Hemipteroids" (PDF). California State University. Archived from the original (PDF) on 4 March 2016. Retrieved 22 August 2015.

[56] Hall, Donald W.; Butler, Jerry F.; Cromroy, Harvey L. (1 September 2014). "Hackberry petiole gall psyllid". Featured Creatures. University of Florida/IFAS. Retrieved 12 August 2015.

[Brisbane-57] "Glossy Shield Bug: Cermatulus nasalis". Brisbane Insects and Spiders. 2010. Retrieved 16 July 2015.

[58] Cohen, Allen C. (1990). "Feeding Adaptations of Some Predaceous Hemiptera" (PDF). Annals of the Entomological Society of America. 83 (6): 1215–1223. doi:10.1093/aesa/83.6.1215.

[59] Goddard, Jerome (2009). "Bed Bugs (Cimex lectularius) and Clinical Consequences of Their Bites". JAMA. 301 (13): 1358–1366. doi:10.1001/jama.2009.405. PMID 19336711.

[Poinar-60] Poinar, G. (2005). "Triatoma dominicana sp. n. (Hemiptera: Reduviidae: Triatominae), and Trypanosoma antiquus sp. n. (Stercoraria: Trypanosomatidae), the First Fossil Evidence of a Triatomine-Trypanosomatid Vector Association". Vector-Borne and Zoonotic Diseases. 5 (1): 72–81. doi:10.1089/vbz.2005.5.72. PMID 15815152.

[Ivens-61] Ivens, A. B. F.; Kronauer, D. J. C.; Pen, I.; Weissing, F. J.; Boomsma, J. J. (2012). "Reproduction and dispersal in an ant-associated root aphid community". Molecular Ecology. 21 (17): 4257–69. Bibcode:2012MolEc..21.4257I. doi:10.1111/j.1365-294X.2012.05701.x. PMID 22804757. S2CID 34429056.

[Choe2012-62] Choe, Jae (14 February 2012). Secret Lives of Ants. JHU Press. pp. 62–64. ISBN 978-1-4214-0521-6.

[63] Evans, Edward W. (November 2009). "Lady beetles as predators of insects other than Hemiptera". Biological Control. 51 (2): 255–267. Bibcode:2009BiolC..51..255E. doi:10.1016/j.biocontrol.2009.05.011.

[Edmunds-64] Edmunds, M. (1974). Defence in Animals. Longman. pp. 172, 191–194, 201, 261 and passim. ISBN 978-0-582-44132-3.

[65] Wierauch C. (2006). "Anatomy of Disguise: Camouflaging Structures in Nymphs of Some Reduviidae (Heteroptera)". Am. Mus. Novit. 3542: 1–18. doi:10.1206/0003-0082(2006)3542[1:aodcsi]2.0.co;2. hdl:2246/5820. S2CID 7894145.

[66] Cott, Hugh B. (1940). Adaptive Coloration in Animals. Methuen. pp. 375–376.

[67] "Better IDs for E.A. Seguy Cicada Illustrations". Retrieved 24 August 2015.

[Wheeler100-69] Wheeler, Alfred George (2001). Biology of the Plant Bugs (Hemiptera: Miridae): Pests, Predators, Opportunists. Cornell University Press. pp. 100ff. ISBN 978-0-8014-3827-1.

[70] Exnerova, Alice; et al. (April 2003). "Reactions of passerine birds to aposematic and non-aposematic firebugs (Pyrrhocoris apterus ; Heteroptera)". Biological Journal of the Linnean Society. 78 (4): 517–525. doi:10.1046/j.0024-4066.2002.00161.x.

[71] Lucchi, Andrea; Mazzoni, Emanuele (2004). "Wax Production in Adults of Planthoppers (Homoptera: Fulgoroidea) with Particular Reference to Metcalfa pruinosa (Flatidae)" (PDF). Annals of the Entomological Society of America. 97 (6): 1294–1298. doi:10.1603/0013-8746(2004)097[1294:wpiaop]2.0.co;2.

[72] Rakitov, Roman A. (2002). "Structure and Function of the Malpighian Tubules, and Related Behaviors in Juvenile Cicadas: Evidence of Homology with Spittlebugs (Hemiptera: Cicadoidea & Cercopoidea)". Zoologischer Anzeiger. 241 (2): 117–130. Bibcode:2002ZooAn.241..117R. doi:10.1078/0044-5231-00025.

[73] Tallamy, Douglas W.; Schaefer, Carl (1997). "Maternal care in the Hemiptera: ancestry, alternatives, and current adaptive value". In Jae C. Choe; Bernard J. Crespi (eds.). The Evolution of Social Behaviour in Insects and Arachnids. Cambridge University Press. pp. 91–115.

[74] Crowl, T.A.; Alexander Jr., J.E. (1989). "Parental care and foraging ability in male water bugs (Belostoma flumineum)". Canadian Journal of Zoology. 67 (2): 513–515. doi:10.1139/z89-074.

[AES-75] "Insect groups (Orders)". Amateur Entomologists' Society. Retrieved 16 July 2015.

[76] "Sooty moulds". Royal Horticultural Society. Retrieved 16 July 2015.

[77] David L. Green (10 August 2003). "Cottony cushion scale: The pest that launched a revolution in pest control methods". Retrieved 16 July 2015.

[78] Capinera, John L. (October 2005). "Featured creatures". University of Florida website – Department of Entomology and Nematology. University of Florida. Retrieved 16 July 2015.

[79] Mahr, Susan (1997). "Know Your Friends: Damsel Bugs". Biological Control News. IV (2). Archived from the original on 20 April 2018. Retrieved 16 July 2015.

[80] James Hagler. Weeden, Catherine R.; Shelton, Anthony M.; Hoffman, Michael P. (eds.). "Geocoris spp. (Heteroptera: Lygaeidae): Bigeyed Bug". Biological Control: A Guide to Natural Enemies in North America. Cornell University. Retrieved 16 July 2015.

[81] Torres, Jorge Braz; Boyd, David W. (2009). "Zoophytophagy in predatory Hemiptera". Brazilian Archives of Biology and Technology. 52 (5): 1199–1208. doi:10.1590/S1516-89132009000500018.

[82] Coll, M.; Ruberson, J.R., eds. (1998). Predatory Heteroptera: their ecology and use in biological control. Entomological Society of America. ISBN 978-0-938522-62-1.^{[permanent dead link]}

[83] FDA Color Additives,"Guidance for Industry: Cochineal Extract and Carmine: Declaration by Name on the Label of All Foods and Cosmetic Products That Contain These Color Additives; Small Entity Compliance Guide". www.fda.gov. Retrieved 2016-02-22.

[84] "Cochineal and Carmine". Major colourants and dyestuffs, mainly produced in horticultural systems. FAO. Retrieved 16 June 2015.

[85] "How Shellac Is Manufactured". The Mail (Adelaide, SA : 1912 – 1954). 18 December 1937. Retrieved 17 July 2015.

[86] Pearnchob, N.; Siepmann, J.; Bodmeier, R. (2003). "Pharmaceutical applications of shellac: moisture-protective and taste-masking coatings and extended-release matrix tablets". Drug Development and Industrial Pharmacy. 29 (8): 925–938. doi:10.1081/ddc-120024188. PMID 14570313. S2CID 13150932.

[CDC-87] "American Trypanosomiasis". Centers for Disease Control (CDC). Retrieved 17 July 2015.

[NEJM2009-88] Goddard, J; deShazo, R (1 April 2009). "Bed bugs (Cimex lectularius) and clinical consequences of their bites". JAMA: The Journal of the American Medical Association. 301 (13): 1358–66. doi:10.1001/jama.2009.405. PMID 19336711.

[bioofbedbug-89] Reinhardt, Klaus; Siva-Jothy, Michael T. (January 2007). "Biology of the Bed Bugs (Cimicidae)" (PDF). Annual Review of Entomology. 52: 351–374. doi:10.1146/annurev.ento.52.040306.133913. PMID 16968204. Archived from the original (PDF) on 5 July 2010. Retrieved 26 May 2010.

[carayon1959-90] Carayon, J. 1959 Insémination par "spermalège" et cordon conducteur de spermatozoids chez Stricticimex brevispinosus Usinger (Heteroptera, Cimicidae). Rev. Zool. Bot. Afr. 60, 81–104.

[91] Greenaway, Twilight. "The Best Way to Handle the Coming Cicada Invasion? Heat Up the Deep Fryer". Smithsonian. Retrieved 12 July 2015.

[92] Anthes, Emily (14 October 2014). "Could insects be the wonder food of the future?". BBC. Retrieved 17 July 2015.

[93] "Future Prospects for Food and Feed Security". Archived from the original on 4 April 2016. Retrieved 13 April 2016.

[94] Myers, J. G. (1929). Insect Singers (PDF). G. Routledge and Sons.

[95] "Cicada". Britannica. Retrieved 12 July 2015.

[96] Cunha, Erlane Jose; Montag, Luciano; Juen, Leandro (2015). "Oil palm crops effects on environmental integrity of Amazonian streams and Heteropteran (Hemiptera) species diversity". Ecological Indicators. 52: 422–429. Bibcode:2015EcInd..52..422C. doi:10.1016/j.ecolind.2014.12.024.

[97] Baker, Mitchell B.; Venugopal, P. Dilip; Lamp, William O. (2015). "Climate Change and Phenology: Empoasca fabae (Hemiptera: Cicadellidae) Migration and Severity of Impact". PLOS ONE. 10 (5): e0124915. Bibcode:2015PLoSO..1024915B. doi:10.1371/journal.pone.0124915. PMC 4430490. PMID 25970705.

[1]

[2]

[3]

[4]

[5]

[a]

[9]

[10]

[11]

[12]

[13]

[14]

[15]

[16]

[17]

[18]

[19]

[20]

[21]

[22]

[23]

[24]

[25]

[26]

[27]

[28]

[29]

[30]

[31]

[32]

[33]

[34]

[35]

[36]

[37]

[38]

[39]

[40]

[41]

[42]

[43]

[44]

[45]

[46]

[47]

[48]

[49]

[50]

[51]

[52]

[53]

[54]

[55]

[56]

[57]

[58]

[59]

[60]

[61]

[62]

[63]

[64]

[65]

[b]

[67]

[68]

[69]

[70]

[71]

[72]

[73]

[74]

[75]

[76]

[77]

[78]

[79]

[80]

[81]

[82]

[83]

[84]

[85]

[86]

[87]

[88]

[89]

[90]

[91]

[92]

[93]

[94]

[95]

[6]

[7]

[8]

[66]

