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Asian giant hornet

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Asian giant hornet
Temporal range: Miocene–present, 15.97–0 Ma
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Vespidae
Genus: Vespa
V. mandarinia
Binomial name
Vespa mandarinia
Smith, 1852[1]
  • Vespa magnifica Smith, 1852
  • Vespa japonica Radoszkowski, 1857
  • Vespa bellona Smith, 1871
  • Vespa magnifica var. latilineata Cameron, 1903
  • Vespa mandarina Dalla Torre, 1894 (misspelling)
  • Vespa mandarinia nobilis Sonan, 1929
  • Vespa magnifica sonani Matsumura, 1930

The Asian giant hornet (Vespa mandarinia) or northern giant hornet,[2][3] including the color form referred to as the Japanese giant hornet,[4][5] is the world's largest hornet. It is native to temperate and tropical East Asia, South Asia, Mainland Southeast Asia, and parts of the Russian Far East. It was also found in the Pacific Northwest of North America in late 2019[6][7] with a few more additional sightings in 2020,[8][9] and nests found in 2021,[10][11] prompting concern that it could become an invasive species.[12][Ala 1] However, by the end of the season in November 2022, there were no confirmed sightings in North America at all,[13] suggesting they may have been eradicated in that region.[14]

Asian giant hornets prefer to live in low mountains and forests, while almost completely avoiding plains and high-altitude climates. V. mandarinia creates nests by digging, co-opting pre-existing tunnels dug by rodents, or occupying spaces near rotten pine roots.[15][Arc 1] It feeds primarily on larger insects, colonies of other eusocial insects, tree sap, and honey from honey bee colonies.[16] The hornet has a body length of 45 mm (1+34 in), a wingspan around 75 mm (3 in), and a stinger 6 mm (14 in) long, which injects a large amount of potent venom.[17]

Taxonomy and phylogeny

form "magnifica"

V. mandarinia is a species in the genus Vespa, which comprises all true hornets. Along with seven other species, V. mandarinia is a part of the V. tropica species group, defined by the single notch located on the apical margin of the seventh gastral sternum of the male. The most closely related species within the species group is V. soror.[Arc 2][Arc 3] The triangular shape of the apical margin of the clypeus of the female is diagnostic, the vertex of both species is enlarged, and the shape of the apex of the aedeagus is distinct and similar.[18]

Division of the genus into subgenera has been attempted in the past,[19] but has been abandoned, due to the anatomical similarity among species and because behavioral similarity is not associated with phylogeny.[15] The species has existed since the Miocene epoch, as indicated by fossils found in the Shanwang Formation.[20]

As of 2012, three subspecies were recognized:[21] V. m. mandarinia, V. m. magnifica, and V. m. nobilis. The former subspecies referred to as V. m. japonica has not been considered valid since 1997.[22] The most recent revision in 2020 eliminated all of the subspecies rankings entirely, with "japonica", "magnifica", and "nobilis" now relegated to informal non-taxonomic names for different color forms.[4]

Common names


Since its discovery in North America, the scientific literature and official government sources refer to this species by its established common name, Asian giant hornet, whilst the mainstream media have taken to using the nickname "murder hornet".[23][24][25] In July 2022, the Entomological Society of America stated that they will adopt the common name northern giant hornet for the species to avoid potentially discriminatory language, citing Xenophobia and racism related to the COVID-19 pandemic in the wake of the COVID-19 pandemic.[2][26][27][28]


Head detail
Hornet specimen held in a human hand to illustrate its size

Regardless of sex, the hornet's head is a light shade of orange and its antennae are brown with a yellow-orange base. Its eyes and ocelli are dark brown to black. V. mandarinia is distinguished from other hornets by its pronounced clypeus and large genae. Its orange mandible contains a black tooth that it uses for digging.[29] The thorax is dark brown, with two grey wings varying in span from 35 to 76 mm (1+38 to 3 in).[29]

Its fore legs are brighter than the mid and hind legs. The base of the fore legs is darker than the rest. The abdomen alternates between bands of dark brown or black, and a yellow-orange hue (consistent with its head color). The sixth segment is yellow. Its stinger is typically 6 mm (14 in) long and delivers a potent venom that, in cases of multiple hornets stinging simultaneously, can kill a human.[29]

Queens and workers


The queens are considerably larger than workers. Queens can exceed 50 mm (2 in), while workers are between 35 and 40 mm (1+25 and 1+35 in). The reproductive anatomy is consistent between the two, but workers do not reproduce.[29]



Drones (males) are similar to females, and can attain 38 millimetres (1+12 in) in length, but lack stingers. This is a consistent feature among the Hymenoptera.[29]



Larvae spin a silk cocoon when they complete development and are ready to pupate.[30] Larval silk proteins have a wide variety of potential applications due to their wide variety of potential morphologies, including the native fiber form, but also sponge, film, and gel.[30]



The mitochondrial genome is provided by Chen et al., 2015.[31] This data has also been important to confirm the place of the wider Vespidae family in the Vespoidea superfamily, and confirms that Vespoidea is monophyletic.[31]



Within two days of the initial 2020 news report on V. mandarinia, insect identification centers in the eastern United States (where the wasp does not occur) began getting identification requests, and were swamped for the next several months, even though not one of the thousands of submitted photos or samples was of V. mandarinia, but were instead primarily wasps such as the European hornet (V. crabro), the eastern cicada killer (Sphecius speciosus), or the southern yellowjacket (Vespula squamosa).[32][33]

Submissions suspected by laypeople to be V. mandarinia also include other wasps of various sizes, bees, sawflies, horntails, wasp-mimicking flies, beetles, Jerusalem crickets, cicadas, and even a plastic children's toy that was wasp-like in appearance, all of which were routinely estimated to be 130-185% of their actual size.[32]

Reports of this species from other parts of the world appear to be erroneous identifications of other introduced hornet species, such as V. orientalis in several locations around the world, and V. velutina in Europe.[34]



Ecological distribution


V. mandarinia is primarily a forest dweller.[35][36] When it does live in urban landscapes, V. mandarinia is highly associated with green space.[35][Ala 2] It is the most dependent upon green space of the Vespa species (with V. analis the least).[35] Extremely urbanized areas provide a refuge for V. analis, whereas V. mandarinia – its predator – is entirely absent.[35]

Geographic distribution

An Asian giant hornet



The Asian giant hornet can be found in:

North America


The first confirmed sightings of the Asian giant hornet in North America were confirmed in 2019 and were mainly concentrated in the Vancouver area, with nests also discovered in neighboring Whatcom County, Washington, in the United States.

