Vespa

Superregnum: Eukaryota
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Cladus: Metapterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Endopterygota
Superordo: Hymenopterida
Ordo: Hymenoptera
Subordo: Apocrita
Superfamilia: Vespoidea

Familia: Vespidae
Subfamilia: Vespinae
Genus: Vespa
Species (22 + †7): V. affinis – V. analis – V. basalis – V. bellicosa – V. bicolor – †V. bilineata – V. binghami – †V. ciliata – †V. cordifera – V. crabro – †V. crabroniformis – †V. dasypodia – V. ducalis – V. dybowskii – V. fervida – V. fumida – V. luctuosa – V. mandarinia – V. mocsaryana – V. multimaculata – †V. nigra – V. orientalis – V. philippinensis – †V. picea – V. simillima – V. soror – V. tropica – V. velutina – V. vivax

Source: Catalogue of Life: 2013 Annual Checklist
Name

Vespa Linnaeus, 1758

Synonyms

Nyctovespa van der Vecht, 1959
Original status: valid subgenus of Vespa Linnaeus, 1758
Gender: feminine
Type species: Vespa binghami du Buysson, 1905
Fixation: original designation [p. 210] [and monotypy]
Authority for synonymy: Carpenter, 1987: 426–427 ('syn.n.')

References

Carpenter, J.M. 1987: Phylogenetic relationships and classification of the Vespinae (Hymenoptera: Vespidae). Systematic entomology, 12(4): 413–431. DOI: 10.1111/j.1365-3113.1987.tb00213.x
Carpenter, J.M. et al. 2011: Taxonomic notes on the Vespinae of Yunnan (Hymenoptera: Vespidae). American Museum novitates, (3709) DOI: 10.1206/3709.2
Perrard, A. et al. 2013: Phylogeny of hornets: a total evidence approach (Hymenoptera, Vespidae, Vespinae, Vespa). Journal of Hymenoptera research, 32: 1–15. DOI: 10.3897/JHR.32.4685
Vecht, J. van der 1959: Notes on oriental Vespinae, including some species from China and Japan (Hymenoptera, Vespidae). Zoologische Mededelingen, 36(13): 205–232.

Vernacular names
беларуская: Шэршні
čeština: Sršeň
Deutsch: Hornissen
English: Hornets
suomi: Herhiläiset
français: Frelon
hrvatski: Stršljeni
Bahasa Indonesia: Tabuhan
日本語: スズメバチ属
lietuvių: Širšės
latviešu: Sirseņi
Bahasa Melayu: Tebuan
Nederlands: Hoornaars
русский: Ше́ршни
svenska: Bålgetingar
Türkçe: Eşek arısı
українська: Шершень
中文: 胡蜂亞科

Hornets (insects in the genus Vespa) are the largest of the eusocial wasps, and are similar in appearance to their close relatives yellowjackets. Some species can reach up to 5.5 cm (2.2 in) in length. They are distinguished from other vespine wasps by the relatively large top margin of the head and by the rounded segment of the abdomen just behind the waist. Worldwide, 22 species of Vespa are recognized.[2][3] Most species only occur in the tropics of Asia, though the European hornet (V. crabro), is widely distributed throughout Europe, Russia, North America, and Northeast Asia. Wasps native to North America in the genus Dolichovespula are commonly referred to as hornets (e.g., baldfaced hornets), but are actually yellowjackets.

Like other social wasps, hornets build communal nests by chewing wood to make a papery pulp. Each nest has one queen, which lays eggs and is attended by workers that, while genetically female, cannot lay fertile eggs. Most species make exposed nests in trees and shrubs, but some (such as Vespa orientalis) build their nests underground or in other cavities. In the tropics, these nests may last year-round, but in temperate areas, the nest dies over the winter, with lone queens hibernating in leaf litter or other insulative material until the spring. Male hornets are docile and do not have stingers.

Hornets are often considered pests, as they aggressively guard their nesting sites when threatened and their stings can be more dangerous than those of bees.[4]

Classification
European hornet with the remnants of a honey bee.
Main article: Characteristics of common wasps and bees

While taxonomically well defined, some confusion may remain about the differences between hornets and other wasps of the family Vespidae, specifically the yellowjackets, which are members of the same subfamily. Also, a related genus of Asian nocturnal vespines, Provespa, is referred to as "night wasps" or "night hornets",[5] though they are not true hornets.[6]

Some other large wasps are sometimes referred to as hornets, most notably the bald-faced hornet (Dolichovespula maculata) found in North America. It is set apart by its black and ivory coloration. The name "hornet" is used for this species primarily because of its habit of making aerial nests (similar to some of the true hornets) rather than subterranean nests. Another example is the Australian hornet (Abispa ephippium), which is actually a species of potter wasp.
Distribution

