Entomologists reserve the termbug for Hemiptera or Heteroptera,[5] which does not include other arthropods or insects of other orders such asants,bees,beetles, orbutterflies. In some varieties of English, allterrestrialarthropods (including non-insectarachnids andmyriapods) also fall under thecolloquial understanding ofbug.[a]
Most hemipterans feed on plants, using their sucking and piercing mouthparts to extractplant sap. Some are bloodsucking, orhematophagous, while others arepredators that feed on other insects or smallinvertebrates. 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 arehemimetabolous, with youngnymphs that somewhat resemble adults. Many aphids are capable ofparthenogenesis, 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 significantagricultural pests, damaging crops by sucking the sap. Others harm humans more directly asvectors of serious viral diseases. Thebed bug is a persistent parasite of humans, and somekissing bugs can transmitChagas disease. Some species have been used forbiological control of insect pests or of invasive plants. A few hemipterans, have been cultivated for the extraction of dyestuffs such ascochineal andcarmine, and forshellac.Cicadas have been used as food, and have appeared in literature since theIliad inAncient Greece.
Hemiptera is the largest order ofhemimetabolous insects (not undergoing complete metamorphosis; though some examples such as malescale insects do undergo a form of complete metamorphosis[11]), containing over 95,000 named species. Other insect orders with more species are allholometabolous, meaning they have apupal stage and undergo completemetamorphosis. The majority of species are terrestrial, including a number of important agricultural pests, but some are found in freshwater habitats. These include thewater boatmen,backswimmers,pond skaters, andgiant water bugs.[12]
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 therostrum. "Homoptera" was established asparaphyletic group and an obsolete name.[17][18][19]
Thecladogram shows Hemiptera's placement withinParaneoptera, as well as how Hemiptera's four suborders are related. English names are given in parentheses where possible.[13]
The defining feature of hemipterans is their "beak" in which the modifiedmandibles andmaxillae form a "stylet" which is sheathed within a modifiedlabium. The stylet is capable of piercingtissues and sucking liquids, while the labium supports it. The stylet contains a channel for the outward movement ofsaliva and another for the inward movement of liquid food. A salivary pump drives saliva into the prey; acibarial 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 typicallyplant sap, but some hemipterans such asassassin bugs arepredators.[29][30]
Both herbivorous and predatory hemipterans injectenzymes to begin digestion extra-orally (before the food is taken into the body). These enzymes includeamylase to hydrolysestarch,polygalacturonase to weaken the toughcell walls of plants, andproteinases to break down proteins.[31]
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 somePhthiraptera, but for other reasons they generally are easy to recognize as non-hemipteran. Similarly, the mouthparts ofSiphonaptera, someDiptera andThysanoptera 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 includecockroaches andpsocids, both of which have longer, many-segmented antennae, and somebeetles, but these have fully hardened forewings which do not overlap.[32]
Theforewings of Hemiptera are either entirely membranous, as in theSternorrhyncha andAuchenorrhyncha, or partially hardened, as in mostHeteroptera. The name "Hemiptera" is from theGreekἡμι- (hemi; "half") andπτερόν (pteron; "wing"), referring to the forewings of many heteropteranswhich are hardened near the base, but membranous at the ends. Wings modified in this manner are termedhemelytra (singular:hemelytron), by analogy with the completely hardenedelytra ofbeetles, and occur only in the suborderHeteroptera. 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 ofSternorrhyncha andAuchenorrhyncha),[33] or held flat on the back, with the ends overlapping (typical ofHeteroptera).[12] Theantennae in Hemiptera typically consist of four or five segments, although they can still be quite long, and the tarsi of thelegs have two or three segments.[34]
Many hemipterans can produce sound for communication.[35] The "song" of male cicadas, the loudest of any insect, is produced bytymbal 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.[36]
Hemipterans arehemimetabolous, meaning that they do not undergometamorphosis, the complete change of form between alarval phase and anadult phase. Instead, their young are callednymphs, and resemble the adults to a greater or lesser degree. The nymphs moult several times as they grow, and eachinstar 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 interveningpupal stage as inholometabolous insects.[39]
Manyaphids areparthenogenetic during part of the life cycle, such that females can produce unfertilized eggs, which areclones 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 alsoviviparous: the young are born live rather than laid as eggs. These adaptations enable aphids to reproduce extremely rapidly when conditions are suitable.[40]
Hemipterans make use of a variety of modes oflocomotion including swimming, skating on a water surface and jumping, as well as walking and flying like other insects.[41][42][43]
Several families of Heteroptera arewater bugs, adapted to anaquatic lifestyle, such as thewater boatmen (Corixidae),water scorpions (Nepidae), andbackswimmers (Notonectidae). They are mostly predatory, and have legs adapted aspaddles to help the animal move through the water.[41] Thepondskaters or water striders (Gerridae) are also associated with water, but use thesurface tension of standing water to keep them above the surface;[44] they include the sea skaters in the genusHalobates, the only truly marine group of insects.[41]
