Awasp is anyinsect of the narrow-waisted suborderApocrita of the orderHymenoptera which is neither abee nor anant; this excludes the broad-waistedsawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. The wasps do not constitute aclade, a complete natural group with a single ancestor, as bees and ants are deeply nested within the wasps, having evolved from wasp ancestors. Wasps that are members of the cladeAculeata cansting their prey.
The most commonly known wasps, such asyellowjackets andhornets, are in the familyVespidae and areeusocial, living together in a nest with an egg-laying queen and non-reproducing workers. Eusociality is favoured by the unusualhaplodiploid system ofsex determination in Hymenoptera, as it makes sisters exceptionally closely related to each other. However, the majority of wasp species are solitary, with each adult female living and breeding independently. Females typically have anovipositor for laying eggs in or near a food source for the larvae, though in theAculeata the ovipositor is often modified instead into asting used for defense or prey capture.
Wasps first appeared in the fossil record in theJurassic, and diversified into many surviving superfamilies by theCretaceous. They are a successful and diverse group of insects with tens of thousands of described species; wasps have spread to all parts of the world except for the polar regions. The largest social wasp is theAsian giant hornet, at up to 5 centimetres (2.0 in) in length; among the largest solitary wasps is a group of species known astarantula hawks, along with the giant scoliid of Indonesia (Megascolia procer). The smallest wasps are solitaryparasitoid wasps in the familyMymaridae, including the world's smallest known insect, with a body length of only 0.139 mm (0.0055 in), and the smallest known flying insect, only 0.15 mm (0.0059 in) long.
The wasps are a cosmopolitanparaphyletic grouping of hundreds of thousands of species,[1][2] consisting of the narrow-waistedcladeApocrita without theants andbees.[3] The Hymenoptera also contain the somewhat wasplike but unwaistedSymphyta, the sawflies.
Hymenoptera in the form of Symphyta (Xyelidae) first appeared in the fossil record in theLower Triassic. Apocrita, wasps in the broad sense, appeared in theJurassic, and had diversified into many of the extant superfamilies by theCretaceous; they appear to have evolved from the Symphyta.[4]Fig wasps with modern anatomical features first appeared in theLower Cretaceous of the Crato Formation in Brazil, some 65 million years before the first fig trees.[5]
Wasps are a diverse group, estimated at well over a hundred thousanddescribed species around the world, and a great many more as yet undescribed.[9][a] For example, almost every one of some 1000 species of tropicalfig trees has its own specificfig wasp (Chalcidoidea) that has co-evolved with it and pollinates it.[10]
Many wasp species are parasitoids; the females deposit eggs on or in a hostarthropod on which the larvae then feed. Some larvae start off as parasitoids, but convert at a later stage to consuming the plant tissues that their host is feeding on. In other species, the eggs are laid directly into plant tissues and formgalls, which protect the developing larvae from predators, but not necessarily from other parasitic wasps. In some species, the larvae are predatory themselves; the wasp eggs are deposited in clusters of eggs laid by other insects, and these are then consumed by the developing wasp larvae.[10]
The largest social wasp is theAsian giant hornet, at up to 5 centimetres (2.0 in) in length.[11] The varioustarantula hawk wasps are of a similar size[12] and can overpower a spider many times its own weight, and move it to its burrow, with a sting that is excruciatingly painful to humans.[13] The solitary giantscoliid,Megascolia procer, with a wingspan of 11.5 cm,[14] has subspecies inSumatra andJava;[15] it is aparasitoid of the Atlas beetleChalcosoma atlas.[16] The female giant ichneumon waspMegarhyssa macrurus is 12.5 centimetres (5 in) long including its very long but slenderovipositor which is used for boring into wood and inserting eggs.[17] The smallest wasps are solitaryparasitoid wasps in the familyMymaridae, including the world's smallest known insect,Dicopomorpha echmepterygis (139 micrometres long) andKikiki huna with a body length of only 158 micrometres, the smallest known flying insect.[18]