Hornets are found mainly in the Northern Hemisphere. The common European hornet (V. crabro) is the best-known species, widely distributed in Europe (but is never found north of the 63rd parallel), Ukraine, and European Russia (except in extreme northern areas). In the east, the species' distribution area stretches over the Ural Mountains to western Siberia (found in the vicinity of Khanty-Mansiysk). In Asia, the common European hornet is found in southern Siberia, as well as in eastern China. The common European hornet was accidentally introduced to eastern North America about the middle of the 19th century and has lived there since at about the same latitudes as in Europe. However, it has never been found in western North America.

The Asian giant hornet (V. mandarinia) lives in the Primorsky Krai, Khabarovsky Krai (southern part), and Jewish AO regions of Russia, and China, Korea, Taiwan, Cambodia, Laos, Vietnam, Indochina, India, Nepal, Sri Lanka, and Thailand, but is most commonly found in the mountains of Japan, where they are commonly known as the giant sparrow bee.

The Oriental hornet (V. orientalis) occurs in semidry, subtropical areas of Central Asia (Armenia, Dagestan in Russia, Iran, Afghanistan, Oman, Pakistan, Turkmenistan, Uzbekistan, Tajikistan, Kyrghyzstan, southern Kazakhstan), and southern Europe (Italy, Malta, Albania, Romania, Turkey, Greece, Bulgaria, Cyprus).[7]

The Asian hornet (V. velutina) has been introduced to France, Spain, Portugal, and Italy.

Stings

Hornets have stingers used to kill prey and defend nests. Hornet stings are more painful to humans than typical wasp stings because hornet venom contains a large amount (5%) of acetylcholine.[8][9] Individual hornets can sting repeatedly; unlike honey bees, hornets do not die after stinging because their stingers are very finely barbed (only visible under high magnification) and can easily be withdrawn, so are not pulled out of their bodies when disengaging.

The toxicity of hornet stings varies according to hornet species; some deliver just a typical insect sting, while others are among the most venomous known insects.[10] Single hornet stings are not in themselves fatal, except sometimes to allergic victims.[10] Multiple stings by hornets (other than V. crabro) may be fatal because of highly toxic species-specific components of their venom.[11]

The stings of the Asian giant hornet (V. mandarinia) are among the most venomous known,[10] and are thought to cause 30–50 human deaths annually in Japan. Between July and September 2013, hornet stings caused the death of 42 people in China.[12] Asian giant hornet's venom can cause allergic reactions and multiple organ failure leading to death, though dialysis can be used to remove the toxins from the bloodstream.[12]

People who are allergic to wasp venom may also be allergic to hornet stings. Allergic reactions are commonly treated with epinephrine (adrenaline) injection using a device such as an epinephrine autoinjector, with prompt follow-up treatment in a hospital. In severe cases, allergic individuals may go into anaphylactic shock and die unless treated promptly.[13] In general, Vespa stings induce the release of histamine due to the various mastoparans that they contain.[Herrera et al 2020 1] However V. orientalis mastoparan is the interesting exception because it does not induce histamine increase in victim tissue[Jalaei et al 2016 1] - because it does not cause mast cell degranulation[Jalaei et al 2016 2] - and is not immunogenic.[Jalaei et al 2016 3]
Attack pheromone

Hornets, like many social wasps, can mobilize the entire nest to sting in defense, which is highly dangerous to humans and other animals. The attack pheromone is released in case of threat to the nest. In the case of the Asian giant hornet (V. mandarinia), this is also used to mobilize many workers at once when attacking colonies of their prey, honey bees and other Vespa species.[14] Three biologically active chemicals, 2-pentanol, 3-methyl-1-butanol, and 1-methylbutyl 3-methylbutanoate, have been identified for this species. In field tests, 2-pentanol alone triggered mild alarm and defensive behavior, but adding the other two compounds increased aggressiveness in a synergistic effect.[14] In the European hornet (V. crabro) the major compound of the alarm pheromone is 2-methyl-3-butene-2-ol.[15]

If a hornet is killed near a nest, it may release pheromones that can cause the other hornets to attack. Materials that come into contact with these pheromones, such as clothes, skin, and dead prey or hornets, can also trigger an attack, as can certain food flavorings, such as banana and apple flavorings, and fragrances that contain C5 alcohols and C10 esters.[14]
Life cycle
The structure of an incipient nest