@@ Line 12: / Line 12: @@
 | display_parents=2
 | taxon=Hemiptera
-| authority=[[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]]
+| authority=[[Linnaeus]], [[10th edition of Systema Naturae|1758]]
 | subdivision_ranks=Suborders
 | subdivision_ref=<ref>{{ITIS |id=103359 |taxon=Hemiptera}}</ref>
@@ Line 21: / Line 21: @@
 }}
-'''Hemiptera''' ({{IPAc-en|h|ɛ|ˈ|m|ɪ|p|t|ər|ə}}; {{etymology|Ancient Greek|hemipterus|half-winged}}) is an [[order (biology)|order]] of [[insect]]s, commonly called '''true bugs''', comprising over 80,000 [[species]] within groups such as the [[cicada]]s, [[aphid]]s, [[planthopper]]s, [[leafhopper]]s, [[Reduviidae|assassin bugs]], [[Cimex|bed bug]]s, and [[shield bug]]s. They range in size from {{cvt|1|mm|in|2}} to around {{cvt|15|cm|in|0}}, and share a common arrangement of piercing-sucking [[Insect mouthparts|mouthparts]].<ref>{{cite web |url=http://www.ento.csiro.au/education/insects/hemiptera.html |title=Hemiptera: bugs, aphids and cicadas |publisher=Commonwealth Scientific and Industrial Research Organisation |access-date=May 8, 2007}}</ref> The name "true bugs" is often limited to the suborder [[Heteroptera]].<ref>{{cite web |url=http://bugguide.net/node/view/94266 |title=Suborder Heteroptera – True Bugs |work= Bug guide |publisher=Iowa State University Entomology |date=n.d.}}</ref>
+'''Hemiptera''' ({{IPAc-en|h|ɛ|ˈ|m|ɪ|p|t|ər|ə}}; {{etymology|Ancient Greek|hemipterus|half-winged}}) is an [[order (biology)|order]] of [[insect]]s, commonly called '''true bugs''', comprising over 80,000 [[species]] within groups such as the [[cicada]]s, [[aphid]]s, [[planthopper]]s, [[leafhopper]]s, [[assassin bugs]], [[Cimex|bed bug]]s, and [[shield bug]]s. They range in size from {{cvt|1|mm|in|2}} to around {{cvt|15|cm|in|0}}, and share a common arrangement of piercing-sucking [[Insect mouthparts|mouthparts]].<ref>{{cite web |url=http://www.ento.csiro.au/education/insects/hemiptera.html |title=Hemiptera: bugs, aphids and cicadas |publisher=Commonwealth Scientific and Industrial Research Organisation |access-date=May 8, 2007}}</ref> The name "true bugs" is often limited to the suborder [[Heteroptera]].<ref>{{cite web |url=http://bugguide.net/node/view/94266 |title=Suborder Heteroptera – True Bugs |work= Bug guide |publisher=Iowa State University Entomology |date=n.d.}}</ref>
-Entomologists reserve the term ''bug'' for Hemiptera or Heteroptera,<ref name="Walbauer1">Gilbert Waldbauer. ''The Handy Bug Answer Book.'' Visible Ink, 1998. [https://archive.org/details/handybuganswerbo00wald/page/1/mode/2up p. 1.] {{ISBN|9781578590490}}</ref> which does not include other arthropods or insects of other orders such as [[Ant|ants]], [[Bee|bees]], [[Beetle|beetles]], or [[Butterfly|butterflies]]. In some variations of English, all [[Terrestrial animal|terrestrial]] [[arthropod]]s (including non-insect [[arachnid]]s, and [[myriapod]]s) also fall under the [[Colloquialism|colloquial]] understanding of ''bug''.{{efn|The [[Museum of New Zealand]] notes that "in everyday conversation", ''bug'' "refers to land arthropods with at least six legs, such as insects, spiders, and centipedes".<ref>[https://www.tepapa.govt.nz/discover-collections/read-watch-play/science/your-bug-questions-answered/what-bug-insects-arachnids "What is a bug? Insects, arachnids, and myriapods"] at Museum of New Zealand Te Papa Tongarewa website. Accessed 10 March 2022.</ref>  In a chapter on "Bugs That Are Not Insects", the entomologist Gilbert Walbauer specifies centipedes, millipedes, arachnids (spiders, [[Opiliones|daddy longlegs]], scorpions, [[mite]]s, [[chigger]]s and ticks) as well as the few terrestrial crustaceans ([[sowbug]]s and [[pillbug]]s),<ref name="Walbauer5">Gilbert Waldbauer. ''The Handy Bug Answer Book.'' Visible Ink, 1998. [https://archive.org/details/handybuganswerbo00wald/page/5/mode/2up pp. 5–26.] {{ISBN|9781578590490}}</ref> but argues that "including legless creatures such as worms, slugs, and snails among the bugs stretches the word too much".<ref name="Walbauer1"/> "Bug" is used for a harmful microorganism, typically a bacterium.<ref>{{cite web |url=https://www.lexico.com/definition/bug |archive-url=https://web.archive.org/web/20191211214247/https://www.lexico.com/definition/bug |url-status=dead |archive-date=December 11, 2019 |title=BUG &#124; Meaning & Definition for UK English |publisher=Lexico.com |accessdate=2022-08-03}}</ref>}}
+Entomologists reserve the term ''bug'' for Hemiptera or Heteroptera,<ref name="Walbauer1">Gilbert Waldbauer. ''The Handy Bug Answer Book.'' Visible Ink, 1998. [https://archive.org/details/handybuganswerbo00wald/page/1/mode/2up p. 1.] {{ISBN|9781578590490}}</ref> which does not include other arthropods or insects of other orders such as [[ants]], [[bees]], [[beetles]], or [[butterflies]]. In some variations of English, all [[Terrestrial animal|terrestrial]] [[arthropod]]s (including non-insect [[arachnid]]s, and [[myriapod]]s) also fall under the [[colloquial]] understanding of ''bug''.{{efn|The [[Museum of New Zealand]] notes that "in everyday conversation", ''bug'' "refers to land arthropods with at least six legs, such as insects, spiders, and centipedes".<ref>[https://www.tepapa.govt.nz/discover-collections/read-watch-play/science/your-bug-questions-answered/what-bug-insects-arachnids "What is a bug? Insects, arachnids, and myriapods"] at Museum of New Zealand Te Papa Tongarewa website. Accessed 10 March 2022.</ref>  In a chapter on "Bugs That Are Not Insects", the entomologist Gilbert Walbauer specifies centipedes, millipedes, arachnids (spiders, [[Opiliones|daddy longlegs]], scorpions, [[mite]]s, [[chigger]]s and ticks) as well as the few terrestrial crustaceans ([[sowbug]]s and [[pillbug]]s),<ref name="Walbauer5">Gilbert Waldbauer. ''The Handy Bug Answer Book.'' Visible Ink, 1998. [https://archive.org/details/handybuganswerbo00wald/page/5/mode/2up pp. 5–26.] {{ISBN|9781578590490}}</ref> but argues that "including legless creatures such as worms, slugs, and snails among the bugs stretches the word too much".<ref name="Walbauer1"/> "Bug" is used for a harmful microorganism, typically a bacterium.<ref>{{cite web |url=https://www.lexico.com/definition/bug |archive-url=https://web.archive.org/web/20191211214247/https://www.lexico.com/definition/bug |url-status=dead |archive-date=December 11, 2019 |title=BUG &#124; Meaning & Definition for UK English |publisher=Lexico.com |accessdate=2022-08-03}}</ref>}}
-Many insects with "bug" in their [[common name]], especially in [[American English]], belong to other orders; for example, the [[lovebug]] is a [[Diptera|fly]]<ref name="denmark">{{cite web |url=http://entomology.ifas.ufl.edu/creatures/misc/lovebug.htm |title=Lovebug, ''Plecia nearctica'' Hardy |first=Harold |last=Denmark |author2=Mead, Frank |author3=Fasulo, Thomas |publisher=University of Florida/IFAS |date= April 2010 |work=Featured Creatures |access-date=10 March 2022 }}</ref> and the [[May bug|Maybug]] and [[Coccinellidae|ladybug]] are [[beetle]]s.<ref>{{cite web |title=''Melolontha melolontha'' (cockchafer or May bug) |url=http://www.nhm.ac.uk/nature-online/species-of-the-day/biodiversity/economic-impact/melolontha-melolontha/|publisher=Natural History Museum |archive-url=https://web.archive.org/web/20150703134124/http://www.nhm.ac.uk/nature-online/species-of-the-day/biodiversity/economic-impact/melolontha-melolontha/ |archive-date=3 July 2015 |access-date=12 July 2015}}</ref>  The term is occasionally extended to colloquial names for freshwater or marine [[Crustacean|crustaceans]] (e.g. [[Balmain bug]], [[Moreton Bay bug]], [[mudbug]]) and used by physicians and bacteriologists for disease-causing germs (e.g. [[Antimicrobial resistance|superbug]]s).<ref name="Walbauer1"/>
+Many insects with "bug" in their [[common name]], especially in [[American English]], belong to other orders; for example, the [[lovebug]] is a [[fly]]<ref name="denmark">{{cite web |url=http://entomology.ifas.ufl.edu/creatures/misc/lovebug.htm |title=Lovebug, ''Plecia nearctica'' Hardy |first=Harold |last=Denmark |author2=Mead, Frank |author3=Fasulo, Thomas |publisher=University of Florida/IFAS |date= April 2010 |work=Featured Creatures |access-date=10 March 2022 }}</ref> and the [[Maybug]] and [[ladybug]] are [[beetle]]s.<ref>{{cite web |title=''Melolontha melolontha'' (cockchafer or May bug) |url=http://www.nhm.ac.uk/nature-online/species-of-the-day/biodiversity/economic-impact/melolontha-melolontha/|publisher=Natural History Museum |archive-url=https://web.archive.org/web/20150703134124/http://www.nhm.ac.uk/nature-online/species-of-the-day/biodiversity/economic-impact/melolontha-melolontha/ |archive-date=3 July 2015 |access-date=12 July 2015}}</ref>  The term is occasionally extended to colloquial names for freshwater or marine [[crustaceans]] (e.g. [[Balmain bug]], [[Moreton Bay bug]], [[mudbug]]) and used by physicians and bacteriologists for disease-causing germs (e.g. [[Antimicrobial resistance|superbug]]s).<ref name="Walbauer1"/>
-Most hemipterans feed on plants, using their sucking and piercing mouthparts to extract [[Sap|plant sap]]. Some are bloodsucking, or [[hematophagous]], while others are [[predation|predators]] that feed on other insects or small [[invertebrate]]s. 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. [[Gerridae|pondskaters]], [[Corixidae|water boatmen]], [[Belostomatidae|giant water bugs]]). Hemipterans are [[hemimetabolous]], with young [[Nymph (biology)|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.
+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 [[invertebrate]]s. 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. [[Gerridae|pondskaters]], [[Corixidae|water boatmen]], [[giant water bugs]]). Hemipterans are [[hemimetabolous]], with young [[Nymph (biology)|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 [[pest (organism)|agricultural pests]], damaging crops by sucking the sap. Others harm humans more directly as [[Vector (epidemiology)|vectors]] of serious viral diseases. The [[Bed bug (insect)|bed bug]] is a persistent parasite of humans, and some [[kissing bug]]s 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 [[shellac]]. [[Cicada]]s have been used as food, and have appeared in literature since the ''[[Iliad]]'' in [[Ancient Greece]].
+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 [[Vector (epidemiology)|vectors]] of serious viral diseases. The [[bed bug]] is a persistent parasite of humans, and some [[kissing bug]]s 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 [[shellac]]. [[Cicada]]s have been used as food, and have appeared in literature since the ''[[Iliad]]'' in [[Ancient Greece]].
 == Diversity ==
-Hemiptera is the largest order of [[Hemimetabolism|hemimetabolous]] insects (not undergoing complete metamorphosis; though some examples such as male [[scale insects]] do undergo a form of complete metamorphosis <ref name=Gullan1997>{{cite journal|last1=Gullan |first1=Penny |last2=Kosztarab |first2=Michael |title=Adaptations in scale insects |journal=Annual Review of Entomology |date=1997 |volume=42 |pages=23–50 |doi=10.1146/annurev.ento.42.1.23 |pmid=15012306 }}</ref>), containing over 95,000 named species. Other insect orders with more species are all [[Holometabolism|holometabolous]], meaning they have a [[pupa]]l 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 [[Corixidae|water boatmen]], [[Notonectidae|backswimmers]], [[Gerridae|pond skaters]], and [[Belostomatidae|giant water bugs]].<ref name=Coulson/>
+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 <ref name=Gullan1997>{{cite journal|last1=Gullan |first1=Penny |last2=Kosztarab |first2=Michael |title=Adaptations in scale insects |journal=Annual Review of Entomology |date=1997 |volume=42 |pages=23–50 |doi=10.1146/annurev.ento.42.1.23 |pmid=15012306 }}</ref>), containing over 95,000 named species. Other insect orders with more species are all [[holometabolous]], meaning they have a [[pupa]]l 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 [[Corixidae|water boatmen]], [[backswimmers]], [[Gerridae|pond skaters]], and [[giant water bugs]].<ref name=Coulson/>
 == Taxonomy and phylogeny ==
@@ Line 43: / Line 43: @@
 The [[fossil record]] of hemipterans goes back to the [[Carboniferous]] ([[Moscovian (Carboniferous)|Moscovian]]).<ref name=natureholometabolousinsects>{{Cite journal |author1=André Nel |author2=Patrick Roques |author3=Patricia Nel |author4=Alexander A. Prokin |author5=Thierry Bourgoin |author6=Jakub Prokop |author7=Jacek Szwedo |author8=Dany Azar |author9=Laure Desutter-Grandcolas |author10=Torsten Wappler |author11=Romain Garrouste |author12=David Coty |author13=Diying Huang |author14=Michael S. Engel |author15=Alexander G. Kirejtshuk |display-authors=5 |year=2013 |title=The earliest known holometabolous insects |journal=Nature |volume=503 |issue=7475 |pages=257–261 |doi=10.1038/nature12629 |pmid=24132233 |bibcode=2013Natur.503..257N |s2cid=205235859 }}</ref> The oldest fossils are of the [[Archescytinidae]] from the [[Lower Permian]] and are thought to be basal to the [[Auchenorrhyncha]]. [[Fulgoromorpha]] and [[Cicadomorpha]] appear in the [[Upper Permian]], as do [[Sternorrhyncha]] of the [[Psylloidea]] and [[Aleyrodoidea]]. [[Aphids]] and [[Coccoidea|Coccoids]] appear in the [[Triassic]]. The [[Coleorrhyncha]] extend back to the [[Lower Jurassic]].<ref>{{cite book| page=213|author=Gillott, Cedric| title=Entomology|edition=3| year=2005| publisher= Springer}}</ref> The [[Heteroptera]] first appeared in the [[Triassic]].<ref>{{cite journal |author=Shcherbakov, D. E. |year=2000 |title=Permian faunas of Homoptera (Hemiptera) in relation to phytogeography and the Permo-Triassic crisis |journal=Paleontological Journal |volume=34 |issue=3 |pages=S251–S267 |url=http://palaeoentomolog.ru/Publ/PALS251.pdf}}</ref>
