Wasp#Social wasp caste structure

File:Wasps are Paraphyletic.svg, consisting of the clade Apocrita without ants and bees, which are not usually considered to be wasps. The Hymenoptera also contain the somewhat wasplike Symphyta, the sawflies. The familiar common wasps and yellowjackets belong to one family, the Vespidae.|left]]

The wasps are a cosmopolitan paraphyletic grouping of hundreds of thousands of species,{{cite web | last1=Broad | first1=Gavin | title=What's the point of wasps? | url=http://www.nhm.ac.uk/natureplus/community/research/life_sciences_news/wasps/blog/tags/species?fromGateway=true | publisher=Natural History Museum | access-date=18 June 2015 | date=25 June 2014}} consisting of the narrow-waisted clade Apocrita without the ants and bees.{{cite journal |last1=Johnson |first1=Brian R. |author2=Borowiec, Marek L. |author3=Chiu, Joanna C. |author4=Lee, Ernest K. |author5=Atallah, Joel |author6=Ward, Philip S. | date=2013 | title=Phylogenomics Resolves Evolutionary Relationships among Ants, Bees, and Wasps | journal=Current Biology | volume=23 | issue=20 | pages=2058–2062 | url=http://www.cell.com/current-biology/pdf/S0960-9822(13)01056-7.pdf |doi=10.1016/j.cub.2013.08.050 | pmid=24094856|s2cid=230835 |doi-access=free |bibcode=2013CBio...23.2058J }} The Hymenoptera also contain the somewhat wasplike but unwaisted Symphyta, the sawflies.

The term wasp is sometimes used more narrowly for members of the Vespidae, which includes several eusocial wasp lineages, such as yellowjackets (the genera Vespula and Dolichovespula), hornets (genus Vespa), and members of the subfamily Polistinae.

File:Neotype male of Electrostephanus petiolatus Brues in Baltic amber (AMNH B-JWJ-260).png fossil from the Middle Eocene, preserved in Baltic amber]]

Hymenoptera in the form of Symphyta (Xyelidae) first appeared in the fossil record in the Lower Triassic. Apocrita, wasps in the broad sense, appeared in the Jurassic, and had diversified into many of the extant superfamilies by the Cretaceous; they appear to have evolved from the Symphyta.{{cite book | last=Gillott |first=Cedric | title=Entomology | url=https://books.google.com/books?id=sFLaBwAAQBAJ&pg=PA302 | date=6 December 2012 | publisher=Springer | isbn=978-1-4615-6915-2 | pages=302–318}} Fig wasps with modern anatomical features first appeared in the Lower Cretaceous of the Crato Formation in Brazil, some 65 million years before the first fig trees.{{cite web |title=World's oldest fig wasp fossil proves that if it works, don't change it |url=http://www.leeds.ac.uk/news/article/834/worlds_oldest_fig_wasp_fossil_proves_that_if_it_works_dont_change_it |access-date=5 August 2015 |date=15 June 2010 |publisher=University of Leeds |archive-date=7 September 2015 |archive-url=https://web.archive.org/web/20150907095104/http://www.leeds.ac.uk/news/article/834/worlds_oldest_fig_wasp_fossil_proves_that_if_it_works_dont_change_it |url-status=dead }}

The Vespidae include the extinct genus Palaeovespa, seven species of which are known from the Eocene rocks of the Florissant fossil beds of Colorado and from fossilised Baltic amber in Europe.{{cite journal | last1=Poinar | first1=G. | author-link=George Poinar Jr. | year=2005 | title=Fossil Trigonalidae and Vespidae (Hymenoptera) in Baltic amber | journal=Proceedings of the Entomological Society of Washington | volume=107 | issue=1 | pages=55–63 | url=http://biostor.org/reference/55642}} Also found in Baltic amber are crown wasps of the genus Electrostephanus.{{Cite journal | last1=Engel | first1=M.S. | last2=Ortega-Blanco | first2=J. | year=2008 | title=The fossil crown wasp Electrostephanus petiolatus Brues in Baltic Amber (Hymenoptera, Stephanidae): designation of a neotype, revised classification, and a key to amber Stephanidae | journal= ZooKeys | issue=4 | pages = 55–64 |doi=10.3897/zookeys.4.49| doi-access=free | bibcode=2008ZooK....4...55E | hdl=2445/36428 | hdl-access=free }}{{Cite journal | last1=Yunakov | first1=N.N. | last2=Kirejtshuk | first2=A.G. | year=2011 | title=New genus and species of broad-nosed weevils from Baltic amber and notes on fossils of the subfamily Entiminae (Coleoptera, Curculionidae) | journal= ZooKeys | issue=160 | pages = 73–96 |doi= 10.3897/zookeys.160.2108 | pmid=22303121 |pmc=3253632| doi-access=free | bibcode=2011ZooK..160...73Y }}{{-}}

File:Indian yellow wasp.jpg sp., India]]

Wasps are a diverse group, estimated at well over a hundred thousand described species around the world, and a great many more as yet undescribed.{{cite journal | last1=Dolphin | first1=Konrad | last2=Quicke | first2=Donald L. J. | title=Estimating the global species richness of an incompletely described taxon: an example using parasitoid wasps (Hymenoptera: Braconidae) | journal=Biological Journal of the Linnean Society | date=July 2001 | volume=73 | issue=3 | pages=279–286 | doi=10.1111/j.1095-8312.2001.tb01363.x| doi-access=free }}{{efn|Methods to estimate species diversity include extrapolating the rate of species descriptions by subfamily (as in the Braconidae) 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.}} For example, almost every one of some 1000 species of tropical fig trees has its own specific fig wasp (Chalcidoidea) that has co-evolved with it and pollinates it.

Many wasp species are parasitoids; the females deposit eggs on or in a host arthropod 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 form galls, 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.