In V. crabro, the nest is founded in spring by a fertilized female known as the queen. She generally selects sheltered places such as dark, hollow tree trunks. She first builds a series of cells (up to 50) out of chewed tree bark. The cells are arranged in horizontal layers named combs, each cell being vertical and closed at the top. An egg is then laid in each cell. After 5–8 days, the egg hatches. Over the following two weeks, the larva progresses through five stages of development. During this time, the queen feeds it a protein-rich diet of insects. Then, the larva spins a silk cap over the cell's opening, and during the next two weeks, transforms into an adult, a process called metamorphosis. The adult then eats its way through the silk cap. This first generation of workers, invariably females, now gradually undertakes all the tasks formerly carried out by the queen (foraging, nest building, taking care of the brood, etc.) with the exception of egg-laying, which remains exclusive to the queen.
Life history of Vespa crabro

As the colony size grows, new combs are added, and an envelope is built around the cell layers until the nest is entirely covered, with the exception of an entry hole. To be able to build cells in total darkness, they apparently use gravity to aid them.[clarification needed] At the peak of its population, which occurs in late summer, the colony can reach a size of 700 workers.

At this time, the queen starts producing the first reproductive individuals. Fertilized eggs develop into females (called "gynes" by entomologists), and unfertilized ones develop into males (sometimes called "drones" as with honeybee drones). Adult males do not participate in nest maintenance, foraging, or caretaking of the larvae. In early to mid-autumn, they leave the nest and mate during "nuptial flights".

Other temperate species (e.g., the yellow hornet, V. simillima, or the Oriental hornet, V. orientalis) have similar cycles. In the case of tropical species (e.g., V. tropica), life histories may well differ, and in species with both tropical and temperate distributions (such as the Asian giant hornet, V. mandarinia), the cycle likely depends on latitude.
Diet and feeding

Adult hornets and their relatives (e.g., yellowjackets) feed themselves with nectar and sugar-rich plant foods. Thus, they can often be found feeding on the sap of oak trees, rotting sweet fruits, honey, and any sugar-containing foodstuffs. Hornets frequently fly into orchards to feed on overripe fruit, and tend to gnaw a hole in fruit to become totally immersed in its pulp. A person who accidentally picks fruit with a feeding hornet can be attacked by the disturbed insect.

The adults also attack various insects, which they kill with stings and jaws. Due to their size and the power of their venom, hornets are able to kill large insects such as honey bees, grasshoppers, locusts, and katydids without difficulty. The victim is fully masticated and then fed to the larvae developing in the nest, rather than consumed by the adult hornets. Given that some of their prey are considered pests, hornets may be considered beneficial under some circumstances.

The larvae of hornets produce a sweet secretion containing sugars and amino acids that is consumed by the workers and queens.[16]
Species

While a history of recognizing subspecies exists within many of the Vespa species, the most recent taxonomic revision of the genus treats all subspecific names in the genus Vespa as synonyms, effectively relegating them to no more than informal names for regional color forms.[3]

Vespa affinis
Vespa analis
Vespa basalis
Vespa bellicosa
Vespa bicolor
†Vespa bilineata
Vespa binghami
†Vespa ciliata
†Vespa cordifera
Vespa crabro
†Vespa crabroniformis
†Vespa dasypodia
Vespa ducalis
Vespa dybowskii
Vespa fervida
Vespa fumida
Vespa luctuosa
Vespa mandarinia
Vespa mocsaryana
Vespa multimaculata
†Vespa nigra
Vespa orientalis
Vespa philippinensis
†Vespa picea
Vespa simillima
Vespa soror
Vespa tropica
Vespa velutina
Vespa vivax

Notable species

Asian giant hornet (V. mandarinia) (one of its color forms is also known as the Japanese giant hornet)
Asian hornet (V. velutina) (also known as the yellow-legged hornet or Asian predatory wasp)
black hornet (V. dybowskii)
black-bellied hornet (V. basalis)
black shield wasp (V. bicolor)
European hornet (V. crabro) (also known as the Old World hornet or brown hornet)
greater banded hornet (V. tropica)
lesser banded hornet (V. affinis)
Oriental hornet (V. orientalis)
yellow hornet (V. simillima) (one of its color forms is also known as the Japanese yellow hornet or Japanese hornet)
Vespa luctuosa (a species which has the most lethal wasp venom (per volume))[10]

As food and medicine

Hornet larvae are widely accepted as food in mountainous regions in China. Hornets and their nests are treated as medicine in traditional Chinese medicine.
Gallery

Black-bellied hornet (Vespa basalis), Xingping

Asian hornet (Vespa velutina), Shanghai

See also

Australian hornet
Bald-faced hornet
Potter wasp
Vespid
Yellowjacket
Characteristics of common wasps and bees