-The present members of the order Hemiptera (sometimes referred to as Rhynchota) were historically placed into two orders, the so-called Homo<!--unlinked, group is paraphyletic-->ptera and Heteroptera/Hemiptera, based on differences in wing structure and the position of the [[Rostrum (anatomy)|rostrum]]. The order is now divided into four suborders, after the "Homoptera" were established as [[paraphyletic]] (now the [[Auchenorrhyncha]] and the [[Sternorrhyncha]]).<ref name=Song2012>{{cite journal|last1=Song |first1=Nan |last2=Liang |first2=Ai-Ping |last3=Bu |first3=Cui-Ping |title=A Molecular Phylogeny of Hemiptera Inferred from Mitochondrial Genome Sequences |journal=PLOS ONE |date=2012 |volume=7 |issue=11 |pages=e48778 |doi=10.1371/journal.pone.0048778 |pmid=23144967 |bibcode=2012PLoSO...748778S |pmc=3493603 |doi-access=free }}</ref><ref name=Forero>{{cite journal|title=The Systematics of Hemiptera |author=Forero, Dimitri |journal=Revista Colombiana de Entomología |volume=34 |issue=1 |date=2008 |pages=1–21 |doi=10.25100/socolen.v34i1.9244 |s2cid=85830606 |url=http://www.scielo.org.co/pdf/rcen/v34n1/v34n1a01.pdf |url-status=dead |archive-url=https://web.archive.org/web/20150113135356/http://www.scielo.org.co/pdf/rcen/v34n1/v34n1a01.pdf |archive-date=January 13, 2015 }}</ref><ref>{{cite journal |last1=Cryan |first1=Jason C. |last2=Urban |first2=Julie M. |title=Higher-level phylogeny of the insect order Hemiptera: is Auchenorrhyncha really paraphyletic? |journal=Systematic Entomology |date=January 2012 |volume=37 |issue=1 |pages=7–21 |doi=10.1111/j.1365-3113.2011.00611.x|s2cid=86564882 }}</ref>
+The present members of the order Hemiptera (sometimes referred to as Rhynchota) were historically placed into two orders, the so-called Homo<!--unlinked, group is paraphyletic-->ptera and Heteroptera/Hemiptera, based on differences in wing structure and the position of the [[Rostrum (anatomy)|rostrum]]. The order is now divided into four suborders, after the "Homoptera" were established as [[paraphyletic]] (now the [[Auchenorrhyncha]] and the [[Sternorrhyncha]]).<ref name=Song2012>{{cite journal|last1=Song |first1=Nan |last2=Liang |first2=Ai-Ping |last3=Bu |first3=Cui-Ping |title=A Molecular Phylogeny of Hemiptera Inferred from Mitochondrial Genome Sequences |journal=PLOS ONE |date=2012 |volume=7 |issue=11 |pages=e48778 |doi=10.1371/journal.pone.0048778 |pmid=23144967 |bibcode=2012PLoSO...748778S |pmc=3493603 |doi-access=free }}</ref><ref name=Forero>{{cite journal|title=The Systematics of Hemiptera |author=Forero, Dimitri |journal=Revista Colombiana de Entomología |volume=34 |issue=1 |date=2008 |pages=1–21 |doi=10.25100/socolen.v34i1.9244 |s2cid=85830606 |url=http://www.scielo.org.co/pdf/rcen/v34n1/v34n1a01.pdf |url-status=dead |archive-url=https://web.archive.org/web/20150113135356/http://www.scielo.org.co/pdf/rcen/v34n1/v34n1a01.pdf |archive-date=January 13, 2015 }}</ref><ref>{{cite journal |last1=Cryan |first1=Jason C. |last2=Urban |first2=Julie M. |title=Higher-level phylogeny of the insect order Hemiptera: is Auchenorrhyncha really paraphyletic? |journal=Systematic Entomology |date=January 2012 |volume=37 |issue=1 |pages=7–21 |doi=10.1111/j.1365-3113.2011.00611.x|bibcode=2012SysEn..37....7C |s2cid=86564882 }}</ref>
 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.<ref name="Johnson"/>
@@ Line 53: / Line 53: @@
   |label1='''[[Psocodea]]'''
   |1={{clade
-   |1=[[Trogiomorpha]] (barklice) [[File:Psocoptera icon.png |70px]]
+   |1=[[Trogiomorpha]] (granary booklice and relatives) <!--[[File:Psocoptera icon.png |70px]] -->
    |2={{clade
     |1=[[Psocomorpha]] (barklice) [[File:Psocoptera icon.png |70px]]
@@ Line 89: / Line 89: @@
 |-
 | [[Heteroptera]] || over 45,000<ref name="Cassis2019">{{cite book|last1=Cassis|first1=Gerasimos|title=Reference Module in Life Sciences|chapter=True Bugs (Insecta: Hemiptera: Heteroptera): Evolution, Classification, Biodiversity and Biology|year=2019|doi=10.1016/B978-0-12-809633-8.20710-3|isbn=9780128096338|s2cid=214379746}}</ref>
-|| [[Triassic]] || [[shield bug]]s, [[Lygaeoidea|seed bug]]s, [[assassin bug]]s, [[Anthocoridae|flower bugs]], [[Coreidae|leaf-footed bugs]], water bugs, [[Miridae|plant bugs]]|| larger bugs; some are [[predator]]y, others are plant-sucking
+|| [[Triassic]] || [[shield bug]]s, [[Lygaeoidea|seed bug]]s, [[assassin bug]]s, [[Anthocoridae|flower bugs]], [[leaf-footed bugs]], water bugs, [[Miridae|plant bugs]]|| larger bugs; some are [[predator]]y, others are plant-sucking
 |-
 | [[Sternorrhyncha]] || 12,500 || [[Upper Permian]] || [[aphids]], [[whiteflies]], [[scale insects]] || plant-sucking bugs, some major horticultural pests; most are small and [[sedentary]] or fully sessile;<ref>{{cite web |title=Sternorrhyncha |url=http://www.amentsoc.org/insects/fact-files/orders/hemiptera-sternorrhyncha.html |publisher=Amateur Entomologists' Society |access-date=13 July 2015}}</ref>
@@ Line 100: / Line 100: @@
 [[File:HemipteraMouthparts.jpg|thumb|Hemipteran mouthparts are distinctive, with [[Mandible (insect)|mandibles]] and [[Insect mouthparts|maxillae]] modified to form a piercing "stylet" sheathed within a modified [[Labium (insect)|labium]].]]
-The defining feature of hemipterans is their "beak" in which the modified [[Mandible (insect)|mandibles]] and [[Insect mouthparts|maxillae]] form a "stylet" which is sheathed within a modified [[Labium (insect)|labium]]. The stylet is capable of piercing [[Tissue (biology)|tissues]] and sucking liquids, while the labium supports it. The stylet contains a channel for the outward movement of [[saliva]] and another for the inward movement of liquid food. A salivary pump drives saliva into the prey; a [[cibarial pump]] extracts liquid from the prey. Both pumps are powered by substantial dilator muscles in the head. The beak is usually folded under the body when not in use. The diet is typically [[Sap|plant sap]], but some hemipterans such as [[Reduviidae|assassin bugs]] are [[Predation|predators]].<ref name=Ruppert>{{cite book |title=Invertebrate Zoology, 7th edition |last1=Ruppert |first1=Edward E. |last2=Fox |first2=Richard, S. |last3=Barnes |first3=Robert D. |year=2004 |publisher=Cengage Learning |isbn=978-81-315-0104-7 |pages=728, 748 }}</ref><ref name=Gullan/>
+The defining feature of hemipterans is their "beak" in which the modified [[Mandible (insect)|mandibles]] and [[Insect mouthparts|maxillae]] form a "stylet" which is sheathed within a modified [[Labium (insect)|labium]]. The stylet is capable of piercing [[Tissue (biology)|tissues]] and sucking liquids, while the labium supports it. The stylet contains a channel for the outward movement of [[saliva]] and another for the inward movement of liquid food. A salivary pump drives saliva into the prey; a [[cibarial pump]] extracts liquid from the prey. Both pumps are powered by substantial dilator muscles in the head. The beak is usually folded under the body when not in use. The diet is typically [[plant sap]], but some hemipterans such as [[assassin bugs]] are [[predators]].<ref name=Ruppert>{{cite book |title=Invertebrate Zoology, 7th edition |last1=Ruppert |first1=Edward E. |last2=Fox |first2=Richard, S. |last3=Barnes |first3=Robert D. |year=2004 |publisher=Cengage Learning |isbn=978-81-315-0104-7 |pages=728, 748 }}</ref><ref name=Gullan/>
 Both herbivorous and predatory hemipterans inject [[enzyme]]s to begin digestion extra-orally (before the food is taken into the body). These enzymes include [[amylase]] to hydrolyse [[starch]], [[polygalacturonase]] to weaken the tough [[cell wall]]s of plants, and [[proteinase]]s to break down proteins.<ref name="Wheeler2001">{{cite book|last=Wheeler|first=Alfred George|title=Biology of the Plant Bugs (Hemiptera: Miridae): Pests, Predators, Opportunists|url=https://books.google.com/books?id=0szFXwGVlo4C&pg=PA105|year=2001|publisher=Cornell University Press|isbn=978-0-8014-3827-1|pages=105–135}}</ref>
-Although the Hemiptera vary widely in their overall form, their mouthparts form a distinctive "rostrum". Other insect orders with mouthparts modified into anything like the rostrum and stylets of the Hemiptera include some [[Phthiraptera]], but for other reasons they generally are easy to recognize as non-hemipteran. Similarly, the mouthparts of [[Siphonaptera]], some [[Diptera]] and [[Thysanoptera]] superficially resemble the rostrum of the Hemiptera, but on closer inspection the differences are considerable. Aside from the mouthparts, various other insects can be confused with Hemiptera, but they all have biting mandibles and maxillae instead of the rostrum. Examples include [[cockroach]]es and [[Psocoptera|psocids]], both of which have longer, many-segmented antennae, and some [[beetle]]s, but these have fully hardened forewings which do not overlap.<ref name="Chinery">{{cite book |author=Chinery, Michael |title=Insects of Britain and Northern Europe |edition=3rd |publisher=Collins |year=1993 |isbn=978-0-00-219918-6}}</ref>
+Although the Hemiptera vary widely in their overall form, their mouthparts form a distinctive "rostrum". Other insect orders with mouthparts modified into anything like the rostrum and stylets of the Hemiptera include some [[Phthiraptera]], but for other reasons they generally are easy to recognize as non-hemipteran. Similarly, the mouthparts of [[Siphonaptera]], some [[Diptera]] and [[Thysanoptera]] superficially resemble the rostrum of the Hemiptera, but on closer inspection the differences are considerable. Aside from the mouthparts, various other insects can be confused with Hemiptera, but they all have biting mandibles and maxillae instead of the rostrum. Examples include [[cockroach]]es and [[psocids]], both of which have longer, many-segmented antennae, and some [[beetle]]s, but these have fully hardened forewings which do not overlap.<ref name="Chinery">{{cite book |author=Chinery, Michael |title=Insects of Britain and Northern Europe |edition=3rd |publisher=Collins |year=1993 |isbn=978-0-00-219918-6}}</ref>
 === Wing structure ===
-The [[insect wing|forewings]] of Hemiptera are either entirely membranous, as in the [[Sternorrhyncha]] and [[Auchenorrhyncha]], or partially hardened, as in most [[Heteroptera]]. The name "Hemiptera" is from the [[Greek language|Greek]] {{lang|el|[[:wikt:ἡμι-|ἡμι-]]}} (''{{lang|el|[[:wikt:hemi-|hemi]]}}''; "half") and {{lang|el|[[:wikt:πτερόν|πτερόν]]}} (''{{lang|el|pteron}}''; "wing"), referring to the forewings of many heteropterans [[Insect wing#Adaptations|which are hardened]] near the base, but membranous at the ends. Wings modified in this manner are termed ''hemelytra'' (singular: ''hemelytron''), by analogy with the completely hardened [[elytron|elytra]] of [[beetle]]s, and occur only in the suborder [[Heteroptera]]. In all suborders, the hindwings – if present at all – are entirely membranous and usually shorter than the forewings.<ref name=Coulson>{{cite book|author1=Coulson, Robert N.|author2=Witter, John A.|title=Forest Entomology: Ecology and Management |url=https://books.google.com/books?id=-QsVpIEsuLcC&pg=PA71 |year=1984 |publisher=John Wiley & Sons |isbn=978-0-471-02573-3 |pages=71–72}}</ref>  The forewings may be held "roofwise" over the body (typical of [[Sternorrhyncha]] and [[Auchenorrhyncha]]),<ref name=Alford>{{cite book|author=Alford, David V.|title=Pests of Ornamental Trees, Shrubs and Flowers: A Color Handbook |url=https://books.google.com/books?id=c9PwiScwmPIC&pg=PA12 |year=2012 |publisher=Academic Press |isbn=978-0-12-398515-6 |page=12}}</ref> or held flat on the back, with the ends overlapping (typical of [[Heteroptera]]).<ref name=Coulson/> The [[antenna (biology)|antennae]] in Hemiptera typically consist of four or five segments, although they can still be quite long, and the tarsi of the [[arthropod leg|legs]] have two or three segments.<ref name=Discover>{{cite web |url=http://www.discoverlife.org/mp/20q?search=Hemiptera |title=Hemiptera |publisher=Discover Life |access-date=13 July 2015}}</ref>
+The [[forewings]] of Hemiptera are either entirely membranous, as in the [[Sternorrhyncha]] and [[Auchenorrhyncha]], or partially hardened, as in most [[Heteroptera]]. The name "Hemiptera" is from the [[Greek language|Greek]] {{lang|el|[[:wikt:ἡμι-|ἡμι-]]}} (''{{lang|el|[[:wikt:hemi-|hemi]]}}''; "half") and {{lang|el|[[:wikt:πτερόν|πτερόν]]}} (''{{lang|el|pteron}}''; "wing"), referring to the forewings of many heteropterans [[Insect wing#Adaptations|which are hardened]] near the base, but membranous at the ends. Wings modified in this manner are termed ''hemelytra'' (singular: ''hemelytron''), by analogy with the completely hardened [[elytra]] of [[beetle]]s, and occur only in the suborder [[Heteroptera]]. In all suborders, the hindwings – if present at all – are entirely membranous and usually shorter than the forewings.<ref name=Coulson>{{cite book|author1=Coulson, Robert N.|author2=Witter, John A.|title=Forest Entomology: Ecology and Management |url=https://books.google.com/books?id=-QsVpIEsuLcC&pg=PA71 |year=1984 |publisher=John Wiley & Sons |isbn=978-0-471-02573-3 |pages=71–72}}</ref>  The forewings may be held "roofwise" over the body (typical of [[Sternorrhyncha]] and [[Auchenorrhyncha]]),<ref name=Alford>{{cite book|author=Alford, David V.|title=Pests of Ornamental Trees, Shrubs and Flowers: A Color Handbook |url=https://books.google.com/books?id=c9PwiScwmPIC&pg=PA12 |year=2012 |publisher=Academic Press |isbn=978-0-12-398515-6 |page=12}}</ref> or held flat on the back, with the ends overlapping (typical of [[Heteroptera]]).<ref name=Coulson/> The [[antenna (biology)|antennae]] in Hemiptera typically consist of four or five segments, although they can still be quite long, and the tarsi of the [[arthropod leg|legs]] have two or three segments.<ref name=Discover>{{cite web |url=http://www.discoverlife.org/mp/20q?search=Hemiptera |title=Hemiptera |publisher=Discover Life |access-date=13 July 2015}}</ref>