The largest social wasp is the Asian giant hornet, at up to {{convert | 5 | cm}} in length.{{cite book | url = https://books.google.com/books?id=qOyV7KnJRCYC&pg=PA147 | title = Steve Backshall's venom: poisonous animals in the natural world | first = Steve | last = Backshall | publisher = New Holland Publishers | year = 2007 | page=147 | isbn = 978-1-84537-734-2 }} The various tarantula hawk wasps are of a similar size{{cite book | last=Carwardine | first=Mark | title=Animal Records | url=https://books.google.com/books?id=T3FEKopUFkUC&pg=PA218 | year=2008 | publisher=Sterling Publishing | isbn=978-1-4027-5623-8 | page=218 | quote=Pepsis heros, which has a body length of up to 5.7cm (2 1/4 in) and a maximum wingspan of 11.4 cm (4 1/2 in).}} 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.{{cite web | url=http://www.desertusa.com/insects/tarantula-hawks.html | title=Tarantula Hawk | last=Williams |first=David B. | publisher=DesertUSA | access-date=13 June 2015}} The solitary giant scoliid, Megascolia procer, with a wingspan of 11.5 cm, has subspecies in Sumatra and Java;{{cite journal | last1=Betrem | first1=J. G. | last2=Bradley | first2=J. Chester | title= Annotations on the genera Triscolia, Megascolia and Scolia (Hymenoptera, Scoliidae) | journal=Zoologische Mededelingen | date=1964 | volume=39 | issue=43 | pages=433–444 | url=http://www.repository.naturalis.nl/document/149949}} it is a parasitoid of the Atlas beetle Chalcosoma atlas.{{cite book | last=Piek | first=Tom | title=Venoms of the Hymenoptera: Biochemical, Pharmacological and Behavioural Aspects | url=https://books.google.com/books?id=xBQlBQAAQBAJ&pg=PA161 | year=2013 | publisher=Elsevier | isbn=978-1-4832-6370-0 | page=173}} The female giant ichneumon wasp Megarhyssa macrurus is {{convert | 12.5 | cm | in | 0}} long including its very long but slender ovipositor which is used for boring into wood and inserting eggs.{{cite web | url=http://www.ext.colostate.edu/pubs/insect/05604.html | title=Pigeon Tremex Horntail and the Giant Ichneumon Wasp | last=Cranshaw |first=W. | date=5 August 2014 | publisher=Colorado State University Extension | access-date=15 June 2015 | archive-date=24 September 2015 | archive-url=https://web.archive.org/web/20150924005235/http://www.ext.colostate.edu/pubs/insect/05604.html | url-status=dead }} The smallest wasps are solitary parasitoid wasps in the family Mymaridae, including the world's smallest known insect, Dicopomorpha echmepterygis (139 micrometres long) and Kikiki huna with a body length of only 158 micrometres, the smallest known flying insect.{{Cite journal |last1=Huber |first1=John |last2=Noyes |first2=John |author-link2=John Noyes (entomologist)|year=2013 |title=A new genus and species of fairyfly, Tinkerbella nana (Hymenoptera, Mymaridae), with comments on its sister genus Kikiki, and discussion on small size limits in arthropods |journal=Journal of Hymenoptera Research |volume=32 |pages=17–44 |doi=10.3897/jhr.32.4663|doi-access=free }}

There are estimated to be 100,000 species of ichneumonoid wasps in the families Braconidae and Ichneumonidae. These are almost exclusively parasitoids, mostly using other insects as hosts. Another family, the Pompilidae, is a specialist parasitoid of spiders.{{cite book | last=Godfray |first=H.C.J. | title=Parasitoids: Behavioral and Evolutionary Ecology | url=https://books.google.com/books?id=B1ZxjLyGQiQC | year=1994 | publisher=Princeton University Press | isbn=978-0-691-00047-3 | pages=3–24}} Some wasps are even parasitoids of parasitoids; the eggs of Euceros are laid beside lepidopteran larvae and the wasp larvae feed temporarily on their haemolymph, but if a parasitoid emerges from the host, the hyperparasites continue their life cycle inside the parasitoid.{{cite web | url=http://www.landcareresearch.co.nz/science/plants-animals-fungi/animals/invertebrates/systematics/hymenoptera/ichneumonidae/factsheets/euceros | title=Ichneumonidae: Eucerotinae: Euceros Gravenhorst 1829 |last1=Ward |first1=D.F. |last2=Schnitzler |first2=F.R. | year=2013 | publisher=Landcare Research | access-date=15 June 2015}} Parasitoids maintain their extreme diversity through narrow specialism. In Peru, 18 wasp species were found living on 14 fly species in only two species of Gurania climbing squash.{{cite web | last1=Westlake | first1=Casey | title=More to biological diversity than meets the eye: Specialization by insect species is the key | url=http://now.uiowa.edu/2014/03/more-biological-diversity-meets-eye | website=Iowa Now | publisher=University of Iowa | access-date=18 June 2015 | date=13 March 2014}}{{cite journal | last1=Condon | first1=M. A. | last2=Scheffer | first2=S. J. | last3=Lewis | first3=M. L. | last4=Wharton | first4=R. | last5=Adams | first5=D. C. | last6=Forbes | first6=A. A. | title=Lethal interactions between parasites and prey increase niche diversity in a tropical community | journal=Science | year=2014 | volume= 343| issue=6176 | pages=1240–1244 | doi=10.1126/science.1245007 | pmid=24626926| bibcode=2014Sci...343.1240C | s2cid=13911928 | url=https://dr.lib.iastate.edu/bitstreams/7a842bad-e3b1-4674-b1e9-49d78105778f/download }}

File:Megascolia procer MHNT dos.jpg|Megascolia procer, a giant solitary species from Java in the Scoliidae. This specimen's length is {{cvt|77|mm|in}} and its wingspan is {{cvt|115|mm|in}}.{{efn | Specimen measured from photograph.}}{{cite journal | last1=Sarrazin | first1=Michael | last2=Vigneron | first2=Jean Pol | last3=Welch | first3=Victoria | last4=Rassart | first4=Marie | title=Nanomorphology of the blue iridescent wings of a giant tropical wasp Megascolia procer javanensis (Hymenoptera) | journal=Phys. Rev. | date=5 November 2008 | volume=E 78 | issue=5 | pages=051902 | doi=10.1103/PhysRevE.78.051902 | pmid=19113150 | arxiv=0710.2692 | bibcode=2008PhRvE..78e1902S | s2cid=30936410 }} Measurement scale on Figure 1.