  • In August 2019, three hornets were found in Nanaimo on Vancouver Island, and a large nest was found and destroyed shortly thereafter;[39][40]
  • At the end of September, a worker was reported in Blaine, Washington.[41]
  • Another worker was found in Blaine in October;[41]
  • In December 2019, another worker was found in Blaine;[8]
  • Two specimens were collected in May 2020, one from Langley, British Columbia, about 13 kilometres (8 miles) north of Blaine, and one from Custer, Washington, 14 km (9 mi) southeast of Blaine.[41]
  • One queen sighting in June 2020, from Bellingham, Washington, 24 km (15 mi) south of Custer[41]
  • An unmated queen was trapped in July 2020, near Birch Bay, Washington, 10 km (6 mi) west of Custer.[8]
  • A male hornet was captured in Custer, Washington in July 2020.[42]
  • A hornet of unknown caste was reported in August 2020, in Birch Bay, and another was trapped in the same area the following day.[41]
  • Three hornets were seen (and two killed) southeast of Blaine on 21 and 25 September 2020,[43] and three more were found in the same area on 29 and 30 September,[44] prompting officials to report that attempts were underway to pinpoint and destroy a nest believed to be in the area.[45]
  • In October 2020, the Washington State Department of Agriculture announced that a nest was found 2.5 metres (8.3 ft) above ground[46] in a cavity of a tree in Blaine, with dozens of hornets entering and leaving.[47] The nest was eradicated the next day, including the immediate discovery and removal of about 100 hornets.[48][49][50] At first the owner of the land required the nest to be returned, and he advertised it for sale.[51] A local beekeeper bought it from him and gave it back to the state entomology team.[51] After further analysis, it was determined that the nest had contained about 500 live specimens, including about 200 queens.[46][52] Some of these specimens were sent to the Smithsonian Institution to become a part of the NMNH Biorepository permanent cryogenic collection.[53][54] It was announced that several undiscovered live nests were also believed to exist within Washington State, because the captures of individual hornets in Birch, Blaine, and Custer were all relatively far from the discovered nest.[46][52][55] However, officials expressed cautious optimism, adding that it might still be possible to eradicate the hornets before they became established in the area.[46] A Canadian official said that although individual specimens had been found in Canada and some nests were suspected to exist there, the hornets' presence seemed to be only in areas near the US-Canadian border, while the center of the invasion appeared to be in Washington State.[46]
  • In November 2020, one individual was found in Abbotsford, BC.[56][57][58][59] As a result the BC government asked Abbotsford beekeepers and residents to report any sightings.[60][61]
  • In November 2020, a queen was found in Aldergrove, BC.[62][63][64][65][66][67]
  • In August 2021, a nest was discovered in Whatcom County, Washington near Blaine, only 2 miles (3.2 km) from the nest WSDA eradicated in 2020.[10][68] This nest was destroyed two weeks later on 25 August, before it could produce new queens.[69]
  • In September 2021, two more nests were found near Blaine, in the vicinity of the nest found in August,[70][11] and a "potential sighting" was reported from near Everson, some 25 miles east of Blaine.[71]

A mitochondrial DNA analysis was performed to determine the maternal population(s) ancestral to the British Columbia and Washington introduced populations.[Wil 1] The high dissimilarity between these two was similar to the mutual distances between each of the Chinese, Japanese, and Korean native populations[Wil 2] suggesting the specimens collected in 2019 were from two different maternal populations,[Wil 3] Japanese in BC[Wil 4] and South Korean in Washington.[Wil 5] This suggests that two separate introductions of the Asian giant hornet occurred in North America within about 80 km (50 mi) of one another within a few months.

In April 2020, authorities in Washington State asked members of the public to be alert and report any sightings of these hornets, which are expected to become active in April if they are in the area.[72] If they become established, the hornets "could decimate bee populations in the United States and establish such a deep presence that all hope for eradication could be lost." A "full-scale hunt" for the species by the WSDA was then underway.[23] Two assessment models of their potential to spread from their present location on the US–Canadian border suggested that they could spread northward into coastal British Columbia and Southeast Alaska, and southward as far as southern Oregon.[12][Ala 3] The USDA's Agricultural Research Service is engaged in lure/attractant development and molecular genetics research, both as part of its normal research mission, but also to further the near-term eradication goal in Washington.[73]

In 2020, the United States Congress considered specific legislation to eradicate V. mandarinia[74] including a proposal by the Interior Secretary, the Fish and Wildlife Director, and the other relevant agencies, which has been introduced as an amendment to the appropriations omnibus.[75][76] British Columbia Agriculture is prepared for a "long fight" lasting years, if necessary.[77] One advantage humans will have is the lack of diversity of such an invasive population – leaving the hornets less prepared for novel environments and challenges.[77]

In June 2021 a dead, desiccated male was found near Marysville, Snohomish County, Washington and reported to WSDA. Its different, more reddish color form immediately suggested yet another parental population from the Japanese and Korean ones already known. USDA APHIS (Animal and Plant Health Inspection Service) performed a genetic analysis several days later and, together with WSDA, confirmed it was of a third, unrelated population. The discovery of a male in June is "perplexing" given that the earliest male emergence in 2020 was July, which was already earlier than normal for the home range. This and its desiccated state indicate it did not emerge in 2021 at all, but is instead a dead specimen that had already emerged in a previous year.[78]

The WSDA announced in December of 2022 that there were "no confirmed sightings" of the hornet in the state for that year,[13] and in December of 2023 stated there were no sightings in 2023, and went on to say that if there are no sightings in 2024, the species will be declared "eradicated".[79]



V. mandarinia nests in low mountain foothills and lowland forests.[35][Arc 1] As a particularly dominant species, no efforts are directed toward conserving V. mandarinia or its habitats, as they are common in areas of low human disturbance.[35] Unlike other species of Vespa, V. mandarinia almost exclusively inhabits subterranean nests[35][Arc 1] – in 1978 it was still doubted that aerial nests were possible, as Matsuura and Sakagami reported this to be unknown in Japan in 1973[80] and aerial nesting is still described as extremely rare in Japan,[37] and yet as of 2021 all nests in the invasive range have been aerial.[citation needed]

In a study of 31 nests, 25 were found around rotten pine roots, and another study found only 9 of 56 nests above ground.[Arc 1] Additionally, rodents, snakes, or other burrowing animals previously made some of the tunnels.[Arc 1] The depth of these nests was between 6 and 60 cm (2 and 24 in). The entrance at the ground surface varies in length from 2 to 60 cm (1 to 24 in) either horizontally, inclined, or vertically. The queens that found the nest prefer narrow cavities.[36]

Nests of V. mandarinia typically lack a developed envelope. During the initial stages of development, the envelope is in an inverted-bowl shape.[Arc 4] As the nest develops, one to three rough sheets of combs are created. Often, single primordial combs are created simultaneously and then fused into a single comb.[36]

A system of one main pillar and secondary pillars connects the combs. Nests usually have four to seven combs.[Arc 4] The top comb is abandoned after summer and left to rot. The largest comb is at the middle to bottom portion of the nest. The largest combs created by V. mandarinia measured 49.5 by 45.5 cm (19+12 by 18 in) with 1,192 cells (no obstacles, circular) and 61.0 by 48.0 cm (24 by 19 in) (elliptical; wrapped around a root system).[36]

Colony cycle


The nesting cycle of V. mandarinia is fairly consistent with that of other eusocial insects. Six phases occur in each cycle.[36]

Pre-nesting period


Inseminated and uninseminated queens enter hibernation following a cycle. They first appear in early to mid-April and begin feeding on the sap of Quercus (oak) trees. Although this timing is consistent among hornets, V. mandarinia dominates the order, receiving preference for premium sap sources. Among the V. mandarinia queens is a dominance hierarchy. The top-ranked queen begins feeding, while the other queens form a circle around her. Once the top queen finishes, the second-highest-ranking queen feeds. This process repeats until the last queen feeds at a poor hour.[36]

Solitary, cooperative, and polyethic periods


Inseminated queens start to search for nesting sites in late April. The uninseminated queens do not search for nests, since their ovaries never fully develop. They continue to feed, but then disappear in early July.