 === Sound production ===
@@ Line 116: / Line 116: @@
 Many hemipterans can produce sound for communication.<ref>{{cite journal|last1=Virant-Doberlet|first1=Meta|last2=Cokl|first2=Andrej|title=Vibrational Communication in Insects|journal=Neotropical Entomology|date=April 2004|volume=33|issue=2|doi=10.1590/S1519-566X2004000200001|pages=121–134|doi-access=free}}</ref> The "song" of male cicadas, the loudest of any insect, is produced by [[tymbal]] organs on the underside of the abdomen, and is used to attract mates. The tymbals are drumlike disks of cuticle, which are clicked in and out repeatedly, making a sound in the same way as popping the metal lid of a jam jar in and out.<ref>{{cite journal |last1=Simmons |first1=Peter |last2=Young |first2=David |title=The Tymbal Mechanism and Song Patterns of the Bladder Cicada, ''Cystosoma saundersii'' |journal=Journal of Experimental Biology |date=1978 |volume=76 |pages=27–45 |doi=10.1242/jeb.76.1.27 |url=http://download.naturkundemuseum-berlin.de/andreas.wessel/Cicada_morphology/simmons_young_1978_cystosoma_saundersii_tymbal_mechanism.pdf |access-date=2015-08-22 |archive-url=https://web.archive.org/web/20151224214649/http://download.naturkundemuseum-berlin.de/andreas.wessel/Cicada_morphology/simmons_young_1978_cystosoma_saundersii_tymbal_mechanism.pdf |archive-date=2015-12-24 |url-status=dead }}</ref>
-[[Stridulation|Stridulatory]] sounds are produced among the aquatic [[Corixidae]] and [[Notonectidae]] (backswimmers) using tibial combs rubbed across rostral ridges.<ref>{{cite journal |last1=Wilcox |first1=R. Stimson |title=Sound-producing Mechanisms of ''Bueno macrotibialis'' Hungerford (Hemiptera: Notonectidae) |journal=International Journal of Insect Morphology & Embryology |year=1975 |volume=4 |issue=21 |pages=169–182 |url=http://deepblue.lib.umich.edu/bitstream/handle/2027.42/22083/0000507.pdf?sequence=1 |doi=10.1016/0020-7322(75)90015-x|hdl=2027.42/22083 |hdl-access=free }}</ref><ref>{{cite book|author1=Gooderham, John|author2=Tsyrlin, Edward|title=The Waterbug Book: A Guide to the Freshwater Macroinvertebrates of Temperate Australia |url=https://books.google.com/books?id=0_Paklse3XcC&pg=PA146|year=2002 |publisher=Csiro Publishing |isbn=978-0-643-06668-7 |page=146}}</ref>
+[[Stridulatory]] sounds are produced among the aquatic [[Corixidae]] and [[Notonectidae]] (backswimmers) using tibial combs rubbed across rostral ridges.<ref>{{cite journal |last1=Wilcox |first1=R. Stimson |title=Sound-producing Mechanisms of ''Bueno macrotibialis'' Hungerford (Hemiptera: Notonectidae) |journal=International Journal of Insect Morphology & Embryology |year=1975 |volume=4 |issue=21 |pages=169–182 |url=http://deepblue.lib.umich.edu/bitstream/handle/2027.42/22083/0000507.pdf?sequence=1 |doi=10.1016/0020-7322(75)90015-x|hdl=2027.42/22083 |hdl-access=free }}</ref><ref>{{cite book|author1=Gooderham, John|author2=Tsyrlin, Edward|title=The Waterbug Book: A Guide to the Freshwater Macroinvertebrates of Temperate Australia |url=https://books.google.com/books?id=0_Paklse3XcC&pg=PA146|year=2002 |publisher=Csiro Publishing |isbn=978-0-643-06668-7 |page=146}}</ref>
 === Life cycle ===
@@ Line 122: / Line 122: @@
 [[File:Ant-mimic predatory bug, Myrmecoris gracilis, on Yarrow.JPG|thumb|upright|left|An [[ant mimicry|ant-mimicking]] predatory bug ''[[Myrmecoris gracilis]]'']]
-Hemipterans are [[Hemimetabolism|hemimetabolous]], meaning that they do not undergo [[metamorphosis]], the complete change of form between a [[larva]]l phase and an [[imago|adult phase]]. Instead, their young are called [[Nymph (biology)|nymphs]], and resemble the adults to a greater or lesser degree. The nymphs moult several times as they grow, and each [[instar]] resembles the adult more than the previous one. Wing buds grow in later stage nymphs; the final transformation involves little more than the development of functional wings (if they are present at all) and functioning sexual organs, with no intervening [[pupa]]l stage as in [[Holometabolism|holometabolous]] insects.<ref>{{cite web |url=https://australian.museum/learn/animals/insects/ |title=Metamorphosis: a remarkable change |author=Britton, David |date=9 July 2009 |publisher=Australian Museum |access-date=13 July 2015}}</ref>
+Hemipterans are [[hemimetabolous]], meaning that they do not undergo [[metamorphosis]], the complete change of form between a [[larva]]l phase and an [[imago|adult phase]]. Instead, their young are called [[Nymph (biology)|nymphs]], and resemble the adults to a greater or lesser degree. The nymphs moult several times as they grow, and each [[instar]] resembles the adult more than the previous one. Wing buds grow in later stage nymphs; the final transformation involves little more than the development of functional wings (if they are present at all) and functioning sexual organs, with no intervening [[pupa]]l stage as in [[holometabolous]] insects.<ref>{{cite web |url=https://australian.museum/learn/animals/insects/ |title=Metamorphosis: a remarkable change |author=Britton, David |date=9 July 2009 |publisher=Australian Museum |access-date=13 July 2015}}</ref>
 ==== Parthenogenesis and viviparity ====
@@ Line 129: / Line 129: @@
 {{further|parthenogenesis|thelytoky|viviparity}}
-Many [[aphid]]s are [[parthenogenesis|parthenogenetic]] during part of the life cycle, such that females can produce unfertilized eggs, which are [[Cloning|clones]] of their mother. All such young are females ([[thelytoky]]), so 100% of the population at these times can produce more offspring. Many species of aphid are also [[Viviparity|viviparous]]: the young are born live rather than laid as eggs. These adaptations enable aphids to reproduce extremely rapidly when conditions are suitable.<ref>{{cite web |url=http://www.biology-resources.com/aphid-01.html |title=Aphids, an Introduction |author=Mackean, D.G. |work=Biology Teaching Resources |access-date=13 July 2015}}</ref>
+Many [[aphid]]s are [[parthenogenetic]] during part of the life cycle, such that females can produce unfertilized eggs, which are [[Cloning|clones]] of their mother. All such young are females ([[thelytoky]]), so 100% of the population at these times can produce more offspring. Many species of aphid are also [[viviparous]]: the young are born live rather than laid as eggs. These adaptations enable aphids to reproduce extremely rapidly when conditions are suitable.<ref>{{cite web |url=http://www.biology-resources.com/aphid-01.html |title=Aphids, an Introduction |author=Mackean, D.G. |work=Biology Teaching Resources |access-date=13 July 2015}}</ref>
 == Locomotion ==
@@ Line 140: / Line 140: @@
 === Swimming and skating ===
-Several families of Heteroptera are ''water bugs'', adapted to an [[aquatic animal|aquatic lifestyle]], such as the [[Corixidae|water boatmen]] (Corixidae), [[Nepidae|water scorpions]] (Nepidae), and [[Notonectidae|backswimmers]] (Notonectidae). They are mostly predatory, and have legs adapted as [[paddle]]s to help the animal move through the water.<ref name="NHM">{{cite web |url=http://www.nhm.ac.uk/resources-rx/files/26feat_its_a_bugs_life-3013.pdf  |title=Hemiptera...It's a Bug's Life |publisher=Natural History Museum |author1=Jon Martin |author2=Mick Webb |access-date=July 26, 2010}}</ref> The [[Gerridae|pondskaters]] or water striders (Gerridae) are also associated with water, but use the [[surface tension]] of standing water to keep them above the surface;<ref name=Bush/> they include the sea skaters in the genus ''[[Halobates]]'', the only truly marine group of insects.<ref name="NHM"/>
+Several families of Heteroptera are ''water bugs'', adapted to an [[aquatic animal|aquatic lifestyle]], such as the [[Corixidae|water boatmen]] (Corixidae), [[Nepidae|water scorpions]] (Nepidae), and [[backswimmers]] (Notonectidae). They are mostly predatory, and have legs adapted as [[paddle]]s to help the animal move through the water.<ref name="NHM">{{cite web |url=http://www.nhm.ac.uk/resources-rx/files/26feat_its_a_bugs_life-3013.pdf  |title=Hemiptera...It's a Bug's Life |publisher=Natural History Museum |author1=Jon Martin |author2=Mick Webb |access-date=July 26, 2010}}</ref> The [[Gerridae|pondskaters]] or water striders (Gerridae) are also associated with water, but use the [[surface tension]] of standing water to keep them above the surface;<ref name=Bush/> they include the sea skaters in the genus ''[[Halobates]]'', the only truly marine group of insects.<ref name="NHM"/>
-[[File:MicroveliaIndia.jpg|thumb|upright=0.9|Adult and nymph ''[[Microvelia]]'' water bugs using [[Marangoni effect|Marangoni propulsion]]]]
+[[File:MicroveliaIndia.jpg|thumb|upright=0.9|Adult and nymph ''[[Microvelia]]'' water bugs using [[Marangoni propulsion]]]]
 === Marangoni propulsion ===
@@ Line 150: / Line 150: @@
 === Flight ===
-Flight is well developed in the Hemiptera although mostly used for short distance movement and dispersal. Wing development is sometimes related to environmental conditions. In some groups of Hemiptera, there are variations of winged, short-winged, and wingless forms within a single species. This kind of polymorphism tends to be helpful when habitats are temporary with more energy put into reproduction when food is available and into dispersal through flight when food becomes scarce.  In aphids, both winged and wingless forms occur with winged forms produced in greater numbers when food resources are depleted. Aphids and whiteflies can sometimes be transported very long distances by atmospheric updrafts and high altitude winds.<ref>{{cite journal |author1= Loxdale, H. D. |author2= Hardie, J. |author3=Halbert, S. |author4=Foottit, R. |author5=Kidd, N. A. C. |author6= Carter, C. I. |year=1993 | title=The relative importance of short- and long-range movement of flying aphids |journal=Biological Reviews |volume=68 |issue=2 |pages=291–311 |doi=10.1111/j.1469-185X.1993.tb00998.x |s2cid=86569822 |url=https://repository.rothamsted.ac.uk/download/77c13e9643abcf2d2cf90fc3220ffa990d82288834c9054e15a503ee7a0a16de/16412309/The_relative_importance_of_short-_and_long-range_m.pdf}}</ref> Wing-length polymorphism is notably rare in tree-living Hemiptera.<ref>{{cite journal |last1=Waloff |first1=Nadia |author-link=Nadia Waloff |doi=10.1111/j.1365-2311.1983.tb00502.x |title=Absence of wing polymorphism in the arboreal, phytophagous species of some taxa of temperate Hemiptera: An hypothesis |journal=Ecological Entomology |volume=8 |issue=2 |pages=229–232 |year=1983 |s2cid=83833159}}</ref>
+Flight is well developed in the Hemiptera although mostly used for short distance movement and dispersal. Wing development is sometimes related to environmental conditions. In some groups of Hemiptera, there are variations of winged, short-winged, and wingless forms within a single species. This kind of polymorphism tends to be helpful when habitats are temporary with more energy put into reproduction when food is available and into dispersal through flight when food becomes scarce.  In aphids, both winged and wingless forms occur with winged forms produced in greater numbers when food resources are depleted. Aphids and whiteflies can sometimes be transported very long distances by atmospheric updrafts and high altitude winds.<ref>{{cite journal |author1= Loxdale, H. D. |author2= Hardie, J. |author3=Halbert, S. |author4=Foottit, R. |author5=Kidd, N. A. C. |author6= Carter, C. I. |year=1993 | title=The relative importance of short- and long-range movement of flying aphids |journal=Biological Reviews |volume=68 |issue=2 |pages=291–311 |doi=10.1111/j.1469-185X.1993.tb00998.x |s2cid=86569822 |url=https://repository.rothamsted.ac.uk/download/77c13e9643abcf2d2cf90fc3220ffa990d82288834c9054e15a503ee7a0a16de/16412309/The_relative_importance_of_short-_and_long-range_m.pdf}}</ref> Wing-length polymorphism is notably rare in tree-living Hemiptera.<ref>{{cite journal |last1=Waloff |first1=Nadia |author-link=Nadia Waloff |doi=10.1111/j.1365-2311.1983.tb00502.x |title=Absence of wing polymorphism in the arboreal, phytophagous species of some taxa of temperate Hemiptera: An hypothesis |journal=Ecological Entomology |volume=8 |issue=2 |pages=229–232 |year=1983 |bibcode=1983EcoEn...8..229W |s2cid=83833159}}</ref>
 === Jumping ===
@@ Line 156: / Line 156: @@
 {{further|Jumping}}
-Many Auchenorrhyncha including representatives of the cicadas, leafhoppers, treehoppers, planthoppers, and froghoppers are adapted for jumping (saltation). Treehoppers, for example, jump by rapidly depressing their hind legs. Before jumping, the hind legs are raised and the femora are pressed tightly into curved indentations in the coxae. Treehoppers can attain a take-off velocity of up to 2.7 metres per second and an acceleration of up to 250 g. The instantaneous power output is much greater than that of normal muscle, implying that energy is stored and released to [[catapult]] the insect into the air.<ref>{{cite journal |last1=Burrows |first1=M. |title=Jumping mechanisms of treehopper insects (Hemiptera, Auchenorrhyncha, Membracidae) |journal=Journal of Experimental Biology |date=March 2013 |volume= 216|issue=5 |pages=788–799 |doi=10.1242/jeb.078741  |pmid=23155084|doi-access=free }}</ref> Cicadas, which are much larger, extend their hind legs for a jump in under a millisecond, again implying elastic storage of energy for sudden release.<ref>{{cite journal |last1=Gorb |first1=S. N. |title=The jumping mechanism of cicada Cercopis vulnerata (Auchenorrhyncha, Cercopidae): skeleton-muscle organisation, frictional surfaces, and inverse-kinematic model of leg movements |journal=Arthropod Structure & Development |date=July 2004 |volume=33 |issue=3 |pages=201–220  |doi=10.1016/j.asd.2004.05.008 |pmid=18089035}}</ref>