There are estimated to be 100,000 species ofichneumonoid wasps in the familiesBraconidae andIchneumonidae. These are almost exclusively parasitoids, mostly using other insects as hosts. Another family, thePompilidae, is a specialist parasitoid of spiders.[10] Some wasps are even parasitoids of parasitoids; the eggs ofEuceros are laid besidelepidopteran larvae and the wasp larvae feed temporarily on theirhaemolymph, but if a parasitoid emerges from the host, thehyperparasites continue their life cycle inside the parasitoid.[19] Parasitoids maintain their extreme diversity through narrow specialism. In Peru, 18 wasp species were found living on 14 fly species in only two species ofGurania climbing squash.[20][21]
Megascolia procer, a giant solitary species fromJava in theScoliidae. This specimen's length is 77 mm (3.0 in) and its wingspan is 115 mm (4.5 in).[b][14]
Megarhyssa macrurus, a parasitoid. The body of a female is 50 mm (2.0 in) long, with a c. 100 mm (3.9 in) ovipositor
Of the dozens of extant wasp families, only the familyVespidae contains social species, primarily in the subfamiliesVespinae andPolistinae. With their powerful stings and conspicuouswarning coloration, often in black and yellow, social wasps are frequent models forBatesian mimicry by non-stinging insects, and are themselves involved in mutually beneficialMüllerian mimicry of other distasteful insects including bees and other wasps. All species of social wasps construct their nests using some form of plant fiber (mostly wood pulp) as the primary material, though this can be supplemented with mud, plant secretions (e.g.,resin), and secretions from the wasps themselves; multiple fibrous brood cells are constructed, arranged in a honeycombed pattern, and often surrounded by a larger protective envelope. Wood fibres are gathered from weathered wood, softened by chewing and mixing withsaliva. The placement of nests varies from group to group; yellow jackets such asDolichovespula media andD. sylvestris prefer to nest in trees and shrubs;Protopolybia exiguaattaches its nests on the underside of leaves and branches;Polistes erythrocephalus chooses sites close to a water source.[22]
Other wasps, likeAgelaia multipicta andVespula germanica, like to nest in cavities that include holes in the ground, spaces under homes, wall cavities or in lofts. While most species of wasps have nests with multiple combs, some species, such asApoica flavissima, only have one comb.[23] The length of the reproductive cycle depends onlatitude;Polistes erythrocephalus, for example, has a much longer (up to 3 months longer) cycle in temperate regions.[24]
Solitary wasps
Potter wasp building mud nest, France. The latest ring of mud is still wet.
The vast majority of wasp species are solitary insects.[10][25] Having mated, the adult female forages alone and if it builds a nest, does so for the benefit of its own offspring. Some solitary wasps nest in small groups alongside others of their species, but each is involved in caring for its own offspring (except for such actions as stealing other wasps' prey or laying in other wasp's nests). There are some species of solitary wasp that build communal nests, each insect having its own cell and providing food for its own offspring, but these wasps do not adopt the division of labour and the complex behavioural patterns adopted byeusocial species.[25]
Adult solitary wasps spend most of their time in preparing their nests and foraging for food for their young, mostly insects or spiders. Their nesting habits are more diverse than those of social wasps. Many species dig burrows in the ground.[25]Mud daubers andpollen wasps construct mud cells in sheltered places.[26]Potter wasps similarly build vase-like nests from mud, often with multiple cells, attached to the twigs of trees or against walls.[27]
Predatory wasp species normally subdue their prey by stinging it, and then either lay their eggs on it, leaving it in place, or carry it back to their nest where an egg may be laid on the prey item and the nest sealed, or several smaller prey items may be deposited to feed a single developing larva. Apart from providing food for their offspring, no further maternal care is given. Members of the familyChrysididae, the cuckoo wasps, arekleptoparasites and lay their eggs in the nests of unrelated host species.[25]
Like all insects, wasps have a hardexoskeleton which protects their three main body parts, thehead, the mesosoma (including the thorax and the first segment of the abdomen) and the metasoma. There is a narrow waist, thepetiole, joining the first and second segments of the abdomen. The two pairs of membranous wings are held together by small hooks and the forewings are larger than the hind ones; in some species, the females have no wings. In females there is usually a rigid ovipositor which may be modified for injecting venom, piercing or sawing.[28] It either extends freely or can be retracted, and may be developed into a stinger for both defence and for paralysing prey.[29]