An inseminated queen begins to create relatively small cells in which she raises around 40 small workers. Workers do not begin to work outside of the hive until July. Queens participate in activities outside the hive until mid-July, when they stay inside the nest and allow workers to do extranidal activities. Early August marks a fully developed nest, containing three combs holding 500 cells and 100 workers. After mid-September, no more eggs are laid and the focus shifts to caring for larvae. The queens die in late October.[36]

Dissolution and hibernating period


Males and new queens take on their responsibilities in mid-September and mid-October, respectively. During this time, their body color becomes intense and the weights of the queens increase about 20%. Once the males and queens leave the nest, they do not return. In V. mandarinia, males wait outside the nest entrance until the queens emerge, when males intercept them in midair, bring them to the ground, and copulate from 8 to 45 seconds. After this episode, the males return to the entrance for a second chance, while the now-mated queens leave to hibernate. Many queens (up to 65%) attempt to fight off the males and leave unfertilized,[37] at least temporarily. After this episode, pre-hibernating queens are found in moist, subterranean habitats.

When sexed individuals emerge, workers shift their focus from protein and animal foods to carbohydrates. The last sexed individuals to emerge may die of starvation.[36]



The stinger of the Asian giant hornet is about 6 mm (14 in) long.[17]



Their stinger injects an especially potent venom that contains mastoparan-M.[81] Mastoparans are found in many bee and wasp venoms.[81] They are cytolytic peptides that can damage tissue by stimulating phospholipase action, in addition to its own phospholipase.[81][36] Masato Ono, an entomologist at Tamagawa University, described the sensation of being stung as feeling "like a hot nail being driven into my leg".[17] Besides using their stingers to inject venom, Asian giant hornets are apparently able to spray venom into a person's eyes under certain circumstances, with one report in 2020 from Japan of long-term damage, though the exact extent of actual visual impairment still remains unassessed.[82]

The venom contains a neurotoxin called mandaratoxin,[36][Abe 1] a single-chain polypeptide with a molecular weight around 20 kDa.[36][Abe 2] While a single wasp cannot inject a lethal dose, multiple stings can be lethal even to people who are not allergic if the dose is sufficient, and allergy to the venom greatly increases the risk of death. Tests involving mice found that the venom falls short of being the most lethal of all wasp venoms, having an LD50 of 4.0 mg/kg. (In comparison, the deadliest wasp venom (at least to laboratory mice) by weight belongs to V. luctuosa at 1.6 mg/kg.) The potency of the V. mandarinia sting is due, rather, to the relatively large amount of venom injected.[83]



Evidence is insufficient to believe that prophylactic immunotherapy for the venom of other Vespidae will prevent allergic reaction to V. mandarinia venom, because of wide differences in venom chemistry.[84]

Effects on humans


In 1957, van der Vecht was under the impression humans in the native range lived in constant fear of V. mandarinia and Iwata reported in 1976 that research and removal were hampered by its attacks.[80]



The strepsipteran Xenos moutoni is a common parasite among Vespa species. In a study of parasites among species of Vespa, 4.3% of V. mandarinia females were parasitized. Males were not stylopized (parasitization by stylopid strepsipterans, such as X. moutoni) at all. The major consequence of being parasitized is the inability to reproduce, and stylopized queens follow the same fate as uninseminated queens. They do not search for an area to create a new colony and feed on sap until early July, when they disappear. In other species of Vespa, males also have a chance of being stylopized. The consequences between the two sexes are similar, as neither sex is able to reproduce.[85]

Communication and perception


V. mandarinia uses both visual and chemical cues as a means of navigating itself and others to the desired location. Scent marking was discussed as a way for hornets to direct other members of the colony to a food source. Even with antennae damage, V. mandarinia was able to navigate itself. It was unable to find its destination only when vision impairment was induced. This implies that while chemical signaling is important, visual cues play an equally important role in guiding individuals. Other behaviors include the formation of a "royal court" consisting of workers that lick and bite the queen, thereby ingesting her pheromones.

These pheromones could directly communicate between the queen and her court or indirectly between her court and other workers due to the ingested pheromones. This is merely speculation, as no direct evidence has been collected to suggest the latter. V. mandarinia communicates acoustically, as well. When larvae are hungry, they scrape their mandibles against the walls of the cell. Furthermore, adult hornets click their mandibles as a warning to other creatures that encroach upon their territories.[29][86]

Scent marking


V. mandarinia is the only species of social wasp known to apply a scent to direct its colony to a food source. The hornet secretes the chemical from the sixth sternal gland, also known as van der Vecht's gland. This behavior is observed during autumnal raids after the hornets begin hunting in groups instead of individually. The ability to apply scents may have arisen because the Asian giant hornet relies heavily on honey bee colonies as its main food source.[87][80]

A single hornet is unable to take on an entire colony of honey bees because species such as Apis cerana have a well-organized defense mechanism. The honey bees swarm one wasp and flutter their wings to heat up the hornet and raise carbon dioxide to a lethal level. So, organized attacks are much more effective and easily devastate a colony of tens of thousands of honey bees.[87][80]

Interspecies dominance


In an experiment observing four different species of Vespa (V. ducalis, V. crabro, V. analis, and V. mandarinia), V. mandarinia was the dominant species. Multiple parameters were set to determine this. The first set parameter observed interaction-mediated departures, which are defined as scenarios wherein one species leaves its position due to the arrival of a more dominant individual. The proportion of interaction-mediated departures was the lowest for V. mandarinia. Another measured parameter was attempted patch entry. Over the observed time, conspecifics (interactions with the same species) resulted in refused entry far more than heterospecifics (interactions with different species).[88]

Lastly, when feeding at sap flows, fights between these hornets, Pseudotorynorrhina japonica, Neope goschkevitschii, and Lethe sicelis were observed, and once more V. mandarinia was the most dominant species. In 57 separate fights, one loss was observed to Neope goschkevitschii, giving V. mandarinia a win rate of 98.3%. Based on interaction-mediated departures, attempted patch entry, and interspecific fights, V. mandarinia is the most dominant Vespa species.[88]


Feeding on a mantis

The Asian giant hornet is intensely predatory; it hunts medium- to large-sized insects, such as bees,[36][89] other hornet and wasp species, beetles, hornworms,[90] and mantises. The latter are favored targets in late summer and fall. Large insects such as mantises are key protein sources to feed queen and drone larvae. Workers forage to feed their larvae, and since their prey can include crop pests, the hornets are sometimes regarded as beneficial.[90]

This hornet often attacks colonies of other Vespa species (V. simillima being the usual prey species), Vespula species,[90] and honey bee (such as Apis cerana and A. mellifera)[90] hives to obtain the adults, pupae, and larvae as food for their own larvae. Sometimes, they cannibalize each other's colonies. A single scout, sometimes two or three, cautiously approaches the hive, producing pheromones to lead its nest-mates to the hive. The hornets can devastate a colony of honey bees, especially if it is the introduced western honey bee. A single hornet can kill as many as 40 bees per minute due to its large mandibles, which can quickly strike and decapitate prey.[91]

The honey bees' stings are ineffective because the hornets are five times their size and heavily armored. Only a few hornets (under 50) can exterminate a colony of tens of thousands of bees in a few hours. The hornets can fly up to 100 km (60 mi) in a single day, at speeds up to 40 km/h (25 mph).[92] The smaller Asian hornet similarly preys on honey bees, and has been spreading throughout Europe.