+Many Auchenorrhyncha including representatives of the cicadas, leafhoppers, treehoppers, planthoppers, and froghoppers are adapted for jumping (saltation). Treehoppers, for example, jump by rapidly depressing their hind legs. Before jumping, the hind legs are raised and the femora are pressed tightly into curved indentations in the coxae. Treehoppers can attain a take-off velocity of up to 2.7 metres per second and an acceleration of up to 250 g. The instantaneous power output is much greater than that of normal muscle, implying that energy is stored and released to [[catapult]] the insect into the air.<ref>{{cite journal |last1=Burrows |first1=M. |title=Jumping mechanisms of treehopper insects (Hemiptera, Auchenorrhyncha, Membracidae) |journal=Journal of Experimental Biology |date=March 2013 |volume= 216|issue=5 |pages=788–799 |doi=10.1242/jeb.078741  |pmid=23155084|doi-access=free }}</ref> Cicadas, which are much larger, extend their hind legs for a jump in under a millisecond, again implying elastic storage of energy for sudden release.<ref>{{cite journal |last1=Gorb |first1=S. N. |title=The jumping mechanism of cicada Cercopis vulnerata (Auchenorrhyncha, Cercopidae): skeleton-muscle organisation, frictional surfaces, and inverse-kinematic model of leg movements |journal=Arthropod Structure & Development |date=July 2004 |volume=33 |issue=3 |pages=201–220  |doi=10.1016/j.asd.2004.05.008 |pmid=18089035|bibcode=2004ArtSD..33..201G }}</ref>
 ===Sedentary===
@@ Line 171: / Line 171: @@
 Most hemipterans are phytophagous, using their sucking and piercing mouthparts to feed on plant sap. These include cicadas, leafhoppers, treehoppers, planthoppers, froghoppers, aphids, whiteflies, [[scale insect]]s, and some other groups. Some are [[Generalist and specialist species|monophages]], being host specific and only found on one plant taxon, others are [[oligophage]]s, feeding on a few plant groups, while others again are less discriminating [[Generalist and specialist species|polyphages]] and feed on many species of plant.<ref name=Gullan>{{cite book |author1=Gullan, P.J. |author2=Cranston, P.S. |title=The Insects: An Outline of Entomology, 5th Edition |url=https://books.google.com/books?id=lF5hBAAAQBAJ&pg=PT790 |year=2014 |publisher=Wiley |isbn=978-1-118-84616-2 |pages=80–81, 790–}}</ref> The relationship between hemipterans and plants appears to be ancient, with piercing and sucking of plants evident in the [[Early Devonian]] period.<ref>{{cite journal |last1=Labandeira |first1=Conrad C. |title=Early History of Arthropod and Vascular Plant Associations |journal=Annual Review of Earth and Planetary Sciences |date=May 1998 |volume=26 |pages=329–377 |doi=10.1146/annurev.earth.26.1.329 |bibcode=1998AREPS..26..329L }}</ref>
-Hemipterans can dramatically cut the mass of affected plants, especially in major outbreaks. They sometimes also change the mix of plants by predation on seeds or feeding on roots of certain species.<ref name="Verne2007">{{cite book |last=Verne |first=Nole C. |title=Forest Ecology Research Horizons |url=https://books.google.com/books?id=WQvJiiCUGVEC&pg=PA210 |year=2007 |publisher=Nova Publishers |isbn=978-1-60021-490-5 |page=210}}</ref> Some sap-suckers move from one host to another at different times of year. Many aphids spend the winter as eggs on a woody host plant and the summer as [[Parthenogenesis|parthenogenetically]] reproducing females on a herbaceous plant.<ref name=Dixon>{{cite book |last=Dixon |first=A.F.G. |title=Aphid Ecology: An optimization approach |url=https://books.google.com/books?id=YrvwCAAAQBAJ&pg=PA128 |year=2012 |publisher=Springer Science & Business Media |isbn=978-94-011-5868-8 |page=128}}</ref>
+Hemipterans can dramatically cut the mass of affected plants, especially in major outbreaks. They sometimes also change the mix of plants by predation on seeds or feeding on roots of certain species.<ref name="Verne2007">{{cite book |last=Verne |first=Nole C. |title=Forest Ecology Research Horizons |url=https://books.google.com/books?id=WQvJiiCUGVEC&pg=PA210 |year=2007 |publisher=Nova Publishers |isbn=978-1-60021-490-5 |page=210}}</ref> Some sap-suckers move from one host to another at different times of year. Many aphids spend the winter as eggs on a woody host plant and the summer as [[parthenogenetically]] reproducing females on a herbaceous plant.<ref name=Dixon>{{cite book |last=Dixon |first=A.F.G. |title=Aphid Ecology: An optimization approach |url=https://books.google.com/books?id=YrvwCAAAQBAJ&pg=PA128 |year=2012 |publisher=Springer Science & Business Media |isbn=978-94-011-5868-8 |page=128}}</ref>
 [[File:Wilting a twig (4608641762).jpg|thumb|upright|left|A twig wilting bug ([[Coreidae]]) piercing and sucking sap from a ''[[Zinnia]]'']]
@@ Line 179: / Line 179: @@
 The Typhlocybine Cicadellids specialize in feeding on non-vascular [[mesophyll tissue]] of leaves, which is more nutritious than the leaf epidermis. Most Heteroptera also feed on mesophyll tissue where they are more likely to encounter defensive secondary plant metabolites which often leads to the evolution of host specificity.<ref>{{cite book |last1=Tonkyn |first1=David W. |last2=Whitcomb |first2=Robert F. |title=Current Topics in Vector Research |editor=Harris, Kerry F. |chapter=Feeding Strategies and the Guild Concept Among Vascular Feeding Insects and Microorganisms|doi=10.1007/978-1-4612-4712-8_6 |pages=179–199 |year=1987 |isbn= 978-1-4612-9126-8 }}</ref>
-Obligate xylem feeding is a special habit that is found in the Auchenorrhyncha among Cicadoidea, Cercopoidea and in Cicadelline Cicadellids. Some phloem feeders may take to xylem sap facultatively, especially when facing dehydration.<ref>{{cite journal| title=A phloem-sap feeder mixes phloem and xylem sap to regulate osmotic potential| author1=Pompon, Julien |author2=Quiring, Dan| author3=Goyer, Claudia| author4=Giordanengo, Philippe| author5=Pelletier, Yvan |journal=Journal of Insect Physiology| volume=57| issue=9| year=2011| pages=1317–1322| url=https://www.u-picardie.fr/PCP/data/pub/2011-Pompon%20et%20al.J%20Insect%20Physiol.pdf| doi=10.1016/j.jinsphys.2011.06.007| pmid=21726563| access-date=2015-08-22| archive-url=https://web.archive.org/web/20150923191653/https://www.u-picardie.fr/PCP/data/pub/2011-Pompon%20et%20al.J%20Insect%20Physiol.pdf| archive-date=2015-09-23| url-status=dead}}</ref> Xylem feeders tend to be polyphagous;<ref>{{cite journal|last1=Den|first1=Francesca F. |last2=Stewart |first2=Alan J. A. |last3=Gibson |first3=Amos |last4=Weiblen |first4=George D. |last5=Novotny |first5=Vojtech |title=Low host specificity in species-rich assemblages of xylem- and phloem-feeding herbivores (Auchenorrhyncha) in a New Guinea lowland rain forest |journal=Journal of Tropical Ecology |date=2013 |pages=467–476 |doi=10.1017/S0266467413000540 |url=http://geo.cbs.umn.edu/DemEtAl2013.pdf |volume=29|issue=6 |s2cid=12101409 }}</ref> to overcome the negative pressure of xylem requires a special cibarial pump.<ref>{{cite journal |title= Why are there no small species among xylem-sucking insects? |journal=Evolutionary Ecology |year=1997| volume=11|issue=4 | pages=419–437 |author1=Novotny, Vojtech |author2=Wilson, Michael R. |doi=10.1023/a:1018432807165|s2cid=39222473 }}</ref>
+Obligate xylem feeding is a special habit that is found in the Auchenorrhyncha among Cicadoidea, Cercopoidea and in Cicadelline Cicadellids. Some phloem feeders may take to xylem sap facultatively, especially when facing dehydration.<ref>{{cite journal| title=A phloem-sap feeder mixes phloem and xylem sap to regulate osmotic potential| author1=Pompon, Julien |author2=Quiring, Dan| author3=Goyer, Claudia| author4=Giordanengo, Philippe| author5=Pelletier, Yvan |journal=Journal of Insect Physiology| volume=57| issue=9| year=2011| pages=1317–1322| url=https://www.u-picardie.fr/PCP/data/pub/2011-Pompon%20et%20al.J%20Insect%20Physiol.pdf| doi=10.1016/j.jinsphys.2011.06.007| pmid=21726563| bibcode=2011JInsP..57.1317P | access-date=2015-08-22| archive-url=https://web.archive.org/web/20150923191653/https://www.u-picardie.fr/PCP/data/pub/2011-Pompon%20et%20al.J%20Insect%20Physiol.pdf| archive-date=2015-09-23| url-status=dead}}</ref> Xylem feeders tend to be polyphagous;<ref>{{cite journal|last1=Den|first1=Francesca F. |last2=Stewart |first2=Alan J. A. |last3=Gibson |first3=Amos |last4=Weiblen |first4=George D. |last5=Novotny |first5=Vojtech |title=Low host specificity in species-rich assemblages of xylem- and phloem-feeding herbivores (Auchenorrhyncha) in a New Guinea lowland rain forest |journal=Journal of Tropical Ecology |date=2013 |pages=467–476 |doi=10.1017/S0266467413000540 |url=http://geo.cbs.umn.edu/DemEtAl2013.pdf |volume=29|issue=6 |s2cid=12101409 }}</ref> to overcome the negative pressure of xylem requires a special cibarial pump.<ref>{{cite journal |title= Why are there no small species among xylem-sucking insects? |journal=Evolutionary Ecology |year=1997| volume=11|issue=4 | pages=419–437 |author1=Novotny, Vojtech |author2=Wilson, Michael R. |doi=10.1023/a:1018432807165|bibcode=1997EvEco..11..419N |s2cid=39222473 }}</ref>
-Phloem feeding hemiptera typically have symbiotic micro-organisms in their gut that help to convert [[amino acid]]s. Phloem feeders produce honeydew from their anus. A variety of organisms that feed on honeydew form symbiotic associations with phloem-feeders.<ref>{{cite journal |title=Phloem-sap feeding by animals: problems and solutions |author=Douglas, A.E.| journal=Journal of Experimental Botany| year=2006| volume=57| issue=4| pages=747–754| doi=10.1093/jxb/erj067| pmid=16449374| url=https://academic.oup.com/jxb/article-pdf/57/4/747/1487897/erj067.pdf| doi-access=free}}</ref><ref>{{cite journal| title=Mesophyll cell-sucking herbivores (Cicadellidae: Typhlocybinae) on rainforest trees in Papua New Guinea: local and regional diversity of a taxonomically unexplored guild Ecological Entomology| year=2014| volume=39| issue=3| pages=325–333 |doi=10.1111/een.12104 |author1=Baje, Leontine |author2= Stewart, A.J.A. |author3=Novotny, Vojtech | journal=Ecological Entomology| s2cid=55083254}}</ref> Phloem sap is a sugary liquid low in amino acids, so insects have to process large quantities to meet their nutritional requirements. Xylem sap is even lower in amino acids and contains [[monosaccharide]]s rather than [[sucrose]], as well as organic acids and minerals. No digestion is required (except for the hydrolysis of sucrose) and 90% of the nutrients in the xylem sap can be utilised.<ref name=Gullan/><ref name=Panizzi>{{cite book |author1=Panizzi, Antônio Ricardo |author2=Parra, José R.P.|title=Insect Bioecology and Nutrition for Integrated Pest Management |url=https://books.google.com/books?id=NqzcD2r3jBYC&pg=PA108 |year=2012 |publisher=CRC Press |isbn=978-1-4398-3708-5 |page=108}}</ref> Some phloem sap feeders selectively mix phloem and xylem sap to control the osmotic potential of the liquid consumed.<ref>{{cite journal |last1=Pompon |first1=Julien |last2=Quiring |first2=Dan |last3=Goyer |first3=Claudia |last4=Giordanengo |first4=Philippe |last5=Pelletier |first5=Yvan |title=A phloem-sap feeder mixes phloem and xylem sap to regulate osmotic potential |journal=Journal of Insect Physiology |date=2011 |volume=57 |issue=9 |pages=1317–1322 |url=https://www.u-picardie.fr/PCP/data/pub/2011-Pompon%20et%20al.J%20Insect%20Physiol.pdf |doi=10.1016/j.jinsphys.2011.06.007 |pmid=21726563 |access-date=2015-08-22 |archive-url=https://web.archive.org/web/20150923191653/https://www.u-picardie.fr/PCP/data/pub/2011-Pompon%20et%20al.J%20Insect%20Physiol.pdf |archive-date=2015-09-23 |url-status=dead }}</ref>
+Phloem feeding hemiptera typically have symbiotic micro-organisms in their gut that help to convert [[amino acid]]s. Phloem feeders produce honeydew from their anus. A variety of organisms that feed on honeydew form symbiotic associations with phloem-feeders.<ref>{{cite journal |title=Phloem-sap feeding by animals: problems and solutions |author=Douglas, A.E.| journal=Journal of Experimental Botany| year=2006| volume=57| issue=4| pages=747–754| doi=10.1093/jxb/erj067| pmid=16449374| url=https://academic.oup.com/jxb/article-pdf/57/4/747/1487897/erj067.pdf| doi-access=free}}</ref><ref>{{cite journal| title=Mesophyll cell-sucking herbivores (Cicadellidae: Typhlocybinae) on rainforest trees in Papua New Guinea: local and regional diversity of a taxonomically unexplored guild Ecological Entomology| year=2014| volume=39| issue=3| pages=325–333 |doi=10.1111/een.12104 |author1=Baje, Leontine |author2= Stewart, A.J.A. |author3=Novotny, Vojtech | journal=Ecological Entomology| s2cid=55083254}}</ref> Phloem sap is a sugary liquid low in amino acids, so insects have to process large quantities to meet their nutritional requirements. Xylem sap is even lower in amino acids and contains [[monosaccharide]]s rather than [[sucrose]], as well as organic acids and minerals. No digestion is required (except for the hydrolysis of sucrose) and 90% of the nutrients in the xylem sap can be utilised.<ref name=Gullan/><ref name=Panizzi>{{cite book |author1=Panizzi, Antônio Ricardo |author2=Parra, José R.P.|title=Insect Bioecology and Nutrition for Integrated Pest Management |url=https://books.google.com/books?id=NqzcD2r3jBYC&pg=PA108 |year=2012 |publisher=CRC Press |isbn=978-1-4398-3708-5 |page=108}}</ref> Some phloem sap feeders selectively mix phloem and xylem sap to control the osmotic potential of the liquid consumed.<ref>{{cite journal |last1=Pompon |first1=Julien |last2=Quiring |first2=Dan |last3=Goyer |first3=Claudia |last4=Giordanengo |first4=Philippe |last5=Pelletier |first5=Yvan |title=A phloem-sap feeder mixes phloem and xylem sap to regulate osmotic potential |journal=Journal of Insect Physiology |date=2011 |volume=57 |issue=9 |pages=1317–1322 |url=https://www.u-picardie.fr/PCP/data/pub/2011-Pompon%20et%20al.J%20Insect%20Physiol.pdf |doi=10.1016/j.jinsphys.2011.06.007 |pmid=21726563 |bibcode=2011JInsP..57.1317P |access-date=2015-08-22 |archive-url=https://web.archive.org/web/20150923191653/https://www.u-picardie.fr/PCP/data/pub/2011-Pompon%20et%20al.J%20Insect%20Physiol.pdf |archive-date=2015-09-23 |url-status=dead }}</ref>