In addition to their largecompound eyes, wasps have several simple eyes known asocelli, which are typically arranged in a triangle just forward of thevertex of the head. Wasps possessmandibles adapted for biting and cutting, like those of many other insects, such asgrasshoppers, but their other mouthparts are formed into a suctorialproboscis, which enables them to drink nectar.[30]
The larvae of wasps resemblemaggots, and are adapted for life in a protected environment; this may be the body of a host organism or a cell in a nest, where the larva either eats the provisions left for it or, in social species, is fed by the adults. Such larvae have soft bodies with no limbs, and have a blind gut (presumably so that they do not foul their cell).[31]
Adult solitary wasps mainly feed on nectar, but the majority of their time is taken up by foraging for food for their carnivorous young, mostly insects or spiders. Apart from providing food for their larval offspring, no maternal care is given.[25] Some wasp species provide food for the young repeatedly during their development (progressive provisioning).[32] Others, such as potter wasps (Eumeninae)[33] and sand wasps (Ammophila,Sphecidae),[34] repeatedly build nests which they stock with a supply of immobilised prey such as one large caterpillar, laying a single egg in or on its body, and then sealing up the entrance (mass provisioning).[35]
Predatory and parasitoidal wasps subdue their prey by stinging it. They hunt a wide variety of prey, mainly other insects (including other Hymenoptera), both larvae and adults.[25]ThePompilidae specialize in catching spiders to provision their nests.[36]
Some social wasps are omnivorous, feeding on fallen fruit, nectar, and carrion such as dead insects. Adult male wasps sometimes visit flowers to obtainnectar. Some wasps, such asPolistes fuscatus, commonly return to locations where they previously found prey to forage.[37] In many social species, the larvae exude copious amounts ofsalivary secretions that are avidly consumed by the adults. These include bothsugars andamino acids, and may provide essential protein-building nutrients that are otherwise unavailable to the adults (who cannot digest proteins).[38]
Sex determination
In wasps, as in other Hymenoptera,sex is determined by ahaplodiploid system, which means that females are unusually closely related to their sisters, enablingkin selection to favour theevolution of eusocial behaviour. Females arediploid, meaning that they have 2nchromosomes and develop from fertilized eggs. Males, called drones, have ahaploid (n) number of chromosomes and develop from an unfertilized egg.[29] Wasps store sperm inside their body and control its release for each individual egg as it is laid; if a female wishes to produce a male egg, she simply lays the egg without fertilizing it. Therefore, under most conditions in most species, wasps have complete voluntary control over the sex of their offspring.[25] Experimental infection ofMuscidifurax uniraptor with thebacteriumWolbachia inducedthelytokous reproduction and an inability to produce fertile, viable male offspring.[39]
Inbreeding avoidance
Females of the solitary wasp parasitoidVenturia canescens can avoid mating with their brothers throughkin recognition.[40] In experimental comparisons, the probability that a female will mate with an unrelated male was about twice as high as the chance of her mating with brothers. Female wasps appear to recognize siblings on the basis of a chemical signature carried or emitted by males.[40] Sibling-mating avoidance reducesinbreeding depression that is largely due to the expression ofhomozygous deleterious recessive mutations.[41]
Ecology
As pollinators
While the vast majority of wasps play no role in pollination, a few species can effectively transport pollen and pollinate several plant species.[42] Since wasps generally do not have a fur-like covering of soft hairs and a special body part for pollen storage (pollen basket) as some bees do, pollen does not stick to them well.[43] However it has been shown that even without hairs, several wasp species are able to effectively transport pollen, therefore contributing for potential pollination of several plant species.[44]
Pollen wasps in the subfamilyMasarinae gather nectar and pollen in a crop inside their bodies, rather than on body hairs like bees, and pollinate flowers ofPenstemon and the water leaf family,Hydrophyllaceae.[45]
TheAgaonidae (fig wasps) are the only pollinators of nearly 1000 species offigs,[43] and thus are crucial to the survival of their host plants. Since the wasps are equally dependent on their fig trees for survival, thecoevolved relationship is fullymutualistic.[46]