Marangoni effect propulsion exploits the change insurface tension when a soap-likesurfactant is released on to a water surface, in the same way that a toy soap boat propels itself. Water bugs in the genusMicrovelia (Veliidae) can travel at up to 17 cm/s, twice as fast as they can walk, by this means.[44]
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.[42] Wing-length polymorphism is notably rare in tree-living Hemiptera.[45]
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 tocatapult the insect into the air.[46] 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.[43]
Instead of relying on any form of locomotion, most Sternorrhyncha females aresedentary or completely sessile, attached to their host plants by their thin feeding stylets which cannot be taken out of the plant quickly.[47]
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 aremonophages, being host specific and only found on one plant taxon, others areoligophages, feeding on a few plant groups, while others again are less discriminatingpolyphages and feed on many species of plant.[30] The relationship between hemipterans and plants appears to be ancient, with piercing and sucking of plants evident in theEarly Devonian period.[48]
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.[49] 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 asparthenogenetically reproducing females on a herbaceous plant.[50]
A twig wilting bug (Coreidae) piercing and sucking sap from aZinnia
Phloem sap, which has a higher concentration of sugars and nitrogen, is under positive pressure unlike the more dilutexylem 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-vascularmesophyll 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.[51]
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.[52] Xylem feeders tend to be polyphagous;[53] to overcome the negative pressure of xylem requires a special cibarial pump.[54]
Phloem feeding hemiptera typically have symbiotic micro-organisms in their gut that help to convertamino acids. Phloem feeders produce honeydew from their anus. A variety of organisms that feed on honeydew form symbiotic associations with phloem-feeders.[55][56] 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 containsmonosaccharides rather thansucrose, 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.[30][57] Some phloem sap feeders selectively mix phloem and xylem sap to control the osmotic potential of the liquid consumed.[58]
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.[59] 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.[60]
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. Thehackleberry gall psyllid for example, causes a woody gall on the leaf petioles of thehackleberry tree it infests,[61] and the nymph of another psyllid produces a protectivelerp out of hardened honeydew.[30]
Most other hemipterans arepredatory, feeding on other insects, or even small vertebrates. This is true of many aquatic species which are predatory, either as nymphs or adults.[34] Thepredatory shield bug for example stabs caterpillars with its beak and sucks out the body fluids.[62] The saliva of predatory heteropterans containsdigestive enzymes such asproteinase andphospholipase, and in some species alsoamylase. 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.[63]
A few hemipterans arehaematophagicectoparasites[64]), feeding on the blood of larger animals. These includebedbugs and thetriatominekissing bugs of the assassin bug familyReduviidae, which can transmit the dangerousChagas disease.[41] The first known hemipteran to feed in this way on vertebrates was the extinct assassin bugTriatoma dominicana found fossilized inamber and dating back about twenty million years. Faecal pellets fossilised beside it show that it transmitted a disease-causingTrypanosoma and the amber included hairs of the likely host, abat.[65]
Some species ofant protect and farm aphids (Sternorrhyncha) and other sap-sucking hemipterans, gathering and eating thehoneydew that these hemipterans secrete. The relationship ismutualistic, 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 andForda 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.[66] Ants may also protect the plant bugs from their natural enemies, removing the eggs of predatory beetles and preventing access by parasitic wasps.[30]
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.[67]
Hemiptera form prey to predators including vertebrates, such as birds, and other invertebrates such asladybirds.[68][69] In response, hemipterans have evolvedantipredator adaptations.Ranatra may feign death (thanatosis). Others such asCarpocoris purpureipennis secretetoxic fluids to ward off arthropod predators; some Pentatomidae such asDolycoris are able to direct these fluids at an attacker. Toxiccardenolide compounds are accumulated by the heteropteranOncopeltus fasciatus when it consumesmilkweeds, while the coreid stinkbugAmorbus rubiginosus acquires 2-hexenal from its food plant,Eucalyptus. Some long-legged bugsmimic twigs, rocking to and fro to simulate the motion of a plant part in the wind.[69] The nymph of theMasked hunter bugcamouflages itself with sand grains, using its hind legs and tarsal fan to form a double layer of grains, coarser on the outside.[70] The Amazon rain forest cicadaHemisciera maculipennis displays bright reddeimatic 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 fromcryptic to deimatic behaviour.[71][b]
Some hemipterans such asfirebugs have boldaposematic warning coloration, often red and black, which appear to deterpasserine birds.[73][74] 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.[75] 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.[47] Nymphal Cicadoidea and Cercopoidea have glands attached to theMalpighian tubules in their proximal segment that produce mucopolysaccharides, which form the froth around spittlebugs, offering a measure of protection.[76]