File:Ichneumon_wasp_(Megarhyssa_macrurus_lunato)_(7686081848).jpg|Megarhyssa macrurus, a parasitoid. The body of a female is {{cvt|50|mm|in}} long, with a c. {{cvt|100|mm|in}} ovipositor

File:Wasp with Orange-kneed tarantula.JPG|Tarantula hawk wasp dragging the tarantula Megaphobema mesomelas to her burrow; it has the most painful sting of any wasp.

File:Active Wasp Nest.jpg

Of the dozens of extant wasp families, only the family Vespidae contains social species, primarily in the subfamilies Vespinae and Polistinae. With their powerful stings and conspicuous warning coloration, often in black and yellow, social wasps are frequent models for Batesian mimicry by non-stinging insects, and are themselves involved in mutually beneficial Müllerian 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 with saliva. The placement of nests varies from group to group; yellow jackets such as Dolichovespula media and D. sylvestris prefer to nest in trees and shrubs; Protopolybia exigua attaches its nests on the underside of leaves and branches; Polistes erythrocephalus chooses sites close to a water source.{{cite journal | author=Carlos A. Martin P. | author2=Anthony C. Bellotti | url=http://www.bdigital.unal.edu.co/18779/1/14701-44150-1-PB.pdf | title=Biologia y comportamiento de Polistes erythrocephalus | language=es |trans-title=Biology and behavior of Polistes erythrocephalus | journal=Acta Agron | volume=36 | issue=1 | pages=63–76 | year=1986 | access-date=2014-10-14}}

Other wasps, like Agelaia multipicta and Vespula 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 as Apoica flavissima, only have one comb.{{cite journal | author=Sôichi Yamane | author2=Sidnei Mateus | author3=Satoshi Hozumi | author4=Kazuyuki Kudô | author5=Ronaldo Zucchi | title=How does a colony of Apoica flavissima (Hymenoptera: Vespidae, Epiponini) maintain a constant temperature? | journal=Entomological Science | volume=12 | issue=3 | pages=341–345 | year=2009 |doi=10.1111/j.1479-8298.2009.00328.x | s2cid=86577862 }} The length of the reproductive cycle depends on latitude; Polistes erythrocephalus, for example, has a much longer (up to 3 months longer) cycle in temperate regions.{{cite book | last=Ross | first=Kenneth G. | title=The Social Biology of Wasps | year=1991 | publisher=Cornell University Press | isbn=978-0-8014-9906-7 | page=104 }}

File:Potter Wasp building mud nest near completion.JPG building mud nest, France. The latest ring of mud is still wet.]]

The vast majority of wasp species are solitary insects. Having mated, the adult female forages alone and if it builds a nest, doing 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 by eusocial species.{{cite book| last=O'Neill |first=Kevin M. | title=Solitary Wasps: Behavior and Natural History | url=https://books.google.com/books?id=khOmHyUT3AkC | year=2001 | publisher=Cornell University Press | isbn=978-0-8014-3721-2 | pages=1–4, 69}}

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. Mud daubers and pollen wasps construct mud cells in sheltered places.{{cite web | last1=Houston | first1=Terry | title=Slender mud-dauber wasps: genus Sceliphron | url=http://museum.wa.gov.au/research/collections/terrestrial-zoology/entomology-insect-collection/entomology-factsheets/sceliphron | publisher=Western Australian Museum | access-date=12 June 2015 | date=October 2013}} Potter wasps similarly build vase-like nests from mud, often with multiple cells, attached to the twigs of trees or against walls.{{cite web | last1=Grissell | first1=E. E. | title=Potter wasps of Florida | url=http://entomology.ifas.ufl.edu/creatures/misc/wasps/potter_wasps.htm | publisher=University of Florida | access-date=12 June 2015 | date=April 2007}}

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 family Chrysididae, the cuckoo wasps, are kleptoparasites and lay their eggs in the nests of unrelated host species.

File:Vespa crabro09(js), Lodz(Poland).jpg]]

Like all insects, wasps have a hard exoskeleton which protects their three main body parts, the head, the mesosoma (including the thorax and the first segment of the abdomen) and the metasoma. There is a narrow waist, the petiole, 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.{{cite web | url=http://www.ento.csiro.au/education/insects/hymenoptera.html | title=Hymenoptera: ants, bees and wasps |work=Insects and their allies | publisher=CSIRO | access-date=16 June 2015}} It either extends freely or can be retracted, and may be developed into a stinger for both defence and for paralysing prey.

In addition to their large compound eyes, wasps have several simple eyes known as ocelli, which are typically arranged in a triangle just forward of the vertex of the head. Wasps possess mandibles adapted for biting and cutting, like those of many other insects, such as grasshoppers, but their other mouthparts are formed into a suctorial proboscis, which enables them to drink nectar.{{cite journal | last1=Krenn | first1=H. W. | last2=Mauss | first2=V. | last3=Plant | first3=J. | title=Evolution of the suctorial proboscis in pollen wasps (Masarinae, Vespidae) | journal=Arthropod Structure & Development | year=2002 | volume=31 | issue=2 | pages=103–120 | pmid=18088974 | doi=10.1016/s1467-8039(02)00025-7| bibcode=2002ArtSD..31..103K }}

The larvae of wasps resemble maggots, 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).{{cite book |last1=Hoell |first1=H.V. |author2=Doyen, J.T. |author3=Purcell, A.H. | year=1998 | title=Introduction to Insect Biology and Diversity, 2nd ed. | publisher= Oxford University Press | pages= 570–579 | isbn= 978-0-19-510033-4}}

File:Wasp August 2007-23.jpg Bembix oculata (Crabronidae) feeding on a fly after paralysing it with its sting]]

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. Some wasp species provide food for the young repeatedly during their development (progressive provisioning).{{cite journal | last1=Field | first1=Jeremy | title=The evolution of progressive provisioning | journal=Behavioral Ecology | year=2005 | volume=16 | issue=4 | pages=770–778 |doi=10.1093/beheco/ari054 | url=https://www.sussex.ac.uk/webteam/gateway/file.php?name=field2005pp.pdf&site=196| doi-access=free | hdl=10.1093/beheco/ari054 | hdl-access=free }} Others, such as potter wasps (Eumeninae){{cite web | last1=Grissell | first1=E. E.| title=Potter Wasps of Florida | url=http://entomology.ifas.ufl.edu/creatures/misc/wasps/potter_wasps.htm | publisher=University of Florida / IFAS | access-date=15 June 2015}} and sand wasps (Ammophila, Sphecidae),{{cite journal | last=Field |first=Jeremy | year=1989 | title=Intraspecific parasitism and nesting success in the solitary wasp Ammophila sabulosa | journal=Behaviour | volume=110 | issue=1–4 | pages=23–45 |jstor=4534782 |doi=10.1163/156853989X00367}} 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).{{cite journal | last1=Elgar | first1=Mark A. | last2=Jebb | first2=Matthew | title=Nest Provisioning In The Mud-Dauber Wasp Sceliphron laetum (F. Smith): Body Mass And Taxa Specific Prey Selection | journal=Behaviour | date=1999 | volume=136 | issue=2 | pages=147–159 |doi=10.1163/156853999501252 }}

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.