Most solitary wasps are parasitoids.[47] As adults, those that do feed typically only take nectar from flowers.Parasitoid wasps are extremely diverse in habits, many laying their eggs in inert stages of their host (egg orpupa), sometimes paralysing their prey by injecting it with venom through their ovipositor. They then insert one or more eggs into the host or deposit them upon the outside of the host. The host remains alive until the parasitoid larvae pupate or emerge as adults.[48]
TheIchneumonidae are specialized parasitoids, often of Lepidoptera larvae deeply buried in plant tissues, which may bewoody. For this purpose, they have exceptionally long ovipositors; they detect their hosts by smell and vibration. Some of the largest species, includingRhyssa persuasoria andMegarhyssa macrurus, parasitisehorntails, large sawflies whose adult females also have impressively long ovipositors.[49] Some parasitic species have a mutualistic relationship with apolydnavirus that weakens the host'simmune system and replicates in theoviduct of the female wasp.[10]
One family ofchalcidoid wasps, theEucharitidae, has specialized asparasitoids of ants, most species hosted by one genus of ant. Eucharitids are among the few parasitoids that have been able to overcome ants' effective defences against parasitoids.[50][51][52]
Many species of wasp, including especially the cuckoo or jewel wasps (Chrysididae), are kleptoparasites, laying their eggs in the nests of other wasp species to exploit their parental care. Most such species attack hosts that provide provisions for their immature stages (such as paralyzed prey items), and they either consume the provisions intended for the host larva, or wait for the host to develop and then consume it before it reaches adulthood. An example of a truebrood parasite is the paper waspPolistes sulcifer, which lays its eggs in the nests of other paper wasps (specificallyPolistes dominula), and whose larvae are then fed directly by the host.[53][54] Sand waspsAmmophila often save time and energy by parasitising the nests of other females of their own species, either kleptoparasitically stealing prey, or as brood parasites, removing the other female's egg from the prey and laying their own in its place.[55] According toEmery's rule,social parasites, especially among insects, tend to parasitise species or genera to which they are closely related.[56][57] For example, the social waspDolichovespula adulterina parasitises other members of its genus such asD. norwegica andD. arenaria.[58][59]
As predators
Many wasp lineages, including those in the familiesVespidae,Crabronidae,Sphecidae, andPompilidae, attack and sting prey items that they use as food for their larvae; while Vespidae usually macerate their prey and feed the resulting bits directly to their brood, most predatory wasps paralyze their prey and lay eggs directly upon the bodies, and the wasp larvae consume them. Apart from collecting prey items to provision their young, many wasps are also opportunistic feeders, and will suck the body fluids of their prey. Although vespid mandibles are adapted for chewing and they appear to be feeding on the organism, they are often merely macerating it into submission. The impact of the predation of wasps on economicpests is difficult to establish.[60]
The roughly 140 species ofbeewolf (Philanthinae) hunt bees, including honeybees, to provision their nests; the adults feed on nectar and pollen.[61]
With their powerful stings and conspicuouswarning coloration, social wasps are the models for many species of mimic. Two common cases areBatesian mimicry, where the mimic is harmless and is essentially bluffing, andMüllerian mimicry, where the mimic is also distasteful, and the mimicry can be considered mutual. Batesian mimics of wasps include many species ofhoverfly and thewasp beetle. Many species of wasp are involved in Müllerian mimicry, as are many species ofbee.[62]
As prey
While wasp stings deter many potential predators,bee-eaters (in the bird family Meropidae) specialise in eating stinging insects, making aerialsallies from a perch to catch them, and removing the venom from the stinger by repeatedly brushing the prey firmly against a hard object, such as a twig.[63] Thehoney buzzard attacks the nests of social hymenopterans, eating wasp larvae; it is the only known predator of the dangerous[64] Asian giant hornet or "yak-killer" (Vespa mandarinia).[65] Likewise,roadrunners are the only real predators oftarantula hawk wasps.[66]
Bee-eaters such asMerops apiaster specialise in feeding on bees and wasps.