-A striking adaptation to a very dilute diet is found in many hemipterans: a filter chamber, a part of the gut looped back on itself as a countercurrent exchanger, which permits nutrients to be separated from excess water.<ref>{{cite web |title=Digestive System |url=http://bugs.bio.usyd.edu.au/learning/resources/Entomology/internalAnatomy/digestiveSystem.html |website=Entomology |publisher=University of Sydney |access-date=22 August 2015}}</ref> The residue, mostly water with sugars and amino acids, is quickly excreted as sticky "honey dew", notably from aphids but also from other Auchenorrhycha and Sternorrhyncha.<ref>{{cite web|last1=Underwood |first1=D. L. A. |title=Hemipteroids |url=http://web.csulb.edu/~dlunderw/entomology/12_13-Hemipteroids.pdf |publisher=California State University |access-date=22 August 2015}}</ref>
+A striking adaptation to a very dilute diet is found in many hemipterans: a filter chamber, a part of the gut looped back on itself as a countercurrent exchanger, which permits nutrients to be separated from excess water.<ref>{{cite web |title=Digestive System |url=http://bugs.bio.usyd.edu.au/learning/resources/Entomology/internalAnatomy/digestiveSystem.html |website=Entomology |publisher=University of Sydney |access-date=22 August 2015 |archive-date=7 October 2012 |archive-url=https://web.archive.org/web/20121007222727/http://bugs.bio.usyd.edu.au/learning/resources/Entomology/internalAnatomy/digestiveSystem.html |url-status=dead }}</ref> The residue, mostly water with sugars and amino acids, is quickly excreted as sticky "honey dew", notably from aphids but also from other Auchenorrhycha and Sternorrhyncha.<ref>{{cite web |last1=Underwood |first1=D. L. A. |title=Hemipteroids |url=http://web.csulb.edu/~dlunderw/entomology/12_13-Hemipteroids.pdf |publisher=California State University |access-date=22 August 2015 |archive-date=4 March 2016 |archive-url=https://web.archive.org/web/20160304190233/http://web.csulb.edu/~dlunderw/entomology/12_13-Hemipteroids.pdf |url-status=dead }}</ref>
 Some Sternorrhyncha including Psyllids and some aphids are gall formers. These sap-sucking hemipterans inject fluids containing plant hormones into the plant tissues inducing the production of tissue that covers to protects the insect and also act as sinks for nutrition that they feed on. The [[Pachypsylla|hackleberry gall psyllid]] for example, causes a woody gall on the leaf petioles of the [[Celtis occidentalis|hackleberry tree]] it infests,<ref>{{cite web |url=http://entomology.ifas.ufl.edu/creatures/trees/hackberry_psyllid.htm |title=Hackberry petiole gall psyllid |author1=Hall, Donald W. |author2=Butler, Jerry F. |author3=Cromroy, Harvey L. |date=1 September 2014 |work=Featured Creatures |publisher=University of Florida/IFAS |access-date=12 August 2015}}</ref> and the nymph of another psyllid produces a protective [[Lerp (biology)|lerp]] out of hardened honeydew.<ref name=Gullan/>
@@ Line 189: / Line 189: @@
 ====Predators====
-Most other hemipterans are [[predation|predatory]], feeding on other insects, or even small vertebrates. This is true of many aquatic species which are predatory, either as nymphs or adults.<ref name=Discover/> The [[Cermatulus nasalis|predatory shield bug]] for example stabs caterpillars with its beak and sucks out the body fluids.<ref name=Brisbane>{{cite web |url=http://www.brisbaneinsects.com/brisbane_stinkbugs/GlossyShieldBugs.htm |title=Glossy Shield Bug: ''Cermatulus nasalis'' |date=2010 |publisher=Brisbane Insects and Spiders |access-date=16 July 2015}}</ref> The saliva of predatory heteropterans contains [[digestive enzyme]]s such as [[proteinase]] and [[phospholipase]], and in some species also [[amylase]]. The mouthparts of these insects are adapted for predation. There are toothed stylets on the mandibles able to cut into and abrade tissues of their prey. There are further stylets on the maxillae, adapted as tubular canals to inject saliva and to extract the pre-digested and liquified contents of the prey.<ref>{{cite journal |last1=Cohen |first1=Allen C. |title=Feeding Adaptations of Some Predaceous Hemiptera |url=http://insectdiets.com/wp-content/uploads/2012/07/Cohen-47.pdf | journal=Annals of the Entomological Society of America | volume=83 | issue=6 | pages=1215–1223 | date=1990 | doi=10.1093/aesa/83.6.1215}}</ref>
+Most other hemipterans are [[predatory]], feeding on other insects, or even small vertebrates. This is true of many aquatic species which are predatory, either as nymphs or adults.<ref name=Discover/> The [[Cermatulus nasalis|predatory shield bug]] for example stabs caterpillars with its beak and sucks out the body fluids.<ref name=Brisbane>{{cite web |url=http://www.brisbaneinsects.com/brisbane_stinkbugs/GlossyShieldBugs.htm |title=Glossy Shield Bug: ''Cermatulus nasalis'' |date=2010 |publisher=Brisbane Insects and Spiders |access-date=16 July 2015}}</ref> The saliva of predatory heteropterans contains [[digestive enzyme]]s such as [[proteinase]] and [[phospholipase]], and in some species also [[amylase]]. The mouthparts of these insects are adapted for predation. There are toothed stylets on the mandibles able to cut into and abrade tissues of their prey. There are further stylets on the maxillae, adapted as tubular canals to inject saliva and to extract the pre-digested and liquified contents of the prey.<ref>{{cite journal |last1=Cohen |first1=Allen C. |title=Feeding Adaptations of Some Predaceous Hemiptera |url=http://insectdiets.com/wp-content/uploads/2012/07/Cohen-47.pdf | journal=Annals of the Entomological Society of America | volume=83 | issue=6 | pages=1215–1223 | date=1990 | doi=10.1093/aesa/83.6.1215}}</ref>
 ====Haematophagic ectoparasites====
-A few hemipterans are [[Hematophagy|haematophagic]] [[ectoparasite]]s<ref>{{cite journal |last1=Goddard |first1=Jerome |title=Bed Bugs (Cimex lectularius) and Clinical Consequences of Their Bites |journal=JAMA |date=2009 |volume=301 |issue=13 |pages=1358–1366 |doi=10.1001/jama.2009.405 |pmid=19336711|doi-access=free }}</ref>), feeding on the blood of larger animals. These include [[Bed bug (insect)|bedbug]]s and the [[Triatominae|triatomine]] [[kissing bug]]s of the assassin bug family [[Reduviidae]], which can transmit the dangerous ''[[Chagas disease]]''.<ref name="NHM"/> The first known hemipteran to feed in this way on vertebrates was the extinct assassin bug ''[[Triatoma dominicana]]'' found fossilized in [[amber]] and dating back about twenty million years. Faecal pellets fossilised beside it show that it transmitted a disease-causing ''[[Trypanosoma]]'' and the amber included hairs of the likely host, a [[bat]].<ref name=Poinar>{{cite journal |doi= 10.1089/vbz.2005.5.72 |last= Poinar |first=G. |title=''Triatoma dominicana'' sp. n. (Hemiptera: Reduviidae: Triatominae), and ''Trypanosoma antiquus'' sp. n. (Stercoraria: Trypanosomatidae), the First Fossil Evidence of a Triatomine-Trypanosomatid Vector Association |journal= Vector-Borne and Zoonotic Diseases |volume= 5|issue= 1|pages= 72–81|year= 2005 |pmid= 15815152}}</ref>
+A few hemipterans are [[haematophagic]] [[ectoparasite]]s<ref>{{cite journal |last1=Goddard |first1=Jerome |title=Bed Bugs (Cimex lectularius) and Clinical Consequences of Their Bites |journal=JAMA |date=2009 |volume=301 |issue=13 |pages=1358–1366 |doi=10.1001/jama.2009.405 |pmid=19336711|doi-access=free }}</ref>), feeding on the blood of larger animals. These include [[bedbug]]s and the [[Triatominae|triatomine]] [[kissing bug]]s of the assassin bug family [[Reduviidae]], which can transmit the dangerous ''[[Chagas disease]]''.<ref name="NHM"/> The first known hemipteran to feed in this way on vertebrates was the extinct assassin bug ''[[Triatoma dominicana]]'' found fossilized in [[amber]] and dating back about twenty million years. Faecal pellets fossilised beside it show that it transmitted a disease-causing ''[[Trypanosoma]]'' and the amber included hairs of the likely host, a [[bat]].<ref name=Poinar>{{cite journal |doi= 10.1089/vbz.2005.5.72 |last= Poinar |first=G. |title=''Triatoma dominicana'' sp. n. (Hemiptera: Reduviidae: Triatominae), and ''Trypanosoma antiquus'' sp. n. (Stercoraria: Trypanosomatidae), the First Fossil Evidence of a Triatomine-Trypanosomatid Vector Association |journal= Vector-Borne and Zoonotic Diseases |volume= 5|issue= 1|pages= 72–81|year= 2005 |pmid= 15815152}}</ref>
 === As symbionts ===
@@ Line 201: / Line 201: @@
 [[File:Common jassid nymphs and ants02.jpg|thumb|upright=0.6|[[Leafhopper]]s protected by [[meat ant]]s]]
-Some species of [[ant]] protect and farm aphids (Sternorrhyncha) and other sap-sucking hemipterans, gathering and eating the [[Honeydew (secretion)|honeydew]] that these hemipterans secrete. The relationship is [[mutualism (biology)|mutualistic]], as both ant and aphid benefit. Ants such as the yellow anthill ant, ''[[Lasius flavus]]'', breed aphids of at least four species, ''Geoica utricularia'', ''Tetraneura ulmi'', ''Forda marginata'' and ''Forda formicaria'', taking eggs with them when they found a new colony; in return, these aphids are obligately associated with the ant, breeding mainly or wholly asexually inside anthills.<ref name=Ivens>{{cite journal |last1=Ivens |first1=A. B. F. |last2=Kronauer |first2=D. J. C. |last3=Pen |first3=I. |last4=Weissing |first4=F. J. |last5=Boomsma |first5=J. J. |title=Reproduction and dispersal in an ant-associated root aphid community |journal=Molecular Ecology|date=2012 |pages=4257–69 |doi=10.1111/j.1365-294X.2012.05701.x |url=https://www.researchgate.net/publication/229163242 |volume=21 |issue=17 |pmid=22804757|s2cid=34429056 }}</ref> Ants may also protect the plant bugs from their natural enemies, removing the eggs of predatory beetles and preventing access by parasitic wasps.<ref name=Gullan/>
+Some species of [[ant]] protect and farm aphids (Sternorrhyncha) and other sap-sucking hemipterans, gathering and eating the [[Honeydew (secretion)|honeydew]] that these hemipterans secrete. The relationship is [[mutualism (biology)|mutualistic]], as both ant and aphid benefit. Ants such as the yellow anthill ant, ''[[Lasius flavus]]'', breed aphids of at least four species, ''Geoica utricularia'', ''Tetraneura ulmi'', ''Forda marginata'' and ''Forda formicaria'', taking eggs with them when they found a new colony; in return, these aphids are obligately associated with the ant, breeding mainly or wholly asexually inside anthills.<ref name=Ivens>{{cite journal |last1=Ivens |first1=A. B. F. |last2=Kronauer |first2=D. J. C. |last3=Pen |first3=I. |last4=Weissing |first4=F. J. |last5=Boomsma |first5=J. J. |title=Reproduction and dispersal in an ant-associated root aphid community |journal=Molecular Ecology|date=2012 |pages=4257–69 |doi=10.1111/j.1365-294X.2012.05701.x |url=https://www.researchgate.net/publication/229163242 |volume=21 |issue=17 |pmid=22804757|bibcode=2012MolEc..21.4257I |s2cid=34429056 }}</ref> Ants may also protect the plant bugs from their natural enemies, removing the eggs of predatory beetles and preventing access by parasitic wasps.<ref name=Gullan/>
 Some leafhoppers (Auchenorrhyncha) are similarly "milked" by ants. In the Corcovado rain forest of Costa Rica, wasps compete with ants to protect and milk leafhoppers; the leafhoppers preferentially give more honeydew, more often, to the wasps, which are larger and may offer better protection.<ref name="Choe2012">{{cite book |last=Choe |first=Jae |title=Secret Lives of Ants |url=https://books.google.com/books?id=gACBQhye2wsC&pg=PT62|date=14 February 2012 |publisher=JHU Press |isbn=978-1-4214-0521-6 |pages=62–64}}</ref>
@@ Line 209: / Line 209: @@
 {{further|antipredator adaptation}}
-[[File:Reduvius personatus, Masked Hunter Bug nymph camouflaged with sand grains.JPG|thumb|left|[[Reduvius personatus|Masked hunter]] nymph has [[Self-decoration camouflage|camouflaged itself]] with sand grains.]]
+[[File:Reduvius personatus, Masked Hunter Bug nymph camouflaged with sand grains.JPG|thumb|left|[[Masked hunter]] nymph has [[Self-decoration camouflage|camouflaged itself]] with sand grains.]]