Hornet larvae, but not adults, can digest solid protein. The adult hornets can only drink the juices of their victims, and they chew their prey into a paste to feed to their larvae. The workers dismember the bodies of their prey to return only the most nutrient-rich body parts, such as flight muscles, to the nest.[5] Larvae of predatory social vespids generally, not just Vespa, secrete a clear liquid, sometimes referred to as Vespa amino acid mixture, the exact amino acid composition of which varies considerably from species to species, and which they produce to feed the adults on demand.[93]

Native honey bees

A defensive ball of Japanese honey bees (A. c. japonica) in which two Japanese hornets (V. simillima xanthoptera) are engulfed, incapacitated, heated, and eventually killed. This sort of defense is also used against the Asian giant hornet.

Beekeepers in Japan attempted to introduce western honey bees (Apis mellifera) for the sake of their high productivity. Western honey bees have no innate defense against the hornets, which can rapidly destroy their colonies.[5] Kakugo virus infection, though, may provide an extrinsic defence.[94] Although a handful of Asian giant hornets can easily defeat the uncoordinated defenses of a western honey bee colony, the Japanese honey bee (Apis cerana japonica) has an effective strategy. When a hornet scout locates and approaches a Japanese honey bee hive, she emits specific pheromonal hunting signals. When the Japanese honey bees detect these pheromones, 100 or so gather near the entrance of the nest and set up a trap, keeping the entrance open.[95]

This permits the hornet to enter the hive. As the hornet enters, a mob of hundreds of bees surrounds it in a ball, completely covering it and preventing it from reacting effectively. The bees violently vibrate their flight muscles in much the same way as they do to heat the hive in cold conditions.[95] This raises the temperature in the ball to the critical temperature of 46 °C (115 °F).[95]

In addition, the exertions of the honey bees raise the level of carbon dioxide (CO2) in the ball.[95] The bees can tolerate up to 50 °C (122 °F) even at that concentration of CO2, but the hornet cannot survive the combination of high temperature and high carbon dioxide level.[95] Some honey bees do die along with the intruder, much as happens when they attack other intruders with their stings, but by killing the hornet scout, they prevent it from summoning reinforcements that would wipe out the entire colony.[96]

Detailed research suggests this account of the behavior of the honey bees and a few species of hornets is incomplete and that the honey bees and the predators are developing strategies to avoid expensive and mutually unprofitable conflict. Instead, when honey bees detect scouting hornets, they transmit an "I see you" signal that commonly warns off the predator.[97] Another defence used by Apis cerana is speeding up dramatically when returning to the colony, to avoid midair attacks.[citation needed]

Diet in North America


The Washington State Department of Agriculture found V. mandarinia to be preying on cluster fly, orange legged drone fly, bristle fly, bronze birch borer beetle, western honey bee, western yellowjacket, German yellowjacket, aerial yellowjacket, bald faced hornet, European paper wasp, golden paper wasp, paddle-tailed darner dragonfly, shadow darner dragonfly, large yellow underwing moth, blinded sphinx moth, and red admiral butterfly (Vanessa atalanta). They had also eaten cow's meat, but the WSDA assumes this to have been beef from a hamburger.[98]



The Asian giant hornet has very few natural predators. However, V. mandarinia nests are attacked by conspecific colonies, and honey buzzards may prey on this hornet.[99] Besides the Honey Buzzard and each other, there are also instances of other insects such as the praying mantis killing Asian giant hornets.[100]



V. mandarinia is not solely carnivorous, but also a pollinator. It is among the diurnal pollinators of the obligate plant parasite Mitrastemon yamamotoi.[101] It is among the most common pollinators of Musella lasiocarpa in the Yunnan Province of China.[102]

Extermination methods


As of 1973, six different methods were used to control hornets in Japan; these methods decrease damage done by V. mandarinia.[citation needed]



Hornets are crushed with wooden sticks with flat heads. Hornets do not counterattack when they are in the bee-hunting phase or the hive-attack phase ("slaughter"), but they aggressively guard a beehive once they kill the defenders and occupy it. The biggest expenditure in this method is time, as the process is inefficient.[36]

Nest removal


Applying poisons or fires at night is an effective way of exterminating a colony. The most difficult part about this tactic is finding the subterranean nests. The most common method of discovering nests is giving a piece of frog or fish meat attached to a cotton ball to a wasp and following it back to its nest. With V. mandarinia, this is particularly difficult considering its common home flight radius of 1–2 kilometres (0.62–1.24 mi). V. mandarinia travels up to 8 kilometres (5.0 mi) away from the nest.[36][103]

For the rare nest that is up in a tree, wrapping the tree in plastic and vacuuming the hornets out is used.[50]

Bait traps


Bait traps can be placed in apiaries. The system consists of multiple compartments that direct the hornet into a one-sided hole which is difficult to return through once it is in the cul-de-sac compartment, an area located at the top of the box from which honey bees can escape through a mesh opening, but wasps cannot due to their large size. Baits used to attract the hornets include a diluted millet jelly solution or a crude sugar solution with a mixture of intoxicants, vinegar, or fruit essence.[36]

The WSDA has been using plastic bottle traps, baited with fruit juice and added alcohol. The alcohol is used because it repels bees, but not V. mandarinia, thus reducing the bycatch.[104]

Mass poisoning


Hornets at the apiary are captured and fed a sugar solution or bee that has been poisoned with malathion. The toxin is expected to spread through trophallaxis. This method is good in principle, but has not been tested extensively.[36]

Trapping at hive entrances


The trap is attached to the front of beehives. The effectiveness of the trap is determined by its ability to capture hornets while allowing honey bees to escape easily. The hornet enters the trap and catches a bee. When it tries to fly back through the entrance of the hive, it hits the front of the trap. The hornet flies upwards to escape and enters the capture chamber, where the hornets are left to die. Some hornets find a way to escape the trap through the front, so these traps can be very inefficient.[36]

Protective screens


As explained in the trapping section, if met by resistance, hornets lose the urge to attack and instead retreat. Different measures of resistance include weeds, wire, or fishing nets or limiting the passage size so only honey bees can make it through. Experienced hornets catch on and eventually stay on these traps, awaiting the arrival of bees. The best method of controlling hornets is to combine protective screens with traps.[36]