The Pompilidae specialize in catching spiders to provision their nests.{{cite web| title=Spider Wasps | url=https://australian.museum/learn/animals/insects/spider-wasps/ | publisher=Australian Museum | access-date=15 June 2015}}

File:IndianSpiderWasp.JPG (Pompilidae) dragging a jumping spider (Salticidae) to provision a nest]]

Some social wasps are omnivorous, feeding on fallen fruit, nectar, and carrion such as dead insects. Adult male wasps sometimes visit flowers to obtain nectar. Some wasps, such as Polistes fuscatus, commonly return to locations where they previously found prey to forage.{{cite journal | last=Richter | first=M. Raveret | title=Social Wasp (Hymenoptera: Vespidae) Foraging Behavior | journal=Annual Review of Entomology | year=2000 | volume=45 | pages=121–150 |doi=10.1146/annurev.ento.45.1.121 | pmid=10761573 }} In many social species, the larvae exude copious amounts of salivary secretions that are avidly consumed by the adults. These include both sugars and amino acids, and may provide essential protein-building nutrients that are otherwise unavailable to the adults (who cannot digest proteins).{{cite book| last=Wilson |first=Edward O. |author-link=Edward O. Wilson | title=Sociobiology: The New Synthesis | url=https://books.google.com/books?id=v7lV9tz8fXAC&pg=PA344 | year=2000 | publisher=Harvard University Press | isbn=978-0-674-00089-6 | page=344}}

In wasps, as in other Hymenoptera, sex is determined by a haplodiploid system, which means that females are unusually closely related to their sisters, enabling kin selection to favour the evolution of eusocial behaviour. Females are diploid, meaning that they have 2n chromosomes and develop from fertilized eggs. Males, called drones, have a haploid (n) number of chromosomes and develop from an unfertilized egg.{{cite book| last1=Grimaldi |first1=David | last2= Engel |first2=Michael S. | title=Evolution of the Insects | url=https://books.google.com/books?id=Ql6Jl6wKb88C | year=2005 | publisher=Cambridge University Press | isbn=978-0-521-82149-0 | page=408}} 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. Experimental infection of Muscidifurax uniraptor with the bacterium Wolbachia induced thelytokous reproduction and an inability to produce fertile, viable male offspring.{{Cite journal |last1=Gottlieb |first1=Yuval |last2=Zchori-Fein |first2=Einat |title=Irreversible thelytokous reproduction in Muscidifurax uniraptor |journal=Entomologia Experimentalis et Applicata |volume=100 |issue=3 |pages=271–278 |year=2001 |url=http://www.nhm.ac.uk/resources/research-curation/projects/chalcidoids/pdf_Y/GottliZc2001.pdf |doi=10.1046/j.1570-7458.2001.00874.x|bibcode=2001EEApp.100..271G |s2cid=54687768 }}

Females of the solitary wasp parasitoid Venturia canescens can avoid mating with their brothers through kin recognition.{{cite journal |last1=Metzger |first1=M. |others=Bernstein, C.; Hoffmeister, T.S.; Desouhant, E. |title=Does kin recognition and sib-mating avoidance limit the risk of genetic incompatibility in a parasitic wasp? |journal=PLOS ONE |volume=5 |issue=10 |pages=e13505 |year=2010 |pmid=20976063 |pmc=2957437 |doi=10.1371/journal.pone.0013505 |bibcode=2010PLoSO...513505M |doi-access=free }} 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. Sibling-mating avoidance reduces inbreeding depression that is largely due to the expression of homozygous deleterious recessive mutations.{{cite journal |last1=Charlesworth |first1=D. |last2=Willis |first2=J.H. |title=The genetics of inbreeding depression |journal=Nat. Rev. Genet. |volume=10 |issue=11 |pages=783–96 |year=2009 |pmid=19834483 |doi=10.1038/nrg2664 |s2cid=771357 }}

While the vast majority of wasps play no role in pollination, a few species can effectively transport pollen and pollinate several plant species.{{cite journal | last1=Sühs | first1=R.B. | last2=Somavilla | first2=A. | last3=Köhler | first3=A | last4=Putzke | first4=J. | year=2009 | title=Vespídeos (Hymenoptera, Vespidae) vetores de pólen de Schinus terebinthifolius Raddi (Anacardiaceae), Santa Cruz do Sul, RS, Brasil | language=pt |trans-title=Pollen vector wasps (Hymenoptera, Vespidae) of Schinus terebinthifolius Raddi (Anacardiaceae), Santa Cruz do Sul, RS, Brazil | journal=Brazilian Journal of Biosciences | volume=7 | issue=2 | pages=138–143 | url=http://www.ufrgs.br/seerbio/ojs/index.php/rbb/article/view/1123}} 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.{{cite web|url=http://www.fs.fed.us/wildflowers/pollinators/animals/wasps.shtml |title=Wasp Pollination |publisher=USDA Forest Service |access-date=5 August 2015}} 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.{{Cite journal |last1=Sühs |last2=Somavilla |last3=Putzke |last4=Köhler |year=2009 |title=Pollen vector wasps (Hymenoptera, Vespidae) of Schinus terebinthifolius Raddi (Anacardiaceae), Santa Cruz do Sul, RS, Brazil |journal=Brazilian Journal of Biosciences |volume=7 |issue=2 |pages=138–143 |via=ABEC}}