References

James M. Carpenter & Jun-ichi Kojima (1997). "Checklist of the species in the subfamily Vespinae (Insecta: Hymenoptera: Vespidae)" (PDF). Natural History Bulletin of Ibaraki University. 1: 51–92.
Archer, M.E. (2012). Penney, D. (ed.). Vespine wasps of the world: behaviour, ecology and taxonomy of the Vespinae. Monograph Series. 4. Siri Scientific. ISBN 9780956779571. OCLC 827754341.
A.H. Smith-Pardo, J.M. Carpenter, L. Kimsey (2020) The diversity of hornets in the genus Vespa (Hymenoptera: Vespidae; Vespinae), their importance and interceptions in the United States. Insect Systematics and Diversity 4(3) https://doi.org/10.1093/isd/ixaa006
Vetter, Richard S.; Visscher, P. Kirk; Camazine, Scott (1999). "Mass Envenomations by Honey Bees and Wasps". Western Journal of Medicine. 170 (4): 223–227. PMC 1305553. PMID 10344177.
"Untitled Document".
Madl, M (2012). "Notes on the genus Provespa Ashmead, 1903 (Insecta: Hymenoptera: Vespidae: Vespinae) based on the material of the Naturhistorisches Museum Wien (Austria)" (PDF). Annals Naturhistorisches Museum Wien. 114: 27–35. Retrieved 10 March 2016.
Dieter Kosmeier. "Vespa orientalis, Oriental Hornet". vespa-crabro.de.
K. D. Bhoola; J. D. Calle; M. Schachter (1961). "Identification of acetylcholine, 5-hydroxytryptamine, histamine, and a new kinin in hornet venom (V. crabro)". J. Physiol. 159 (1): 167–182. doi:10.1113/jphysiol.1961.sp006799. PMC 1359584. PMID 13868844.
Conniff, R. (June 2003). "Stung – How tiny little insects get us to do exactly as they wish". Discover.
J.O. Schmidt; S. Yamane; M. Matsuura; C.K. Starr (1986). "Hornet venoms: lethalities and lethal capacities". Toxicon. 24 (9): 950–4. doi:10.1016/0041-0101(86)90096-6. PMID 3810666.
P. Barss (1989). "Renal failure and death after multiple stings in Papua New Guinea. Ecology, prevention and management of attacks by vespid wasps". Med J Aust. 151 (11–12): 659–63. doi:10.5694/j.1326-5377.1989.tb139643.x. PMID 2593913. S2CID 22208955.
Park, Madison; Zhang, Dayu; Landau, Elizabeth (October 3, 2013). "Deadly giant hornets kill 42 people in China". CNN. Archived from the original on October 4, 2013.
"Insect bites and stings: MedlinePlus Medical Encyclopedia".
"Volatile fragrance chemicals may attract unwanted attention from hornets and bees".
"Vespa". pherobase.com.

Wheeler, W.M. (1922) Social Life Among the Insects. II. The Scientific Monthly 15(1): 68–88

Jalaei, Jafar; Fazeli, Mehdi; Rajaian, Hamid; Ghalehsoukhteh, Somayeh Layeghi; Dehghani, Alireza; Winter, Dominic (2016). "In vitro antihistamine-releasing activity of a peptide derived from wasp venom of Vespa orientalis". Asian Pacific Journal of Tropical Biomedicine. Medknow. 6 (3): 259–264. doi:10.1016/j.apjtb.2015.12.001. ISSN 2221-1691.

p. 263, "Cytotoxicity data from a LDH release assay, which measure damage to the plasma membrane showed no significant increase in percentage of LDH release and its IC50 was nearly 128 μL, so we confirmed that LDH absorbance was not affected by this peptide."
p. 262, "Moreover, the results showed the IC50 of mast cells degranulation at 126 μmol/L, which was approximately high implying that this peptide had high selectivity for normal cells"

p. 262, "implying that this peptide had ... immuno-modulation property which led the prevention of immune system activation (Figure 7)."

Herrera, Cayetano; Leza, Mar; Martínez-López, Emma (2020-07-23). "Diversity of compounds in Vespa spp. venom and the epidemiology of its sting: a global appraisal". Archives of Toxicology. Springer Science and Business Media LLC. 94 (11): 3609–3627. doi:10.1007/s00204-020-02859-3. ISSN 0340-5761. PMID 32700166. S2CID 220716061.

p. 3, "Several studies have shown that mastoparans act on mast cells. ... They are responsible for releasing, mainly, histamine