Glue traps


Some Japanese beekeepers have found that placing glue traps of the sort commonly used against mice atop the bees' artificial nesting box with a disarmed giant hornet stuck to the glue will attract hornets who try to help the stricken hornet, but they also get trapped on the glue sheet.[105]

Human consumption


In some Japanese mountain villages, the nests are excavated and the larvae are considered a delicacy when fried.[5] In the central Chūbu region, these wasps are sometimes eaten as snacks or an ingredient in drinks. The grubs are often preserved in jars, pan-fried or steamed with rice to make a savory dish called hebo-gohan. The adults are fried on skewers, stinger and all, until the body becomes crunchy.[106]

Economic impact


If V. mandarinia settles all suitable habitats in North America, control costs in the United States will be over US$113.7 million/year (possibly significantly higher).[Ala 4]

Agricultural impact


If V. mandarinia reaches all suitable habitat in North America, bee products would bring in US$11.98 ± 0.64 million less per year, and bee-pollinated crops would produce US$101.8 million less per year.[Ala 5] New York, Massachusetts, Pennsylvania, Connecticut, North Carolina, New Jersey, and Virginia would be most severely affected.[Ala 6] By region, New England would be worst hit, and to a lesser degree the entire northeast and the entirety of eastern North America.[Ala 6] New England would become by far the greatest concentration of V. mandarinia in the world, far surpassing the original introduction site (the Pacific Northwest), and even its home range of East Asia.[Ala 6] Alfalfa/other hays, apples, grapes, tobacco, cotton, and blueberries would be the crops most severely affected.[Ala 7]