Relationship with humans
Paper wasp nest on a house
As pests
Social wasps are considered pests when they become excessively common, or nest close to buildings. People are most often stung in late summer and early autumn, when wasp colonies stop breeding new workers; the existing workers search for sugary foods and are more likely to come into contact with humans.[67][68][69] Wasp nests made in or near houses, such as in roof spaces, can present a danger as the wasps may sting if people come close to them.[70] Stings are usually painful rather than dangerous, but in rare cases, people may suffer life-threateninganaphylactic shock.[71]
Some species of parasitic wasp, especially in groups such asAphelinidae,Braconidae,Mymaridae, andTrichogrammatidae, are exploited commercially to providebiological control ofinsect pests.[2][72] One of the first species to be used wasEncarsia formosa, a parasitoid of a range of species ofwhitefly. It entered commercial use in the 1920s in Europe, was overtaken by chemicalpesticides in the 1940s, and again received interest from the 1970s.Encarsia is being tested ingreenhouses to control whitefly pests oftomato andcucumber, and to a lesser extent ofaubergine (eggplant), flowers such asmarigold, andstrawberry.[73] Several species of parasitic wasp are natural predators ofaphids and can help to control them.[74] For instance,Aphidius matricariae is used to control the peach-potato aphid.[75]
Tomato leaf covered with nymphs of whitefly parasitised byEncarsia formosa
In sport
Wasps RFC was an English professionalrugby union team originally based in London but later playing in Coventry; the name dates from 1867 at a time when names of insects were fashionable for clubs. The club's first kit was black with yellow stripes.[76] The club has an amateur side calledWasps FC.[77]
Among the other clubs bearing the name are a basketball club in Wantirna, Australia,[78] andAlloa Athletic F.C., a football club inScotland.[79]
Wasps have been modelled injewellery since at least the nineteenth century, whendiamond andemerald waspbrooches were made ingold andsilver settings.[80] A fashion forwasp waisted female silhouettes with sharply cinched waistlines emphasizing the wearer's hips and bust arose repeatedly in the nineteenth and twentieth centuries.[81][82]
In literature
The Ancient Greek playwrightAristophanes wrote thecomedy play Σφῆκες (Sphēkes),The Wasps, first put on in 422 BC. The "wasps" are the chorus of old jurors.[83]
It flew, he is convinced, within a yard of him, struck the ground, rose again, came down again perhaps thirty yards away, and rolled over with its body wriggling and its sting stabbing out and back in its last agony. He emptiedboth barrels into it before he ventured to go near. When he came to measure the thing, he found it was twenty-seven and a half inches across its open wings, and its sting was three inches long. ... The day after, a cyclist riding, feet up, down the hill between Sevenoaks and Tonbridge, very narrowly missed running over a second of these giants that was crawling across the roadway.[84]
Detail ofBotticelli'sVenus and Mars, 1485, with a wasp's nest on right, probably a symbol of the Vespucci family (Italianvespa, wasp) who commissioned the painting.[85]
Wasp (1957) is ascience fiction book by the English writerEric Frank Russell; it is generally considered Russell's best novel.[86] InStieg Larsson's bookThe Girl Who Played with Fire (2006) and itsfilm adaptation,Lisbeth Salander has adopted her kickboxing ringname, "The Wasp", as her hacker handle and has a wasp tattoo on her neck, indicating her high status among hackers, unlike her real world situation, and that like a small but painfully stinging wasp, she could be dangerous.[87]
Parasitoidal wasps played an indirect role in the nineteenth-centuryevolution debate. The Ichneumonidae contributed toCharles Darwin's doubts about the nature and existence of a well-meaning and all-powerful Creator. In an 1860 letter to the American naturalistAsa Gray, Darwin wrote:
I own that I cannot see as plainly as others do, and as I should wish to do, evidence of design and beneficence on all sides of us. There seems to me too much misery in the world. I cannot persuade myself that a beneficent and omnipotent God would have designedly created the Ichneumonidae with the express intention of their feeding within the living bodies of caterpillars, or that a cat should play with mice.[88]
With its powerful sting and familiar appearance, the wasp has given its name to many ships, aircraft and military vehicles.[89] Nine ships and one shore establishment of theRoyal Navy have been namedHMS Wasp, the first an 8-gunsloop launched in 1749.[90]Eleven ships of theUnited States Navy have similarly borne the nameUSS Wasp, the first a merchant schooner acquired by the Continental Navy in 1775.[91] The eighth of these,an aircraft carrier, gained two Second World War battle stars, promptingWinston Churchill to remark "Who said a Wasp couldn't sting twice?"[89] In the Second World War, a German self-propelled howitzer was namedWespe,[92] while the British developed theWasp flamethrower from theBren Gun Carrier.[93]In aerospace, theWestland Wasp was a military helicopter developed in England in 1958 and used by the Royal Navy and other navies.[94] TheAeroVironment Wasp III is aminiature UAV developed forUnited States Air Force special operations.[95]
^Methods to estimate species diversity include extrapolating the rate of species descriptions by subfamily (as in theBraconidae) until zero is reached; and extrapolating geographically from the species distribution of well-studied taxa to the group of interest (say, the Braconidae). Dolphin et al found a correlation between the predicted numbers of undescribed species by these two methods, doubling or tripling the number of species in the group.[9]
^Carlos A. Martin P.; Anthony C. Bellotti (1986)."Biologia y comportamiento dePolistes erythrocephalus" [Biology and behavior of Polistes erythrocephalus](PDF).Acta Agron (in Spanish).36 (1):63–76. Retrieved14 October 2014.