Pollen wasps in the subfamily Masarinae gather nectar and pollen in a crop inside their bodies, rather than on body hairs like bees, and pollinate flowers of Penstemon and the water leaf family, Hydrophyllaceae.{{cite web | url=http://www.fs.fed.us/wildflowers/pollinators/pollinator-of-the-month/masarines.shtml | title=Pollen Wasps | last=Tepidino |first=Vince | publisher=USDA Forest Service | access-date=5 August 2015}}

The Agaonidae (fig wasps) are the only pollinators of nearly 1000 species of figs, and thus are crucial to the survival of their host plants. Since the wasps are equally dependent on their fig trees for survival, the coevolved relationship is fully mutualistic.{{Cite journal | last1=Machado | first1=Carlos A. | last2=Robbins | first2=Nancy | last3=Gilbert | first3=M. Thomas | last4=Herre | first4=Edward Allen | title=Critical Review of Host Specificity and Its Coevolutionary Implications in the Fig-fig-wasp Mutualism | journal=PNAS | volume=102 | date=April 2005 | issue=Suppl 1 | pages=6558–65 | pmid=15851680 | pmc=1131861 | doi=10.1073/pnas.0501840102| bibcode=2005PNAS..102.6558M | doi-access=free }}

{{main|Parasitoid wasp}}

Most solitary wasps are parasitoids.{{cite web |last1=Sekar |first1=Sandhya |title=Parasitoid wasps may be the most diverse animal group |url=https://www.bbc.co.uk/earth/story/20150522-the-wasps-that-rule-the-world |publisher=BBC |access-date=14 February 2018 |date=22 May 2015}} 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 or pupa), 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.{{cite book | title=Parasitic Wasps | last=Quicke |first=D.L.J. | date=1997 | isbn=978-0-412-58350-6 | pages=Chapter 8 and passim| publisher=Springer }}

The Ichneumonidae are specialized parasitoids, often of Lepidoptera larvae deeply buried in plant tissues, which may be woody. For this purpose, they have exceptionally long ovipositors; they detect their hosts by smell and vibration. Some of the largest species, including Rhyssa persuasoria and Megarhyssa macrurus, parasitise horntails, large sawflies whose adult females also have impressively long ovipositors.{{cite web | last1=Sezen | first1=Uzay | title=Giant Ichneumon Wasp (Megarhyssa macrurus) Ovipositing | url=http://naturedocumentaries.org/3843/giant-ichneumon-megarhyssa-macrurus-ovipositing/ | publisher=Nature Documentaries.org | access-date=15 June 2015 | date=24 March 2015}} Some parasitic species have a mutualistic relationship with a polydnavirus that weakens the host's immune system and replicates in the oviduct of the female wasp.

One family of chalcidoid wasps, the Eucharitidae, has specialized as parasitoids 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.{{cite journal | last1=Lachaud | first1=Jean-Paul | last2=Pérez-Lachaud | first2=Gabriela | title=Impact of natural parasitism by two eucharitid wasps on a potential biocontrol agent ant in southeastern Mexico | journal=Biological Control | date=2009 | volume=48 | issue=1 | pages=92–99 | doi=10.1016/j.biocontrol.2008.09.006| bibcode=2009BiolC..48...92L }}{{cite book | last1=Williams | first1=David F. | title=Exotic ants : biology, impact, and control of introduced species | date=1994 | publisher=Westview Press|location=Boulder, CO | isbn=978-0-8133-8615-7| pages=104–120|chapter=Biology and importance of two eucharitid parasites of Wasmannia and Solenopsis}}{{cite journal | last1=Brues | first1=C. T. | title=A New Chalcid-Fly Parasitic on the Australian Bull-Dog Ant | journal=Annals of the Entomological Society of America | year=1919 | volume=12 | issue=1 | pages=13–21 |doi=10.1093/aesa/12.1.13| url=https://www.biodiversitylibrary.org/partpdf/21997 }}

{{further|Kleptoparasitism|Cuckoo wasp}}

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 true brood parasite is the paper wasp Polistes sulcifer, which lays its eggs in the nests of other paper wasps (specifically Polistes dominula), and whose larvae are then fed directly by the host.{{cite journal | last1=Ortolani | first1=I. | last2=Cervo | first2=R. | year=2009 | title=Coevolution of daily activity timing in a host-parasite system | journal=Biological Journal of the Linnean Society | volume=96 | issue=2 | pages=399–405 | doi=10.1111/j.1095-8312.2008.01139.x| doi-access=free }}{{cite journal | last1=Dapporto | first1=L. | last2=Cervo | first2=R. | last3=Sledge | first3=M. F. | last4=Turillazzi | first4=S. | year=2004 | title=Rank integration in dominance hierarchies of host colonies by the paper wasp social parasite Polistes sulcifer (Hymenoptera, Vespidae) | journal=Journal of Insect Physiology | volume=50 | issue=2–3 | pages=217–223 | doi=10.1016/j.jinsphys.2003.11.012 | pmid=15019524 | bibcode=2004JInsP..50..217D }} Sand wasps Ammophila 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.{{cite book | last=O'Neill |first=Kevin M. | year=2001 | title=Solitary Wasps: Behavior and Natural History | publisher=Cornell University Press | page=129}} According to Emery's rule, social parasites, especially among insects, tend to parasitise species or genera to which they are closely related.{{cite web | url=http://www.nature.com/scitable/knowledge/library/social-parasitism-in-ants-13256421 | title=Social Parasitism in Ants | last=Deslippe |first=Richard | publisher=Nature Education Knowledge | year=2010}}{{cite journal | last=Emery |first=C. | title=Über den Ursprung der dulotischen, parasitischen und myrmekophilen Ameisen |language=German | journal=Biologisches Centralblatt | volume=29 | pages=352–362 | date=1909}} For example, the social wasp Dolichovespula adulterina parasitises other members of its genus such as D. norwegica and D. arenaria.{{cite journal | last1=Carpenter | first1=James M. | last2=Perera | first2=Estelle P. | title=Phylogenetic Relationships among Yellowjackets and the Evolution of Social Parasitism (Hymenoptera: Vespidae, Vespinae) | journal=American Museum Novitates | date=16 March 2006 | issue=3507 | pages=1–19 | doi=10.1206/0003-0082(2006)3507[1:PRAYAT]2.0.CO;2| hdl=2246/5782 | s2cid=53048610 | url=https://www.biodiversitylibrary.org/itempdf/280855 }}{{Cite journal | title=Parasitism of Dolichovespula norwegica by D. adulterina (Hymenoptera: Vespidae) | last=Dvorak | first=L. | date=2007 | volume=13 | issue=1 | pages=65–67 | journal=Silva Gabreta | url=http://www.npsumava.cz/storage/vyzkum/SGpdf/SG13_1_Dvorak.pdf | access-date=23 June 2015 | archive-date=4 March 2016 | archive-url=https://web.archive.org/web/20160304214434/http://www.npsumava.cz/storage/vyzkum/SGpdf/SG13_1_Dvorak.pdf | url-status=dead }}

