Hemiptera
Hemiptera | |
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Species from each of the four suborders of Hemiptera, clockwise from top-left: Aphids (Sternorrhyncha ).
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
(unranked): | Paraneoptera |
Superorder: | Condylognatha |
Order: | Hemiptera Linnaeus, 1758 |
Suborders[2] | |
Hemiptera (
Entomologists reserve the term bug for Hemiptera or Heteroptera,
Many insects with "bug" in their
Most hemipterans feed on plants, using their sucking and piercing mouthparts to extract
Humans have interacted with the Hemiptera for millennia. Some species, including many aphids, are significant
Diversity
Hemiptera is the largest order of
Taxonomy and phylogeny
Hemiptera belong to the
The
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
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]
Paraneoptera |
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Suborder | No. of Species | First appearance | Examples | Characteristics |
---|---|---|---|---|
Auchenorrhyncha | over 42,000[20] | Lower Permian |
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 | larger bugs; some are predatory , others are plant-sucking
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Sternorrhyncha | 12,500 | Upper Permian |
scale insects |
plant-sucking bugs, some major horticultural pests; most are small and sedentary or fully sessile;[22]
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Biology
Mouthparts
The defining feature of hemipterans is their "beak" in which the modified
Both herbivorous and predatory hemipterans inject
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
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
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,
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
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
Some species attack pest insects and are used in biological control. One of these is the
Haematophagic ectoparasites
A few hemipterans are
As symbionts
Some species of
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.[62]
As prey: defences against predators and parasites
Hemiptera form prey to predators including vertebrates, such as birds, and other invertebrates such as
Some hemipterans such as
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.[72] In the aquatic Belostomatidae, females lay their eggs on the back of the male which guards the eggs.[73] 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
For pest control
Members of the families
Insect products
Other hemipterans have positive uses for humans, such as in the production of the dyestuff
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
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
Notes
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