See also



  1. ^ Smith, F. (1852). "VIII. Descriptions of some new and apparently undescribed species of hymenopterous insects from North China, collected by Robert Fortune, Esq". Transactions of the Royal Entomological Society of London. 7 (2). Royal Entomological Society: 33–44. doi:10.1111/j.1365-2311.1852.tb02208.x. (Vespa mandarinia: p. 38)
  2. ^ a b
  3. ^ "Giant hornet gets new name". agr.wa.gov. Retrieved 25 July 2022.
  4. ^ a b Smith-Pardo, Allan H; Carpenter, James M.; Kimsey, Lynn; Hines, Heather (May 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). doi:10.1093/isd/ixaa006.
  5. ^ a b c d e Piper, Ross (2007). Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals. Bloomsbury Academic. pp. 9–11. ISBN 978-0-313-33922-6.
  6. ^ BC Gov News: Asian Giant Hornet nest eradicated in Nanaimo.
  7. ^ USDA New Pest Response Guidelines: Vespa mandarinia Asian giant hornet.
  8. ^ a b c "Hornets". Washington State Department of Agriculture.
  9. ^ "WSDA News Releases". Washington State Department of Agriculture.
  10. ^ a b "Washington state has first live 'murder hornet' sighting of 2021". NBC News. 13 August 2021. Retrieved 13 August 2021.
  11. ^ a b "3rd murder hornet nest of 2021 found in northwest Washington". 11 September 2021.
  12. ^ a b Zhu, Gengping; Gutierrez Illan, Javier; Looney, Chris; Crowder, David W. (2020). "Assessing the ecological niche and invasion potential of the Asian giant hornet". Proceedings of the National Academy of Sciences. 117 (40): 24646–24648. Bibcode:2020PNAS..11724646Z. doi:10.1073/pnas.2011441117. PMC 7547231. PMID 32963093.
  13. ^ a b Bartick, Alex (6 December 2022). "No northern giant hornets found in Washington in 2022". KOMO News (Seattle, Washington). Retrieved 7 December 2022.
  14. ^ Murder Hornets Appear to Have Been Driven Out of the U.S. - Newsweek 10/17/22
  15. ^ a b Yamane, Seiki (July 1976). "Morphological and taxonomic studies on vespine larvae, with reference to the phylogeny of the subfamily Vespinae (Hymenoptera: Vespidae)". Insecta Matsumurana. Entomology. 8. Faculty of Agriculture Hokkaido University HUSCAP: 1–45. hdl:2115/9782. Retrieved 14 January 2021.
  16. ^ Campbell, Dana (11 November 2014). "Vespa mandarinia". Encyclopedia of Life. Archived from the original on 7 October 2013. Retrieved 16 September 2014.
  17. ^ a b c Handwerk, Brian (25 October 2002). ""Hornets From Hell" Offer Real-Life Fright". National Geographic News. Archived from the original on 25 January 2010.
  18. ^ Archer, Michael E. (1993). "A phylogenetic study of the species of the genus Vespa (Hymenoptera: Vespinae)". Insect Systematics & Evolution. 24 (4): 469–478. doi:10.1163/187631293x00226.
  19. ^ Vecht, Jacobus van der (21 May 1959). "Notes on oriental Vespinae, including some species from China and Japan (Hymenoptera, Vespidae)". Zoologische Mededelingen. 36 (13). Naturalis: 205–232.
  20. ^ "Vespa mandarinia Smith 1852 (Asian giant hornet)". PBDB.org.
  21. ^ Archer, M.E. (2012). Penney, D. (ed.). Vespine wasps of the world: behaviour, ecology and taxonomy of the Vespinae. Monograph Series. Vol. 4. Siri Scientific. ISBN 9780956779571. OCLC 827754341.
  22. ^ Carpenter, James M. & Kojima, Jun-ichi (1997). "Checklist of the species in the subfamily Vespinae (Insecta: Hymenoptera: Vespidae)" (PDF). Natural History Bulletin of Ibaraki University. 1: 51–92.
  23. ^ a b Baker, Mike (2 May 2020). "'Murder Hornets' in the U.S.: The Rush to Stop the Asian Giant Hornet". The New York Times. Retrieved 5 May 2020.
  24. ^ "Murder hornet". Collins English Dictionary. Retrieved 1 November 2020.
  25. ^ Garvey, Kathy Keatley (4 May 2020). "About Those Asian Giant Hornets..." UC Davis: San Mateo & San Francisco Counties. University of California, Davis. Archived from the original on 7 September 2021. Retrieved 22 May 2021. "It's ridiculous to call them murder hornets," says noted UC Davis wasp expert and researcher Lynn Kimsey, director of the Bohart Museum of Entomology and professor of entomology, UC Davis Department of Entomology and Nematology.
  26. ^ "'Murder hornets' get a new name; now called northern giant hornets". FOX 13 Seattle. 25 July 2022. Retrieved 25 July 2022.
  27. ^ Schlosser, Kurt (25 July 2022). "Murder hornets get a new name, but desire to wipe them out in Washington state remains the same". GeekWire. Retrieved 25 July 2022.
  28. ^ "'Murder Hornet' Rebranding Makes A Clean Break From Crime". news.yahoo.com. 27 July 2022. Retrieved 27 July 2022.
  29. ^ a b c d e f Barth, Zach; Kearns, Thomas; Wason, Elizabeth. "Vespa mandarinia". Animal Diversity Web. University of Michigan Museum of Zoology. Archived from the original on 8 October 2015. Retrieved 25 September 2014.
  30. ^ a b Kuroda, Reiko; Kameda, Tsunenori (31 January 2018). "Conformation change of hornet silk proteins in the solid phase in response to external stimulation". Chirality. 30 (5). Wiley: 541–547. doi:10.1002/chir.22824. ISSN 0899-0042. PMID 29384590.
  31. ^ a b
  32. ^ a b Michael J Skvarla, Matthew A Bertone, Patrick J Liesch (2022) Murder Hornet Mayhem: The Impact of the 2020 Giant Hornet Panic and COVID-19 Pandemic on Arthropod Identification Laboratories. American Entomologist 68(2): 38–43, https://doi.org/10.1093/ae/tmac029
  33. ^ AGH and look-alikes; WSDA
  34. ^ Osterloff, Emily (12 March 2020). "Why Asian hornets are bad news for British bees". Natural History Museum, London. Retrieved 11 November 2020.
  35. ^ a b c d e f g Azmy, Muna Maryam; Hosaka, Tetsuro; Numata, Shinya (2016). "Responses of four hornet species to levels of urban greenness in Nagoya city, Japan: Implications for ecosystem disservices of urban green spaces". Urban Forestry & Urban Greening. 18. Elsevier BV: 117–125. Bibcode:2016UFUG...18..117A. doi:10.1016/j.ufug.2016.05.014. ISSN 1618-8667. S2CID 89290615.
  36. ^ a b c d e f g h i j k l m n o p q r Matsuura, Makoto; Sakagami, Shôichi F. (1973). "A Bionomic Sketch of the Giant Hornet, Vespa mandarinia, a Serious Pest for Japanese Apiculture (With 12 Text-figures and 5 Tables)". 北海道大學理學部紀要 [Journal of the Faculty of Science Hokkaido University Series VI. Zoology]. 19 (1). 北海道大學 (Hokkaido University): 125–162. hdl:2115/27557. S2CID 55398608.
  37. ^ a b c "Asian Giant Hornets". Penn State Extension. 6 May 2020. Retrieved 19 November 2020.
  38. ^ "broadcast listing". TV Asahi. 28 September 2008. Archived from the original on 16 October 2022. Retrieved 5 May 2020.
  39. ^ "British Columbia Ministry of Agriculture Pest Alert: Asian Giant Hornet" (PDF). September 2019.
  40. ^ Bérubé, Conrad (February 2020). "Giant Alien Insect Invasion Averted – Canadian Beekeepers Thwart Apicultural Disaster" (PDF). American Bee Journal: 209–214.
  41. ^ a b c d e "Confirmed Asian giant hornet detections". Washington State Department of Agriculture.
  42. ^ "First male 'murder hornet' trapped in Wash. state". KOMO News. 17 August 2020. Retrieved 2 November 2020.
  43. ^ Relyea, Kie (30 September 2020). "Three more Asian giant hornets found in Whatcom. Why the warm weather could mean more". Bellingham Herald. Retrieved 2 November 2020..
  44. ^ "Detection and Eradication Data". Washington State Department of Agriculture. Retrieved 2 November 2020.
  45. ^ "Washington State Officials Hunt for Colony of 'Murder Hornets'". The New York Times. 3 October 2020.
  46. ^ a b c d e "Washington State Department of Agriculture Press Conference". TVW. 10 November 2020. Retrieved 11 November 2020.
  47. ^ Geranios, Nicholas (23 October 2020). "Washington state discovers first 'murder hornet' nest in US". AP News. Retrieved 23 October 2020.
  48. ^ "Washington state crews destroy first US murder hornet nest". The Guardian. Associated Press. 24 October 2020. Retrieved 25 October 2020.
  49. ^ Geranios, Nicholas (26 October 2020). "Scientists remove 98 'murder hornets' in Washington state". AP News. Retrieved 26 October 2020.