Many wasp lineages, including those in the families Vespidae, Crabronidae, Sphecidae, and Pompilidae, 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 economic pests is difficult to establish.{{cite book| last1=Fisher |first1=T.W.| author2=Bellows, Thomas S.| author3=Caltagirone, L.E. | author4=Dahlsten, D.L. | author5=Huffaker, Carl B. | author6 = Gordh, G.| title=Handbook of Biological Control: Principles and Applications of Biological Control | url=https://books.google.com/books?id=u2X-rfgU0ewC&pg=PA455 | year=1999 | publisher=Academic Press | isbn=978-0-08-053301-8 | page=455}}

The roughly 140 species of beewolf (Philanthinae) hunt bees, including honeybees, to provision their nests; the adults feed on nectar and pollen.{{cite web | title=Biology of the European beewolf (Philanthus triangulum, Hymenoptera, Crabronidae) | url=http://www.biologie.uni-regensburg.de/Zoologie/Strohm/beewolfbiology/index.html | website=Evolutionary Ecology | publisher=University of Regensburg | access-date=20 June 2015 | date=5 June 2007}}

{{further|Mimicry|Aposematism}}

With their powerful stings and conspicuous warning coloration, social wasps are the models for many species of mimic. Two common cases are Batesian mimicry, where the mimic is harmless and is essentially bluffing, and Müllerian mimicry, where the mimic is also distasteful, and the mimicry can be considered mutual. Batesian mimics of wasps include many species of hoverfly and the wasp beetle. Many species of wasp are involved in Müllerian mimicry, as are many species of bee.{{cite book | last=Edmunds |first=Malcolm | year=1974 | title=Defence in Animals: A Survey of Anti-Predator Defences | url=https://archive.org/details/defenceinanimals0000edmu | url-access=registration | publisher=Longman | isbn=978-0-582-44132-3 | pages=[https://archive.org/details/defenceinanimals0000edmu/page/74 74], 82–83}}

While wasp stings deter many potential predators, bee-eaters (in the bird family Meropidae) specialise in eating stinging insects, making aerial sallies 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.{{cite book | editor=Forshaw, Joseph |last1=Forshaw |first1=J. |last2=Kemp |first2=A. | year=1991 | title=Encyclopaedia of Animals: Birds | publisher= Merehurst Press | pages= 144–145 | isbn= 978-1-85391-186-6}} The honey buzzard attacks the nests of social hymenopterans, eating wasp larvae; it is the only known predator of the dangerous{{cite web | url=https://www.theguardian.com/world/2013/sep/26/hornet-attacks-kill-18-china | work=The Guardian | title=Hornet attacks kill dozens in China | date=26 September 2013 |access-date=18 June 2015 }} Asian giant hornet or "yak-killer" (Vespa mandarinia).{{cite book | last1=Cocker | first1=Mark | last2=Mabey | first2=Richard |author2-link=Richard Mabey | title=Birds Britannica | year=2005 |location=London | publisher=Chatto & Windus | pages=113–114 | isbn=978-0-7011-6907-7}} Likewise, roadrunners are the only real predators of tarantula hawk wasps.{{cite web|work=Avian Web|title=Roadrunners|url=http://www.beautyofbirds.com/roadrunners.html|access-date=3 May 2012}}

File:CSIRO ScienceImage 11188 Fig wasp.jpg|Minute pollinating fig wasps, Pleistodontes: the trees and wasps have coevolved and are mutualistic.

File:Latina rugosa planidia.png|Latina rugosa planidia (arrows, magnified) attached to an ant larva; the Eucharitidae are among the few parasitoids able to overcome the strong defences of ants.

File:Goudwesp.jpg|The Chrysididae, such as this Hedychrum rutilans, are known as cuckoo or jewel wasps for their parasitic behaviour and metallic iridescence.

File:Bee wolf.jpg|European beewolf Philanthus triangulum provisioning her nest with a honeybee

File:Clytus arietis (Linné, 1758) (3989861203).jpg|Wasp beetle Clytus arietis is a Batesian mimic of wasps.

File:Pair of Merops apiaster feeding detail.jpg|Bee-eaters such as Merops apiaster specialise in feeding on bees and wasps.

File:Yellowjacket nest found in Kittery, Maine.jpg

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.{{cite web |last1=Gross |first1=Bob |title=Close encounters with stinging insects common in the fall |url=https://www.thetimesherald.com/story/news/local/2017/09/19/close-encounters-stinging-insects-common-fall/680741001/ |website=Times Herald |access-date=August 23, 2021 |date=September 19, 2017}}{{cite web |last1=Boyles |first1=Margaret |title=Yellow Jacket Alert: Taking the Sting Out of Fall |url=https://www.almanac.com/yellow-jacket-alert-taking-sting-out-fall |website=The Old Farmer's Almanac |access-date=August 23, 2021 |date=August 18, 2021}}{{cite web |last1=Hayes |first1=Kim |title=What's That Buzz? Fall may bring more aggressive hornets and wasps |url=https://www.aarp.org/home-family/your-home/info-2017/hornets-aggressive-early-autumn-fd.html |website=AARP |access-date=August 23, 2021 |date=October 11, 2017}} 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.{{cite web|title=Wasps|url=http://www.bpca.org.uk/pages/?page_id=226|publisher=British Pest Control Association|access-date=5 August 2015}} Stings are usually painful rather than dangerous, but in rare cases, people may suffer life-threatening anaphylactic shock.{{cite web|title=Allergy to Wasp and Bee Stings|url=https://www.allergyuk.org/allergy-to-wasp-and-bee-stings/allergy-to-wasp-and-bee-stings|publisher=Allergy UK|access-date=5 August 2015|archive-date=11 August 2015|archive-url=https://web.archive.org/web/20150811075106/http://www.allergyuk.org/allergy-to-wasp-and-bee-stings/allergy-to-wasp-and-bee-stings|url-status=dead}}