-Hemiptera form prey to predators including vertebrates, such as birds, and other invertebrates such as [[ladybird]]s.<ref>{{cite journal |last1=Evans |first1=Edward W. |title=Lady beetles as predators of insects other than Hemiptera |journal=Biological Control |date=November 2009 |volume=51 |issue=2|pages=255–267 |doi=10.1016/j.biocontrol.2009.05.011}}</ref><ref name=Edmunds>{{cite book |last1=Edmunds |first1=M. |title=Defence in Animals |url=https://archive.org/details/defenceinanimals0000edmu |url-access=registration |date=1974 |publisher=Longman |isbn=978-0-582-44132-3 |pages=[https://archive.org/details/defenceinanimals0000edmu/page/172 172], 191–194, 201, 261 and passim}}</ref> In response, hemipterans have evolved [[antipredator adaptation]]s. ''[[Ranatra]]'' may feign death ([[thanatosis]]). Others such as ''[[Carpocoris purpureipennis]]'' secrete [[toxin|toxic]] fluids to ward off arthropod predators; some Pentatomidae such as ''[[Dolycoris]]'' are able to direct these fluids at an attacker. Toxic [[cardenolide]] compounds are accumulated by the heteropteran ''[[Oncopeltus fasciatus]]'' when it consumes [[milkweeds]], while the coreid stinkbug ''[[Amorbus rubiginosus]]'' acquires 2-[[hexenal]] from its food plant, ''Eucalyptus''. Some long-legged bugs [[mimic]] twigs, rocking to and fro to simulate the motion of a plant part in the wind.<ref name=Edmunds/> The nymph of the [[Reduvius personatus|Masked hunter]] bug [[Self-decoration camouflage|camouflages itself]] with sand grains, using its hind legs and tarsal fan to form a double layer of grains, coarser on the outside.<ref>{{cite journal | author=Wierauch C. | year=2006 |title=Anatomy of Disguise: Camouflaging Structures in Nymphs of Some Reduviidae (Heteroptera) |journal= Am. Mus. Novit. | volume=3542 |pages=1–18 | doi=10.1206/0003-0082(2006)3542[1:aodcsi]2.0.co;2| hdl=2246/5820 | s2cid=7894145 | url=https://www.biodiversitylibrary.org/item/280965 }}</ref> The Amazon rain forest cicada ''[[Hemisciera maculipennis]]'' displays bright red [[deimatic]] flash coloration on its hindwings when threatened; the sudden contrast helps to startle predators, giving the cicada time to escape. The coloured patch on the hindwing is concealed at rest by an olive green patch of the same size on the forewing, enabling the insect to switch rapidly from [[crypsis|cryptic]] to deimatic behaviour.<ref>{{cite book |last1=Cott |first1=Hugh B. |title=Adaptive Coloration in Animals |url=https://archive.org/details/adaptivecolorati00cott |date=1940 |publisher=Methuen |pages=[https://archive.org/details/adaptivecolorati00cott/page/375 375–376]}}</ref>{{efn|The green/red flash coloration earns the insect the name of stop and go cicada.<ref>{{cite web |title=Better IDs for E.A. Seguy Cicada Illustrations |url=http://www.cicadamania.com/cicadas/2015/03/ |access-date=24 August 2015}}</ref>}}
+Hemiptera form prey to predators including vertebrates, such as birds, and other invertebrates such as [[ladybird]]s.<ref>{{cite journal |last1=Evans |first1=Edward W. |title=Lady beetles as predators of insects other than Hemiptera |journal=Biological Control |date=November 2009 |volume=51 |issue=2|pages=255–267 |doi=10.1016/j.biocontrol.2009.05.011|bibcode=2009BiolC..51..255E }}</ref><ref name=Edmunds>{{cite book |last1=Edmunds |first1=M. |title=Defence in Animals |url=https://archive.org/details/defenceinanimals0000edmu |url-access=registration |date=1974 |publisher=Longman |isbn=978-0-582-44132-3 |pages=[https://archive.org/details/defenceinanimals0000edmu/page/172 172], 191–194, 201, 261 and passim}}</ref> In response, hemipterans have evolved [[antipredator adaptation]]s. ''[[Ranatra]]'' may feign death ([[thanatosis]]). Others such as ''[[Carpocoris purpureipennis]]'' secrete [[toxin|toxic]] fluids to ward off arthropod predators; some Pentatomidae such as ''[[Dolycoris]]'' are able to direct these fluids at an attacker. Toxic [[cardenolide]] compounds are accumulated by the heteropteran ''[[Oncopeltus fasciatus]]'' when it consumes [[milkweeds]], while the coreid stinkbug ''[[Amorbus rubiginosus]]'' acquires 2-[[hexenal]] from its food plant, ''Eucalyptus''. Some long-legged bugs [[mimic]] twigs, rocking to and fro to simulate the motion of a plant part in the wind.<ref name=Edmunds/> The nymph of the [[Masked hunter]] bug [[Self-decoration camouflage|camouflages itself]] with sand grains, using its hind legs and tarsal fan to form a double layer of grains, coarser on the outside.<ref>{{cite journal | author=Wierauch C. | year=2006 |title=Anatomy of Disguise: Camouflaging Structures in Nymphs of Some Reduviidae (Heteroptera) |journal= Am. Mus. Novit. | volume=3542 |pages=1–18 | doi=10.1206/0003-0082(2006)3542[1:aodcsi]2.0.co;2| hdl=2246/5820 | s2cid=7894145 | url=https://www.biodiversitylibrary.org/item/280965 }}</ref> The Amazon rain forest cicada ''[[Hemisciera maculipennis]]'' displays bright red [[deimatic]] flash coloration on its hindwings when threatened; the sudden contrast helps to startle predators, giving the cicada time to escape. The coloured patch on the hindwing is concealed at rest by an olive green patch of the same size on the forewing, enabling the insect to switch rapidly from [[crypsis|cryptic]] to deimatic behaviour.<ref>{{cite book |last1=Cott |first1=Hugh B. |title=Adaptive Coloration in Animals |url=https://archive.org/details/adaptivecolorati00cott |date=1940 |publisher=Methuen |pages=[https://archive.org/details/adaptivecolorati00cott/page/375 375–376]}}</ref>{{efn|The green/red flash coloration earns the insect the name of stop and go cicada.<ref>{{cite web |title=Better IDs for E.A. Seguy Cicada Illustrations |url=http://www.cicadamania.com/cicadas/2015/03/ |access-date=24 August 2015}}</ref>}}
-[[File:Pyrrhocoris apterus group.jpg|thumb|Firebugs, ''[[Pyrrhocoris apterus]]'', [[antipredator adaptation|protect themselves from predators]] with bright [[aposematism|aposematic]] warning coloration, and by aggregating in a group.]]
+[[File:Pyrrhocoris apterus group.jpg|thumb|Firebugs, ''[[Pyrrhocoris apterus]]'', [[antipredator adaptation|protect themselves from predators]] with bright [[aposematic]] warning coloration, and by aggregating in a group.]]
-Some hemipterans such as [[Pyrrhocoris apterus|firebugs]] have bold [[aposematic]] warning coloration, often red and black, which appear to deter [[passerine]] birds.<ref name="Wheeler100">{{cite book |last=Wheeler |first=Alfred George |title=Biology of the Plant Bugs (Hemiptera: Miridae): Pests, Predators, Opportunists |url=https://books.google.com/books?id=0szFXwGVlo4C&pg=PA100 |year=2001 |publisher=Cornell University Press |isbn=978-0-8014-3827-1 |pages=100ff}}</ref><ref>{{cite journal |last1=Exnerova |first1=Alice |display-authors=etal|title=Reactions of passerine birds to aposematic and non-aposematic firebugs (Pyrrhocoris apterus ; Heteroptera) |journal=Biological Journal of the Linnean Society |date=April 2003 |volume=78 |issue=4 |pages=517–525 |doi=10.1046/j.0024-4066.2002.00161.x|doi-access=free }}</ref> Many hemipterans including aphids, scale insects and especially the planthoppers secrete wax to protect themselves from threats such as fungi, parasitoidal insects and predators, as well as abiotic factors like desiccation.<ref>{{cite journal |last1=Lucchi |first1=Andrea |last2=Mazzoni |first2=Emanuele |title=Wax Production in Adults of Planthoppers (Homoptera: Fulgoroidea) with Particular Reference to Metcalfa pruinosa (Flatidae) |journal=Annals of the Entomological Society of America |date=2004 |volume=97 |issue=6 |pages=1294–1298 |url=http://ag.udel.edu/delpha/2926.pdf |doi=10.1603/0013-8746(2004)097[1294:wpiaop]2.0.co;2|doi-access=free }}</ref> Hard waxy coverings are especially important in the sedentary Sternorrhyncha such as scale insects, which have no means of escaping from predators; other Sternorrhyncha evade detection and attack by creating and living inside plant galls.<ref name="GrimaldiEngel2005"/> Nymphal Cicadoidea and Cercopoidea have glands attached to the [[Malpighian tubule]]s in their proximal segment that produce mucopolysaccharides, which form the froth around spittlebugs, offering a measure of protection.<ref>{{cite journal|title=Structure and Function of the Malpighian Tubules, and Related Behaviors in Juvenile Cicadas: Evidence of Homology with Spittlebugs (Hemiptera: Cicadoidea & Cercopoidea)|first=Roman A. |last=Rakitov |doi=10.1078/0044-5231-00025 | journal=Zoologischer Anzeiger| volume =241|issue= 2|year=2002|pages =117–130}}</ref>
+Some hemipterans such as [[Pyrrhocoris apterus|firebugs]] have bold [[aposematic]] warning coloration, often red and black, which appear to deter [[passerine]] birds.<ref name="Wheeler100">{{cite book |last=Wheeler |first=Alfred George |title=Biology of the Plant Bugs (Hemiptera: Miridae): Pests, Predators, Opportunists |url=https://books.google.com/books?id=0szFXwGVlo4C&pg=PA100 |year=2001 |publisher=Cornell University Press |isbn=978-0-8014-3827-1 |pages=100ff}}</ref><ref>{{cite journal |last1=Exnerova |first1=Alice |display-authors=etal|title=Reactions of passerine birds to aposematic and non-aposematic firebugs (Pyrrhocoris apterus ; Heteroptera) |journal=Biological Journal of the Linnean Society |date=April 2003 |volume=78 |issue=4 |pages=517–525 |doi=10.1046/j.0024-4066.2002.00161.x|doi-access=free }}</ref> Many hemipterans including aphids, scale insects and especially the planthoppers secrete wax to protect themselves from threats such as fungi, parasitoidal insects and predators, as well as abiotic factors like desiccation.<ref>{{cite journal |last1=Lucchi |first1=Andrea |last2=Mazzoni |first2=Emanuele |title=Wax Production in Adults of Planthoppers (Homoptera: Fulgoroidea) with Particular Reference to Metcalfa pruinosa (Flatidae) |journal=Annals of the Entomological Society of America |date=2004 |volume=97 |issue=6 |pages=1294–1298 |url=http://ag.udel.edu/delpha/2926.pdf |doi=10.1603/0013-8746(2004)097[1294:wpiaop]2.0.co;2|doi-access=free }}</ref> Hard waxy coverings are especially important in the sedentary Sternorrhyncha such as scale insects, which have no means of escaping from predators; other Sternorrhyncha evade detection and attack by creating and living inside plant galls.<ref name="GrimaldiEngel2005"/> Nymphal Cicadoidea and Cercopoidea have glands attached to the [[Malpighian tubule]]s in their proximal segment that produce mucopolysaccharides, which form the froth around spittlebugs, offering a measure of protection.<ref>{{cite journal|title=Structure and Function of the Malpighian Tubules, and Related Behaviors in Juvenile Cicadas: Evidence of Homology with Spittlebugs (Hemiptera: Cicadoidea & Cercopoidea)|first=Roman A. |last=Rakitov |doi=10.1078/0044-5231-00025 | journal=Zoologischer Anzeiger| volume =241|issue= 2|year=2002|pages =117–130|bibcode=2002ZooAn.241..117R }}</ref>
 Parental care is found in many species of Hemiptera especially in members of the Membracidae and numerous Heteroptera. In many species of shield bug, females stand guard over their egg clusters to protect them from egg parasitoids and predators.<ref>{{cite book|chapter= Maternal care in the Hemiptera: ancestry, alternatives, and current adaptive value| author1=Tallamy, Douglas W.|author2= Schaefer, Carl| pages=91–115 | publisher=Cambridge University Press| title=The Evolution of Social Behaviour in Insects and Arachnids |year=1997|editor=Jae C. Choe |editor2=Bernard J. Crespi}}</ref> In the aquatic Belostomatidae, females lay their eggs on the back of the male which guards the eggs.<ref>{{cite journal| title=Parental care and foraging ability in male water bugs (Belostoma flumineum)|author1=Crowl, T.A.| author2=Alexander Jr., J.E.| journal=Canadian Journal of Zoology| year= 1989| volume= 67|issue=2|pages=513–515|doi=10.1139/z89-074}}</ref> Protection provided by ants is common in the Auchenorrhyncha.<ref name=Gullan/>
@@ Line 223: / Line 223: @@
 {{further|Insects in culture}}
-[[Image:Icerya purchasi 1435060.jpg|thumb|Colony of [[Icerya purchasi|cottony cushion scale]], a pest of  [[citrus]] fruits]]
+[[Image:Icerya purchasi 1435060.jpg|thumb|Colony of [[cottony cushion scale]], a pest of  [[citrus]] fruits]]
 === As pests ===
-Although many species of Hemiptera are significant pests of crops and garden plants, including many species of [[aphid]] and [[scale insect]]s, other species are harmless. The damage done is often not so much the deprivation of the plant of its sap, but the fact that they transmit serious viral diseases between plants.<ref name=AES>{{cite web |url=http://www.amentsoc.org/insects/fact-files/orders/ |title=Insect groups (Orders) |publisher=Amateur Entomologists' Society |access-date=16 July 2015}}</ref> They often produce copious amounts of [[honeydew (secretion)|honeydew]] which encourages the growth of [[Sooty mold|sooty mould]].<ref>{{cite web |url=https://www.rhs.org.uk/advice/profile?PID=770 |title=Sooty moulds |publisher=Royal Horticultural Society |access-date=16 July 2015}}</ref> Significant pests include the [[Icerya purchasi|cottony cushion scale]], a pest of [[citrus]] fruit trees,<ref>{{cite web |url=http://gardenbees.com/biological%20control/revolution.htm |title=Cottony cushion scale: The pest that launched a revolution in pest control methods |date=10 August 2003 |author=David L. Green |access-date=16 July 2015}}</ref> the [[Myzus persicae|green peach aphid]] and other aphids which attack crops worldwide and transmit diseases,<ref>{{cite web|url=http://entnemdept.ufl.edu/creatures/veg/aphid/green_peach_aphid.htm|title=Featured creatures|last=Capinera|first=John L. |date=October 2005|work=University of Florida website – Department of Entomology and Nematology|publisher=University of Florida|access-date=16 July 2015}}</ref> and [[Jumping plant louse#Status as pests|jumping plant lice]] which are often host plant-specific and transmit diseases.{{Citation needed|date=December 2019|reason=Previous citation was removed as it was associated with a predatory publisher.}}
+Although many species of Hemiptera are significant pests of crops and garden plants, including many species of [[aphid]] and [[scale insect]]s, other species are harmless. The damage done is often not so much the deprivation of the plant of its sap, but the fact that they transmit serious viral diseases between plants.<ref name=AES>{{cite web |url=http://www.amentsoc.org/insects/fact-files/orders/ |title=Insect groups (Orders) |publisher=Amateur Entomologists' Society |access-date=16 July 2015}}</ref> They often produce copious amounts of [[honeydew (secretion)|honeydew]] which encourages the growth of [[sooty mould]].<ref>{{cite web |url=https://www.rhs.org.uk/advice/profile?PID=770 |title=Sooty moulds |publisher=Royal Horticultural Society |access-date=16 July 2015}}</ref> Significant pests include the [[cottony cushion scale]], a pest of [[citrus]] fruit trees,<ref>{{cite web |url=http://gardenbees.com/biological%20control/revolution.htm |title=Cottony cushion scale: The pest that launched a revolution in pest control methods |date=10 August 2003 |author=David L. Green |access-date=16 July 2015}}</ref> the [[green peach aphid]] and other aphids which attack crops worldwide and transmit diseases,<ref>{{cite web|url=http://entnemdept.ufl.edu/creatures/veg/aphid/green_peach_aphid.htm|title=Featured creatures|last=Capinera|first=John L. |date=October 2005|work=University of Florida website – Department of Entomology and Nematology|publisher=University of Florida|access-date=16 July 2015}}</ref> and [[Jumping plant louse#Status as pests|jumping plant lice]] which are often host plant-specific and transmit diseases.{{Citation needed|date=December 2019|reason=Previous citation was removed as it was associated with a predatory publisher.}}