  50. ^ a b Salp, Karla (30 October 2020). "Stirring up a hornet nest – safely". Washington State Department of Agriculture. Retrieved 3 November 2020.
  51. ^ a b Main, Douglas (2022). "The untold, dramatic story behind the discovery of America's first murder hornet nest". National Geographic. National Geographic Society. Archived from the original on 15 April 2022.
  52. ^ a b "'Murder hornet' nest with nearly 200 queens destroyed 'in the nick of time'". CBS News. 11 November 2020. Retrieved 11 November 2020.
  53. ^ Eisenstadt, Abigail (17 November 2020). "Family Members Follow Original Asian Giant Hornet to Smithsonian". Smithsonian Magazine. Smithsonian Institution. Retrieved 18 November 2020.
  54. ^ "Frozen 'murder hornets' shipped across the country for further study". KATU. 17 November 2020. Retrieved 18 November 2020.
  55. ^ "'Murder hornets': More nests likely to be found in US". BBC News. 11 November 2020. Retrieved 12 November 2020.
  56. ^ "B.C. beekeepers and residents asked to keep lookout after discovery of another Asian giant hornet". CBC. 3 November 2020. Retrieved 4 November 2020.
  57. ^ Mooney, Harrison (3 November 2020). "Residents alerted after 'murder hornet' found in Abbotsford". Vancouver Sun. Retrieved 4 November 2020.
  58. ^ Azpiri, Jon (3 November 2020). "'Murder hornet' spotted in Abbotsford, B.C." Global News. Retrieved 4 November 2020.
  59. ^ ""Murder hornet" discovered in Abbotsford neighbourhood". Daily Hive Vancouver. 3 November 2020. Retrieved 4 November 2020.
  60. ^ "Asian giant hornet found in Abbotsford". Province of British Columbia News Archive. 3 November 2020. Retrieved 4 November 2020.
  61. ^ Georgiou, Aristos (9 November 2020). "Murder hornets discovered 100 miles from Seattle as invasion of North America continues". Newsweek. Retrieved 9 November 2020.
  62. ^ jonazpiri (10 November 2020). "Another 'murder hornet' spotted in B.C. – BC". Global News. Retrieved 13 November 2020.
  63. ^ Daflos, Penny (12 November 2020). "Residents find queen 'murder hornet' in B.C.'s Fraser Valley as ministry traps sit empty". CTV News. Retrieved 13 November 2020.
  64. ^ Mangione, Kendra (10 November 2020). "2 'murder hornets' found about 5 kilometres apart within 5 days in B.C." CTV News. Retrieved 13 November 2020.
  65. ^ "Another 'murder hornet' discovered in B.C.'s Fraser Valley". CTV News. 3 November 2020. Retrieved 13 November 2020.
  66. ^ Claxton, Matthew (28 May 2020). "Asian giant "murder hornets" found in Brookswood". Aldergrove Star. Retrieved 13 November 2020.
  67. ^ "Two more 'murder hornets' turn up on B.C. mainland". AgCanada. 19 August 2023.
  68. ^ "STATE ENTOMOLOGISTS CONFIRM THE FIRST LIVE ASIAN GIANT HORNET SIGHTING OF 2021". Washington State Department of Agriculture. 8 September 2020. Retrieved 13 August 2021.
  69. ^ "State eradicates first Asian giant hornet nest of 2021, asks public to continue reporting".
  70. ^ "3rd Asian giant hornet nest of 2021 found, 2nd nest eradicated in Whatcom County". 10 September 2021.
  71. ^ "Suspected Asian giant hornet spotted farther east than other sightings". 17 September 2021.
  72. ^ Holpuch, Amanda (2 May 2020). "'Murder hornets' in Washington state threaten bees and whip up media swarm; Asian giant hornet, which became more active in the state in April, is the world's largest and can kill humans with multiple stings". The Guardian.
  73. ^ "ARS Takes On the Asian Giant Hornet". USDA ARS AgResearch Magazine. Retrieved 18 November 2020.
  74. ^ Grijalva, Raul M. (8 May 2020). "Murder Hornet Eradication Act of 2020" (PDF).
  75. ^ "Rules Committee Print 116–68 Text of the House Amendment to the Senate Amendment to H.R. 133" (PDF). p. 1606.
  76. ^ Burton, Bonnie (21 December 2020). "US Congress to establish 'murder hornet' eradication pilot program". CNET. Retrieved 22 December 2020.
  77. ^ a b Cecco, Leyland (1 January 2021). "Beekeepers brace for next round with Canada's 'murder hornets'". The Guardian. Retrieved 1 January 2021.
  78. ^ "State and Federal Entomologists Confirm New Asian Giant Hornet Detection in Snohomish County". Washington State Department of Agriculture. 16 June 2021. Retrieved 16 June 2021.
  79. ^ "ANNUAL INVASIVE PEST HUNT CONCLUDES - Second year without northern giant hornet detections"
  80. ^ a b c d Akre, R D; Davis, H G (1978). "Biology and Pest Status of Venomous Wasps". Annual Review of Entomology. 23 (1). Annual Reviews: 215–238. doi:10.1146/annurev.en.23.010178.001243. ISSN 0066-4170. PMID 343706.
  81. ^ a b c
  82. ^ Hirano, Koji; Tanikawa, Atsuhiro (May–August 2020). "Ocular injury caused by the sprayed venom of the Asian Giant Hornet (Vespa mandarinia)". Case Reports in Ophthalmology. 11 (2): 430–435. doi:10.1159/000508911. PMC 7506230. PMID 32999672. S2CID 221882297.
  83. ^ Schmidt, Justin O.; Yamane, Soichi; Matsuura, Makoto; Starr, Christopher K. (1986). "Hornet venoms: Lethalities and lethal capacities" (PDF). Toxicon. 24 (9). Elsevier: 950–954. Bibcode:1986Txcn...24..950S. doi:10.1016/0041-0101(86)90096-6. PMID 3810666.
  84. ^ Pongracic, Jacqueline A. "Vespid allergy and Asian giant hornet". American Academy of Allergy Asthma & Immunology.
  85. ^ Makino, Shun'ichi; Yamashita, Yoshiharu (25 December 1998). "Levels of Parasitism by Xenos moutoni du Buysson (Strepsiptera, Stylopidae) and their Seasonal Changes in Hornets (Hymenoptera: Vespidae, Vespa) Caught with Bait Traps" (PDF). Entomological Science. 1 (4): 537–543. ISSN 1479-8298. NAID 110003374544. Archived (PDF) from the original on 20 March 2019. Retrieved 25 September 2014.
  86. ^ "Vespa mandarinia". University of Wisconsin–La Crosse (UWL). Archived from the original on 10 January 2019. Retrieved 25 September 2014.
  87. ^ a b Taylor, Benjamin J.; Nordheim, Erik V.; Schueller, Teresa I.; Jeanne, Robert L. (2011). "Recruitment in swarm-founding wasps: Polybia occidentalis does not actively scent-mark carbohydrate food sources". Psyche. 2011: 1–7. doi:10.1155/2011/378576.
  88. ^ a b Yoshimoto, J.; Nishida, T. (1 March 2009). "Factors Affecting Behavioral Interactions Among Sap-Attracted Insects". Annals of the Entomological Society of America. 102 (2): 201–209. doi:10.1603/008.102.0203.
  89. ^ Richter, M. Raveret (2000). "Social Wasp (Hymenoptera: Vespidae) Foraging Behavior". Annual Review of Entomology. 45 (1). Annual Reviews: 121–150. doi:10.1146/annurev.ento.45.1.121. ISSN 0066-4170. PMID 10761573. p. 126: Social wasps use masticated arthropod prey and other animal protein to progressively provision their developing brood. Prey items most commonly include a variety of arthropods such as ... bees (...79...)...
  90. ^ a b c d "Adw: : Information". Animal Diversity Web.
  91. ^ Kosmeier, Dieter (27 January 2013). "Vespa mandarinia (Asian Giant Hornet) page". Vespa-crabro.de. Retrieved 18 March 2013.
  92. ^ Kosmeier, Dieter (27 January 2013). "Vespa mandarinia (Asian Giant Hornet) page". Vespa-crabro.de. Retrieved 18 March 2013.
  93. ^ Hunt, James H.; Baker, Irene; Baker, Herbert G. (November 1982). "Similarity of amino acids in nectar and larval saliva: the nutritional basis for trophallaxis in social wasps". Evolution. 36 (6): 1318–1322. doi:10.2307/2408164. JSTOR 2408164. PMID 28563573.
  94. ^ Hamblin, Steven R.; White, Peter A.; Tanaka, Mark M. (2014). "Viral niche construction alters hosts and ecosystems at multiple scales". Trends in Ecology & Evolution. 29 (11). Cell Press: 594–599. Bibcode:2014TEcoE..29..594H. doi:10.1016/j.tree.2014.08.005. ISSN 0169-5347. PMID 25237032.
  95. ^ a b c d e
  96. ^ "Defensive Adaptations: Heat Tolerance As A Weapon". Bio.davidson.edu. 2001. Archived from the original on 4 November 2013. Retrieved 18 March 2013.
  97. ^ Tan, Ken; Wang, Zhenwei; Li, Hua; Yang, Shuang; Hu, Zongwen; Kastberger, Gerald; Oldroyd, Benjamin P. (2012). "An 'I see you' prey–predator signal between the Asian honeybee, Apis cerana, and the hornet, Vespa velutina". Animal Behaviour. 83 (4): 879–882. doi:10.1016/j.anbehav.2011.12.031. S2CID 53192582.