{{further|Biological pest control}}

Some species of parasitic wasp, especially in groups such as Aphelinidae, Braconidae, Mymaridae, and Trichogrammatidae, are exploited commercially to provide biological control of insect pests.{{cite web | title=Wasp | url=http://animals.nationalgeographic.com/animals/bugs/wasp/ | archive-url=https://web.archive.org/web/20101005012148/http://animals.nationalgeographic.com/animals/bugs/wasp/ | url-status=dead | archive-date=5 October 2010 | publisher=National Geographic| date=9 November 2010 }}{{cite web | title=New wasp parasite being studied | url=http://www.royalsociety.org.nz/2000/04/20/122/ | work=The Royal Society of New Zealand | date=20 April 2000 | access-date=15 July 2013 | archive-date=4 March 2016 | archive-url=https://web.archive.org/web/20160304080628/http://www.royalsociety.org.nz/2000/04/20/122/ | url-status=dead }} One of the first species to be used was Encarsia formosa, a parasitoid of a range of species of whitefly. It entered commercial use in the 1920s in Europe, was overtaken by chemical pesticides in the 1940s, and again received interest from the 1970s. Encarsia is being tested in greenhouses to control whitefly pests of tomato and cucumber, and to a lesser extent of aubergine (eggplant), flowers such as marigold, and strawberry.{{cite web | last1=Hoddle | first1=Mark | title=Encarsia formosa | url=http://www.biocontrol.entomology.cornell.edu/parasitoids/encarsia.php | publisher=Cornell University | access-date=12 June 2015}} Several species of parasitic wasp are natural predators of aphids and can help to control them.{{cite web |title=Aphid Predators |url=https://www.rhs.org.uk/advice/profile?pid=507| publisher=Royal Horticultural Society |access-date=5 August 2015}} For instance, Aphidius matricariae is used to control the peach-potato aphid.{{cite book |last1=Adams |first1=C.R. |author2=K.M. Bamford |author3=M.P. Early |title=Principles of Horticulture |url=https://books.google.com/books?id=cQbLBAAAQBAJ&pg=PA71 |date=22 October 2013 |publisher=Elsevier |isbn=978-1-4831-4184-8 |pages=71–72}}

File:Encarsia formosa, an endoparasitic wasp, is used for whitefly control.jpg|Encarsia formosa, a parasitoid, is sold commercially for biological control of whitefly, an insect pest of tomato and other horticultural crops.

File:Tomate Blatt Eier Weiße Fliege parasitiert.jpg|Tomato leaf covered with nymphs of whitefly parasitised by Encarsia formosa

Wasps RFC was an English professional rugby 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.{{cite web | url=http://www.wasps.co.uk/History1.ink | title=History 1867–1930 London Wasps | publisher=Wasps.co.uk |archive-url=https://web.archive.org/web/20140722215008/http://www.wasps.co.uk/History1.ink |archive-date=22 July 2014}} The club has an amateur side called Wasps FC.{{cite web|title=Wasps Football Club|url=http://www.pitchero.com/clubs/waspsfootballclub/|publisher=Pitcher|access-date=5 August 2015}}

Among the other clubs bearing the name are a basketball club in Wantirna, Australia,{{cite web|title=Wantirna Wasps Basketball Club|url=http://www.waspsbb.com.au/history|access-date=5 August 2015}} and Alloa Athletic F.C., a football club in Scotland.{{cite web |url=http://spfl.co.uk/clubs/alloa-athletic/ |title=Alloa Athletic Football Club |publisher=Scottish Professional Football League |access-date=10 September 2015}}

File:Polaire, French actress 5.jpg, c. 1900, demonstrated by Polaire, a French actress famous for this silhouette]]

Wasps have been modelled in jewellery since at least the nineteenth century, when diamond and emerald wasp brooches were made in gold and silver settings.{{cite web | url=http://www.alvr.com/2068/diamond-and-emerald-wasp-brooch-by-fontanna/ | title=Diamond and emerald wasp brooch by Fontanna | publisher=A La Vieille Russie | access-date=10 June 2015 | archive-url=https://web.archive.org/web/20150610204657/http://www.alvr.com/2068/diamond-and-emerald-wasp-brooch-by-fontanna/ | archive-date=10 June 2015 | quote=Crown rose diamond and emerald wasp brooch set in silver and gold. By Fontana French, ca. 1875. Width: 3 inches. $46,500}} A fashion for wasp waisted female silhouettes with sharply cinched waistlines emphasizing the wearer's hips and bust arose repeatedly in the nineteenth and twentieth centuries.{{cite web | last=Kunzle | first=David | url=http://www.corsets.de/Data_from_paper_patterns_advertisem.php | title=Fashion and Fetishism | access-date=17 June 2015 | archive-url=https://web.archive.org/web/20141121022841/http://www.corsets.de/Data_from_paper_patterns_advertisem.php | archive-date=21 November 2014}}{{cite book | last=Klingerman |first=Katherine Marie | url=http://etd.lsu.edu/docs/available/etd-04072006-115441/unrestricted/Klingerman_thesis.pdf | title=Binding Femininity: The Effects of Tightlacing on the Female Pelvis | publisher=University of Vermont (MA Thesis) | date=May 2006 | access-date=17 June 2015 | archive-date=21 February 2017 | archive-url=https://web.archive.org/web/20170221164857/http://etd.lsu.edu/docs/available/etd-04072006-115441/unrestricted/Klingerman_thesis.pdf | url-status=dead }}

The Ancient Greek playwright Aristophanes wrote the comedy play Σφῆκες (Sphēkes), The Wasps, first put on in 422 BC. The "wasps" are the chorus of old jurors.{{cite web | title=Ancient Greece – Aristophanes – The Wasps | url=http://www.ancient-literature.com/greece_aristophanes_wasps.html | access-date=10 June 2015}}