 === For pest control ===
 [[File:Spined soldier bug.webm|thumb|right|thumbtime=40|genus [[Podisus]] [[Spined soldier bug]] eggs and then the recently hatched first instar bugs]]
-Members of the families [[Reduviidae]], [[Phymatidae]] and [[Nabidae]] are obligate predators.  Some predatory species are used in [[biological pest control]]; these include various [[Nabidae|nabids]],<ref>{{cite journal |url=http://www.entomology.wisc.edu/mbcn/kyf402.html |title=Know Your Friends: Damsel Bugs |author=Mahr, Susan |journal=Biological Control News |year=1997 |volume=IV |issue=2 |access-date=16 July 2015}}</ref> and even some members of families that are primarily phytophagous, such as the genus ''[[Geocoris]]'' in the family [[Lygaeidae]].<ref>{{cite web |editor1=Weeden, Catherine R. |editor2=Shelton, Anthony M. |editor3=Hoffman, Michael P. |work=Biological Control: A Guide to Natural Enemies in North America |url=http://www.nysaes.cornell.edu/ent/biocontrol/predators/Geocoris.html |author=James Hagler |publisher=Cornell University |title=''Geocoris'' spp. (Heteroptera: Lygaeidae): Bigeyed Bug |access-date=16 July 2015}}</ref> Other hemipterans are [[omnivore]]s, alternating between a plant-based and an animal-based diet. For example, ''[[Dicyphus hesperus]]'' is used to control [[whitefly]] on [[tomato]]es but also sucks sap, and if deprived of plant tissues will die even if in the presence of whiteflies.<ref>{{cite journal |author1=Torres, Jorge Braz |author2=Boyd, David W. |year=2009 |title=Zoophytophagy in predatory Hemiptera |journal=Brazilian Archives of Biology and Technology |volume=52 |issue=5 |doi=10.1590/S1516-89132009000500018 |pages=1199–1208|doi-access=free }}</ref>
+Members of the families [[Reduviidae]], [[Phymatidae]] and [[Nabidae]] are obligate predators.  Some predatory species are used in [[biological pest control]]; these include various [[Nabidae|nabids]],<ref>{{cite journal |url=http://www.entomology.wisc.edu/mbcn/kyf402.html |title=Know Your Friends: Damsel Bugs |author=Mahr, Susan |journal=Biological Control News |year=1997 |volume=IV |issue=2 |access-date=16 July 2015 |archive-date=20 April 2018 |archive-url=https://web.archive.org/web/20180420161131/http://www.entomology.wisc.edu/mbcn/kyf402.html |url-status=dead }}</ref> and even some members of families that are primarily phytophagous, such as the genus ''[[Geocoris]]'' in the family [[Lygaeidae]].<ref>{{cite web |editor1=Weeden, Catherine R. |editor2=Shelton, Anthony M. |editor3=Hoffman, Michael P. |work=Biological Control: A Guide to Natural Enemies in North America |url=http://www.nysaes.cornell.edu/ent/biocontrol/predators/Geocoris.html |author=James Hagler |publisher=Cornell University |title=''Geocoris'' spp. (Heteroptera: Lygaeidae): Bigeyed Bug |access-date=16 July 2015}}</ref> Other hemipterans are [[omnivore]]s, alternating between a plant-based and an animal-based diet. For example, ''[[Dicyphus hesperus]]'' is used to control [[whitefly]] on [[tomato]]es but also sucks sap, and if deprived of plant tissues will die even if in the presence of whiteflies.<ref>{{cite journal |author1=Torres, Jorge Braz |author2=Boyd, David W. |year=2009 |title=Zoophytophagy in predatory Hemiptera |journal=Brazilian Archives of Biology and Technology |volume=52 |issue=5 |doi=10.1590/S1516-89132009000500018 |pages=1199–1208|doi-access=free }}</ref>
 The [[spined soldier bug]] (''[[Podisus]] maculiventris'') sucks body fluids from several pests including the larvae of the [[Colorado beetle]] and the [[Mexican bean beetle]].<ref>{{cite book |title=Predatory Heteroptera: their ecology and use in biological control |editor=Coll, M. |editor2=Ruberson, J.R. |year=1998 |publisher=Entomological Society of America |isbn=978-0-938522-62-1 |url=http://www.cabdirect.org/abstracts/19981105845.html;jsessionid=B3E7BAA960BF7059CA04F60707516915 }}{{Dead link|date=January 2020 |bot=InternetArchiveBot |fix-attempted=yes }}</ref>
@@ Line 247: / Line 247: @@
 [[File:Bed bug, Cimex lectularius.jpg|thumb|left|Bed bug nymph, ''[[Cimex lectularius]]'', engorged with human blood]]
-[[Chagas disease]] is a modern-day tropical disease caused by ''[[Trypanosoma cruzi]]'' and transmitted by [[Triatominae|kissing bugs]], so-called because they suck human blood from around the lips while a person sleeps.<ref name=CDC>{{cite web|publisher=Centers for Disease Control (CDC) |url= https://www.cdc.gov/dpdx/trypanosomiasisAmerican/ |title= American Trypanosomiasis |access-date=17 July 2015}}</ref>
+[[Chagas disease]] is a modern-day tropical disease caused by ''[[Trypanosoma cruzi]]'' and transmitted by [[kissing bugs]], so-called because they suck human blood from around the lips while a person sleeps.<ref name=CDC>{{cite web|publisher=Centers for Disease Control (CDC) |url= https://www.cdc.gov/dpdx/trypanosomiasisAmerican/ |title= American Trypanosomiasis |access-date=17 July 2015}}</ref>
-The [[Bed bug (insect)|bed bug]], ''Cimex lectularius'', is an external [[parasite]] of humans. It lives in bedding and is mainly active at night, feeding on human blood, generally without being noticed.<ref name=NEJM2009>{{cite journal |last=Goddard |first=J |author2=deShazo, R |title=Bed bugs (Cimex lectularius) and clinical consequences of their bites |journal=JAMA: The Journal of the American Medical Association |date=1 April 2009 |volume=301 |issue=13 |pages=1358–66 |pmid=19336711 |doi=10.1001/jama.2009.405|doi-access=free }}</ref><ref name=bioofbedbug>{{Cite journal| last1=Reinhardt | first1=Klaus| last2=Siva-Jothy| first2=Michael T.| title=Biology of the Bed Bugs (Cimicidae) | journal=[[Annual Review of Entomology]]| volume=52| pages=351–374|date=Jan 2007| pmid=16968204 | url=http://www.falw.vu.nl/nl/Images/siva%202006_tcm19-30750.pdf |doi=10.1146/annurev.ento.52.040306.133913 |access-date=26 May 2010 | archive-url=https://web.archive.org/web/20100705084911/http://www.falw.vu.nl/nl/Images/siva%202006_tcm19-30750.pdf| archive-date=5 July 2010}}</ref> Bed bugs [[Mating|mate]] by [[traumatic insemination]]; the male pierces the female's abdomen and injects his sperm into a secondary genital structure, the [[spermalege]]. The sperm travel in the female's blood ([[haemolymph]]) to [[female sperm storage|sperm storage]] structures (seminal conceptacles); they are released from there to fertilise her eggs inside her [[ovaries]].<ref name=bioofbedbug/><ref name="carayon1959">Carayon, J. 1959 Insémination par "spermalège" et cordon conducteur de spermatozoids chez Stricticimex brevispinosus Usinger (Heteroptera, Cimicidae). Rev. Zool. Bot. Afr. 60, 81–104.</ref>
+The [[Cimex lectularius|bed bug]], ''Cimex lectularius'', is an external [[parasite]] of humans. It lives in bedding and is mainly active at night, feeding on human blood, generally without being noticed.<ref name=NEJM2009>{{cite journal |last=Goddard |first=J |author2=deShazo, R |title=Bed bugs (Cimex lectularius) and clinical consequences of their bites |journal=JAMA: The Journal of the American Medical Association |date=1 April 2009 |volume=301 |issue=13 |pages=1358–66 |pmid=19336711 |doi=10.1001/jama.2009.405|doi-access=free }}</ref><ref name=bioofbedbug>{{Cite journal| last1=Reinhardt | first1=Klaus| last2=Siva-Jothy| first2=Michael T.| title=Biology of the Bed Bugs (Cimicidae) | journal=[[Annual Review of Entomology]]| volume=52| pages=351–374|date=Jan 2007| pmid=16968204 | url=http://www.falw.vu.nl/nl/Images/siva%202006_tcm19-30750.pdf |doi=10.1146/annurev.ento.52.040306.133913 |access-date=26 May 2010 | archive-url=https://web.archive.org/web/20100705084911/http://www.falw.vu.nl/nl/Images/siva%202006_tcm19-30750.pdf| archive-date=5 July 2010}}</ref> Bed bugs [[Mating|mate]] by [[traumatic insemination]]; the male pierces the female's abdomen and injects his sperm into a secondary genital structure, the [[spermalege]]. The sperm travel in the female's blood ([[haemolymph]]) to [[sperm storage]] structures (seminal conceptacles); they are released from there to fertilise her eggs inside her [[ovaries]].<ref name=bioofbedbug/><ref name="carayon1959">Carayon, J. 1959 Insémination par "spermalège" et cordon conducteur de spermatozoids chez Stricticimex brevispinosus Usinger (Heteroptera, Cimicidae). Rev. Zool. Bot. Afr. 60, 81–104.</ref>
 === As food ===
@@ Line 255: / Line 255: @@
 [[File:deepfried cicada.jpg|thumb|Deep-fried [[cicada]]s, ''Cryptotympana atrata'', in Chinese [[Shandong cuisine]]]]
-Some larger hemipterans such as [[cicada]]s are used as food in Asian countries such as [[China]],<ref>{{cite web |last1=Greenaway |first1=Twilight |url=http://www.smithsonianmag.com/arts-culture/the-best-way-to-handle-the-coming-cicada-invasion-heat-up-the-deep-fryer-19372006 |title=The Best Way to Handle the Coming Cicada Invasion? Heat Up the Deep Fryer |publisher=Smithsonian |access-date=12 July 2015}}</ref> and they are much esteemed in Malawi and other African countries. Insects have a high protein content and good food conversion ratios, but most hemipterans are too small to be a useful component of the human diet.<ref>{{cite news |title=Could insects be the wonder food of the future? |author=Anthes, Emily |url=http://www.bbc.com/future/story/20141014-time-to-put-bugs-on-the-menu |newspaper=BBC |date=14 October 2014 |access-date=17 July 2015}}</ref> At least nine species of Hemiptera are eaten worldwide.<ref>{{cite web| url=http://www.fao.org/docrep/018/i3253e/i3253e00.htm |title=Future Prospects for Food and Feed Security |access-date=13 April 2016}}</ref>
+Some larger hemipterans such as [[cicada]]s are used as food in Asian countries such as [[China]],<ref>{{cite web |last1=Greenaway |first1=Twilight |url=http://www.smithsonianmag.com/arts-culture/the-best-way-to-handle-the-coming-cicada-invasion-heat-up-the-deep-fryer-19372006 |title=The Best Way to Handle the Coming Cicada Invasion? Heat Up the Deep Fryer |publisher=Smithsonian |access-date=12 July 2015}}</ref> and they are much esteemed in Malawi and other African countries. Insects have a high protein content and good food conversion ratios, but most hemipterans are too small to be a useful component of the human diet.<ref>{{cite news |title=Could insects be the wonder food of the future? |author=Anthes, Emily |url=http://www.bbc.com/future/story/20141014-time-to-put-bugs-on-the-menu |newspaper=BBC |date=14 October 2014 |access-date=17 July 2015}}</ref> At least nine species of Hemiptera are eaten worldwide.<ref>{{cite web | url=http://www.fao.org/docrep/018/i3253e/i3253e00.htm | title=Future Prospects for Food and Feed Security | access-date=13 April 2016 | archive-date=4 April 2016 | archive-url=https://web.archive.org/web/20160404091437/http://www.fao.org/docrep/018/i3253e/i3253e00.htm | url-status=dead }}</ref>
 === In art and literature ===
@@ Line 269: / Line 269: @@
 === Threats ===
-Large-scale cultivation of the oil palm ''[[Elaeis guineensis]]'' in the [[Amazon basin]] damages freshwater habitats and reduces the diversity of aquatic and semi-aquatic Heteroptera.<ref>{{cite journal |last1=Cunha |first1=Erlane Jose |last2=Montag |first2=Luciano |last3=Juen |first3=Leandro |title=Oil palm crops effects on environmental integrity of Amazonian streams and Heteropteran (Hemiptera) species diversity |journal=Ecological Indicators |date=2015 |volume=52 |pages=422–429 |doi=10.1016/j.ecolind.2014.12.024}}</ref> Climate change may be affecting the global migration of hemipterans including the potato leafhopper, ''[[Empoasca fabae]]''. Warming is correlated with the severity of potato leafhopper infestation, so increased warming may worsen infestations in future.<ref>{{cite journal |last1=Baker |first1=Mitchell B. |last2=Venugopal |first2=P. Dilip |last3=Lamp |first3=William O. |title=Climate Change and Phenology: Empoasca fabae (Hemiptera: Cicadellidae) Migration and Severity of Impact |journal=PLOS ONE |date=2015|volume=10 |issue=5 |page=e0124915 |doi=10.1371/journal.pone.0124915 |pmid=25970705 |pmc=4430490|bibcode=2015PLoSO..1024915B |doi-access=free }}</ref>
+Large-scale cultivation of the oil palm ''[[Elaeis guineensis]]'' in the [[Amazon basin]] damages freshwater habitats and reduces the diversity of aquatic and semi-aquatic Heteroptera.<ref>{{cite journal |last1=Cunha |first1=Erlane Jose |last2=Montag |first2=Luciano |last3=Juen |first3=Leandro |title=Oil palm crops effects on environmental integrity of Amazonian streams and Heteropteran (Hemiptera) species diversity |journal=Ecological Indicators |date=2015 |volume=52 |pages=422–429 |doi=10.1016/j.ecolind.2014.12.024|bibcode=2015EcInd..52..422C }}</ref> Climate change may be affecting the global migration of hemipterans including the potato leafhopper, ''[[Empoasca fabae]]''. Warming is correlated with the severity of potato leafhopper infestation, so increased warming may worsen infestations in future.<ref>{{cite journal |last1=Baker |first1=Mitchell B. |last2=Venugopal |first2=P. Dilip |last3=Lamp |first3=William O. |title=Climate Change and Phenology: Empoasca fabae (Hemiptera: Cicadellidae) Migration and Severity of Impact |journal=PLOS ONE |date=2015|volume=10 |issue=5 |page=e0124915 |doi=10.1371/journal.pone.0124915 |pmid=25970705 |pmc=4430490|bibcode=2015PLoSO..1024915B |doi-access=free }}</ref>
 == Notes ==