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.[77] In the aquatic Belostomatidae, females lay their eggs on the back of the male which guards the eggs.[78] Protection provided by ants is common in the Auchenorrhyncha.[30]
Although many species of Hemiptera are significant pests of crops and garden plants, including many species ofaphid andscale 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.[79] They often produce copious amounts ofhoneydew which encourages the growth ofsooty mould.[80] Significant pests include thecottony cushion scale, a pest ofcitrus fruit trees,[81] thegreen peach aphid and other aphids which attack crops worldwide and transmit diseases,[82] andjumping plant lice which can be plant-specific and transmit diseases, as with theAsian citrus psyllid which transmitscitrus greening disease.[83]
Members of the familiesReduviidae,Phymatidae andNabidae are obligate predators. Some predatory species are used inbiological pest control; these include variousnabids,[84] and even some members of families that are primarily phytophagous, such as the genusGeocoris in the familyLygaeidae.[85] Other hemipterans areomnivores, alternating between a plant-based and an animal-based diet. For example,Dicyphus hesperus is used to controlwhitefly ontomatoes but also sucks sap, and if deprived of plant tissues will die even if in the presence of whiteflies.[86]
Other hemipterans have positive uses for humans, such as in the production of the dyestuffcarmine (cochineal). The FDA has created guidelines for how to declare when it has been added to a product.[88] The scale insectDactylopius coccus produces the brilliant red-colouredcarminic 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.[89] A similar number oflac bugs are needed to make a kilogram ofshellac, a brush-on colourant and wood finish.[90] 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.[91]
Chagas disease is a modern-day tropical disease caused byTrypanosoma cruzi and transmitted bykissing bugs, so-called because they suck human blood from around the lips while a person sleeps.[92]
Thebed bug,Cimex lectularius, is an externalparasite of humans. It lives in bedding and is mainly active at night, feeding on human blood, generally without being noticed.[93][94] Bed bugsmate bytraumatic insemination; the male pierces the female's abdomen and injects his sperm into a secondary genital structure, thespermalege. The sperm travel in the female's blood (haemolymph) tosperm storage structures (seminal conceptacles); they are released from there to fertilise her eggs inside herovaries.[94][95]
Some larger hemipterans such ascicadas are used as food in Asian countries such asChina,[96] 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.[97] At least nine species of Hemiptera are eaten worldwide.[98]
Cicadas have featured in literature since the time ofHomer'sIliad, and as motifs in decorative art from the ChineseShang dynasty (1766–1122 B.C.). They are described byAristotle in hisHistory of Animals and byPliny the Elder in hisNatural History; their mechanism of sound production is mentioned byHesiod in his poemWorks and Days "when the Skolymus flowers, and the tunefulTettix sitting on his tree in the weary summer season pours forth from under his wings his shrill song".[99]
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.[100]
Large-scale cultivation of the oil palmElaeis guineensis in theAmazon basin damages freshwater habitats and reduces the diversity of aquatic and semi-aquatic Heteroptera.[101] 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.[102]
^TheMuseum 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 andpillbugs),[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.[72]
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^Gillott, Cedric (2005).Entomology (3 ed.). Springer. p. 213.
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^Cobben, René Hubert (1978).Evolutionary trends in Heteroptera, part 2: mouthpart-structures and feeding strategies. Verslagen van landbouwkundige onderzoekingen, number 707. Wageningen, Netherlands: H. Veenman for Centrum voor Landbouwpublikaties en Landbouwdocumentatie. p. 7.OCLC852739989.
^Chinery, Michael (1993). "Order Hemiptera – The True Bugs".A Field Guide to the Insects of Britain and Northern Europe (3rd ed.). Collins. p. 115.ISBN978-0-00-219918-6.
^abGorb, 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.PMID18089035.
^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.ISBN978-1-4612-9126-8.
^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.S2CID55083254.
^Underwood, D. L. A."Hemipteroids"(PDF). California State University. Archived fromthe original(PDF) on 4 March 2016. Retrieved22 August 2015.
^Hall, Donald W.; Butler, Jerry F.; Cromroy, Harvey L. (1 September 2014)."Hackberry petiole gall psyllid".Featured Creatures. University of Florida/IFAS. Retrieved12 August 2015.
^Poinar, G. (2005). "Triatoma dominicana sp. n. (Hemiptera: Reduviidae: Triatominae), andTrypanosoma 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.PMID15815152.
^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.
^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. Retrieved16 July 2015.
^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. Retrieved16 June 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.PMID14570313.S2CID13150932.
^Carayon, J. 1959 Insémination par "spermalège" et cordon conducteur de spermatozoids chez Stricticimex brevispinosus Usinger (Heteroptera, Cimicidae). Rev. Zool. Bot. Afr. 60, 81–104.
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