H. G. Wells made use of giant wasps in his novel The Food of the Gods and How It Came to Earth (1904):{{cite book | last=Wells | first=H. G. | title=Food of the Gods | url=https://books.google.com/books?id=zygmCQAAQBAJ&pg=PT7867 | date=1904 | publisher=Macmillan}}

{{quote|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 emptied both 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.}}

File:Botticelli-Venus and Mars (cropped).jpg's Venus and Mars, 1485, with a wasp's nest on right, probably a symbol of the Vespucci family (Italian vespa, wasp) who commissioned the painting.{{cite book |last=Lightbown |first=Ronald |author-link=Ronald Lightbown |title=Sandro Botticelli: Life and Work |year=1989 |publisher=Thames and Hudson |pages=165–168 |isbn=978-0896599314}}]]

Wasp (1957) is a science fiction book by the English writer Eric Frank Russell; it is generally considered Russell's best novel.{{cite book | last=Russell |first=Eric Frank | author-link=Eric Frank Russell | title=Wasp | publisher=Gollancz Science Fiction | date=1957 | isbn=978-0-575-07095-0 | url=https://archive.org/details/wasp00russ }} In Stieg Larsson's book The Girl Who Played with Fire (2006) and its film 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.{{cite book |last1=Rosenberg |first1=Robin S. |last2=O'Neill |first2=Shannon |last3=McDonald-Smith |first3=Lynne |title=The Psychology of the Girl with the Dragon Tattoo: Understanding Lisbeth Salander and Stieg Larsson's Millennium Trilogy |url=https://books.google.com/books?id=XJgWYDTG88wC&pg=PT34 |year=2013 |publisher=BenBella Books |isbn=978-1-936661-35-0|page=34}}

Parasitoidal wasps played an indirect role in the nineteenth-century evolution debate. The Ichneumonidae contributed to Charles Darwin's doubts about the nature and existence of a well-meaning and all-powerful Creator. In an 1860 letter to the American naturalist Asa Gray, Darwin wrote:

{{quote|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.{{cite web| url=http://www.darwinproject.ac.uk/entry-2814 | title=Letter 2814 — Darwin, C. R. to Gray, Asa | date=22 May 1860 | access-date=4 May 2011}}}}

File:HMS Wasp (1880).jpg warships to bear the name]]

With its powerful sting and familiar appearance, the wasp has given its name to many ships, aircraft and military vehicles. Nine ships and one shore establishment of the Royal Navy have been named {{HMS|Wasp}}, the first an 8-gun sloop launched in 1749.{{Cite Colledge2006}}

Eleven ships of the United States Navy have similarly borne the name {{USS|Wasp}}, the first a merchant schooner acquired by the Continental Navy in 1775.{{cite book | author=USS Wasp Veterans | title=USS Wasp | url=https://books.google.com/books?id=I6DKJQMZFxQC&pg=PP10| date=15 June 1999| publisher=Turner Publishing | isbn=978-1-56311-404-5 | pages=8–11}} The eighth of these, an aircraft carrier, gained two Second World War battle stars, prompting Winston Churchill to remark "Who said a Wasp couldn't sting twice?"{{cite web|title=USS Wasp. History|url=http://www.public.navy.mil/surflant/lhd1/Pages/History.aspx|publisher=United States Navy|access-date=1 July 2016|archive-date=10 July 2016|archive-url=https://web.archive.org/web/20160710003103/http://www.public.navy.mil/surflant/lhd1/Pages/History.aspx|url-status=dead}} In the Second World War, a German self-propelled howitzer was named Wespe,{{cite book | last1=Jentz | first1=Thomas L. | last2=Doyle | first2=Hilary Louis | title=Panzer Tracts No. 23: Panzer Production From 1933 to 1945 | date=2011 | publisher=Panzer Tracts}} while the British developed the Wasp flamethrower from the Bren Gun Carrier.{{cite book | last=Bishop | year=2002 | first=Chris | title=The Encyclopedia of Weapons of World War II | publisher=Sterling Publishing | isbn=9781586637620 | page=272}}

In aerospace, the Westland Wasp was a military helicopter developed in England in 1958 and used by the Royal Navy and other navies.{{cite book | last=James |first=Derek N. | title=Westland Aircraft since 1915 | publisher=Putnam | date=1991 | isbn=978-0-85177-847-1 | page=365}} The AeroVironment Wasp III is a miniature UAV developed for United States Air Force special operations.{{cite web | url=https://www.af.mil/About-Us/Fact-Sheets/Display/Article/104480/wasp-iii/ | title=US Air Force Wasp III Fact Sheet | access-date=12 June 2015 | url-status=live |archive-url=https://web.archive.org/web/20150923171712/http://www.af.mil/AboutUs/FactSheets/Display/tabid/224/Article/104480/wasp-iii.aspx |archive-date=23 September 2015}}

• Characteristics of common wasps and bees
• Bee and wasp stings
• Schmidt sting pain index
• Fear of wasps (spheksophobia)

{{notelist}}

{{reflist|32em}}

• {{Cite book | title=The Social Biology of Wasps | last=Ross | first=Kenneth G. | publisher=Cornell Press | year=1991 | isbn=978-0-801-49906-7}}

{{Commons}}

{{Commons category|Wasps}}

{{EB1911 poster|Wasp}}

• [http://www.beekeeping.isgood.ca/resources/differences-between-bees-and-wasps/ Differences between Bees and Wasps]
• [https://www.bbc.co.uk/news/science-environment-49692974 Natural History Museum – Wasps: If you can't love them, at least admire them]
• [https://www.nlm.nih.gov/medlineplus/ency/article/000033.htm N.I.H. Medline Encyclopedia] – Insect bites and stings
• [http://www.waspweb.org/Megalyroidea/Megalyridae/index.htm Waspweb]
• {{DermNet|arthropods/bites}}

{{Hymenoptera|2}}

{{Eusociality}}

{{Insects in culture}}

{{Authority control}}

Category:Biological pest control wasps

Category:Extant Jurassic first appearances

Category:Insects in culture

Category:Paraphyletic groups

Category:Pest insects