^Sôichi Yamane; Sidnei Mateus; Satoshi Hozumi; Kazuyuki Kudô; Ronaldo Zucchi (2009). "How does a colony ofApoica flavissima (Hymenoptera: Vespidae, Epiponini) maintain a constant temperature?".Entomological Science.12 (3):341–345.doi:10.1111/j.1479-8298.2009.00328.x.S2CID86577862.
^Hoell, H.V.; Doyen, J.T.; Purcell, A.H. (1998).Introduction to Insect Biology and Diversity, 2nd ed. Oxford University Press. pp. 570–579.ISBN978-0-19-510033-4.
^Elgar, Mark A.; Jebb, Matthew (1999). "Nest Provisioning In The Mud-Dauber WaspSceliphron laetum (F. Smith): Body Mass And Taxa Specific Prey Selection".Behaviour.136 (2):147–159.doi:10.1163/156853999501252.
^Lachaud, Jean-Paul; Pérez-Lachaud, Gabriela (2009). "Impact of natural parasitism by two eucharitid wasps on a potential biocontrol agent ant in southeastern Mexico".Biological Control.48 (1):92–99.Bibcode:2009BiolC..48...92L.doi:10.1016/j.biocontrol.2008.09.006.
^Williams, David F. (1994). "Biology and importance of two eucharitid parasites ofWasmannia andSolenopsis".Exotic ants: biology, impact, and control of introduced species. Boulder, CO: Westview Press. pp. 104–120.ISBN978-0-8133-8615-7.
^Dapporto, L.; Cervo, R.; Sledge, M. F.; Turillazzi, S. (2004). "Rank integration in dominance hierarchies of host colonies by the paper wasp social parasitePolistes sulcifer (Hymenoptera, Vespidae)".Journal of Insect Physiology.50 (2–3):217–223.Bibcode:2004JInsP..50..217D.doi:10.1016/j.jinsphys.2003.11.012.PMID15019524.
^O'Neill, Kevin M. (2001).Solitary Wasps: Behavior and Natural History. Cornell University Press. p. 129.
^"Diamond and emerald wasp brooch by Fontanna". A La Vieille Russie. Archived fromthe original on 10 June 2015. Retrieved10 June 2015.Crown rose diamond and emerald wasp brooch set in silver and gold. By Fontana French, ca. 1875. Width: 3 inches. $46,500
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![Megascolia procer, a giant solitary species from Java in the Scoliidae. This specimen's length is 77 mm (3.0 in) and its wingspan is 115 mm (4.5 in).[b][14]](/mt/?noimg=&dark=on&url=https%3a%2f%2fen.wikipedia.org%2fwiki%2fFile%3aMegascolia_procer_MHNT_dos.jpg)
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![Tarantula hawk wasp dragging the tarantula Megaphobema mesomelas to her burrow; it has the most painful sting of any wasp.[13]](/mt/?noimg=&dark=on&url=https%3a%2f%2fen.wikipedia.org%2fwiki%2fFile%3aWasp_with_Orange-kneed_tarantula.JPG)
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