  98. ^ Salp, Karla (3 August 2021). "Hungry hornet babies – what's on their menu in the PNW?". Washington State Department of Agriculture AgBriefs. Retrieved 13 August 2021.
  99. ^ Wason, Zach Barth; Thomas Kearns; Elizabeth. "ADW: : INFORMATION". Animal Diversity Web. Retrieved 4 April 2024.{{cite web}}: CS1 maint: multiple names: authors list (link)
  100. ^ "スズメバチを食べるカマキリ - ニホンミツバチの四季". goo blog (in Japanese). Retrieved 7 April 2024.
  101. ^ Suetsugu, K. (6 September 2018). Dafni, A. (ed.). "Social wasps, crickets and cockroaches contribute to pollination of the holoparasitic plant Mitrastemon yamamotoi (Mitrastemonaceae) in southern Japan". Plant Biology. 21 (1). Wiley: 176–182. doi:10.1111/plb.12889. hdl:20.500.14094/90005488. ISSN 1435-8603. PMID 30098096.
  102. ^
  103. ^ Molteni, Megan (24 August 2020). "Inside the Sprint to Map the Murder Hornet Genome". Wired. Retrieved 2 November 2020.
  104. ^ Golembiewski, Kate (27 December 2020). "Murder Hornets Menaced Parts of the U.S. This Year. How Big of a Threat Are They?". Discover Magazine. Retrieved 7 January 2021.
  105. ^ "Giant hornet".
  106. ^ Dooley, Ben (5 May 2020). "In Japan, the 'Murder Hornet' Is Both a Lethal Threat and a Tasty Treat". The New York Times. Retrieved 10 September 2020.
  1. ^ a b c d e p. 51–52, "The queens usually select underground cavities as nest sites. The cavities are either associated with rotten tree roots or are made by small vertebrates such as moles and snakes. The cavities are in well drained soil along a slope or under an overhanging cliff. The nests are found at a depth of six to 60 cm and the entrance tunnel is two to 60 cm long. A few nests are found above ground (e.g. nine nests from a sample of 56) either in tree hollows or mud walls and within one or two meties above the surface of the ground. The nests are found on hillsides, parks and forests but are rare in the lowlands and high mountains. The Taiwan colour form of V. mandarinia also nests underground but the western colour form has been found in tree hollows near the surface of the ground (Bingham, 1888)"
  2. ^ p. 48, "V. soror du Buysson 1905 was described as a variety of V. ducalis Smith, 1852, despite the structural characteristics of the vertex and apical margin of the clypeus being similar to V. magnifica. Van der Vecht (1957) recognised the confusion and proposed the new combination V. mandarinia soror. Archer (1991a) showed that V. mandarinia soror was sympatric for part of its geographical distribution with V. m. mandarinia but still retained its distinctive colour characteristics so should be given specific status, V. soror."
  3. ^ p. 48–49, "V. mandarinia and V. soror cannot be satisfactory separated by structural characteristics but are readily separated by colour characteristics:
    1. Third to the sixth gastral terga in the female and to the seventh gastral terga in the male black, at most with a narrow apical orange band on the third gastral tergum .................................................................... soror du Buysson. 1905
    — Third to the fifth gastral terga in the female and to the sixth gastral terga in the male with either a narrow or broad apical orange band, tergum six in the female and tergum seven in the male largely orange ................................... mandarinia Smith. 1852"
  4. ^ a b p. 52, "The queen builds a comb of about 44 cells (range 37–60) with a mean cell building rate per day of 1.64 (range 0–4) and a mean egg laying rate per day of 1.53 (range 0–5). The envelope is bowl-shaped, not completely enclosing the comb which is ventrally exposed within the nest cavity. The queen is able to excavate soil so as to enlarge the nest cavity as the nest grows in size. The first workers emerge as adults after about 38 days from the queen nest. In southern Japan, nests at maturity consist of four to seven combs although five to six combs are more usual. Mature nests contain about 2700 cells with the largest nest having 4661 cells. The large cells are clearly larger than the small cells although the size of the small cells does increase during the development of the nest. The envelope is thin and absent at the bottom of the nest exposing the lower comb and providing access to the combs. The workers continue to excavate soil to enlarge the nest cavity although stones too large to be carried drop to the bottom of the nest cavity. The ability of the queen and workers to excavate soil probably relates to the lack of relocation behaviour in this species."
  1. ^ p. 1693, "A hornet (Vespa mandarinia) neurotoxin, mandaratoxin (MDTX)"
  2. ^ p. 1696, "Estimations of the molecular weight of purified MDTX in its reduced and unreduced forms with denaturing solvents and its molecular weight in the native form are nearly the same. It is concluded that the toxin is a single polypeptide chain of approximately 20,000 daltons. Thus, the toxin acts on nerve membranes as a monomer protein of similar molecular weight."
  1. ^  • p. 6, "Our results show that the east coast is highly suitable for the establishment and spread of V. mandarinia, a zone where this species has not been recorded yet, making it necessary to implement preventive actions to avoid a possible invasion."
  2. ^  • p. 2, "The abundance of V. mandarinia is positively associated with amounts of green spaces in urban landscapes, suggesting that the control of their populations should be focused on urban green areas.26"
     • p. 6, "For canopy cover, a similar result was found by Azmy et al.26 in urban environments in China, where the quality of green areas benefited the abundance of V. mandarinia."
  3. ^  • p. 4-6: Figure 2B, Figure 3A, Figure 3C, Figure 4
     • Supplemental #2: Table S5
  4. ^  • p. 7, "Here we have estimated an annual projected loss of US$113.7 million per year. However, the costs associated with control actions not included in our study may eventually increase these amounts significantly (i.e. c. US$26 million as estimated by Barbet-Massin et al.12 for V. velutina)."
  5. ^  • p. 1, "If this species spread across the country, it could threaten 95 216 ± 5551 honey bee colonies, threatening an estimated income of US$ ... 101.8 million for ... bee-pollinated crops production ... while colonizing 60 837.8 km2 of bee-pollinated croplands."
     • p. 5, "The potential threatened income associated with bee-pollinated croplands reached US$101.8 million per year (Tables S7 and S8)."
  6. ^ a b c  • p. 4-7: Figure 2, §3.2 Threatened colonies, §3.3 Hive products potential losses, §3.4 Threatened bee-pollinated croplands, Figure 4, Figure 5
     • Supplemental #2: Table S5, Table S6 (mislabeled S5), Table S7, Table S8, Table S9
  7. ^  • p. 5, "We also identified that alfalfa/hay, apples, grapes and tobacco are the crops with the largest threatened areas of 58 484.1, 522.9, 468.5 and 432.9 km2, respectively (Table S8)."
     • Supplemental #2: Table S3, Table S8, Table S9
  • Secondary support of Alaniz:
  1. ^ p. 2, "Complete mitochondrial DNA was sequenced with Illumina’s MiSeq platform (ILLUMINA, United States)."
  2. ^ p. 3, "These genetic differences corresponded to the values found between native V. mandarinia from Japan, South Korea, and China."
  3. ^ p. 3-4, "A high pairwise distance of 0.0071 was also confirmed between the 13 PCGs of mitochondrial DNA sequences of V. mandarinia specimens from the United States and Canada, suggesting that the hornets differ in maternal origin (Table 1). ... The ML tree also revealed that V. mandarinia from the United States and Canada were not monophyletic (Fig. 2). Molecular phylogenetic analysis of the mitochondrial genomes revealed that V. mandarinia from the United States was genetically distant from that of Canada. ... The observed genetic differences between the Canada and U.S. mitochondrial genomes suggest that the two V. mandarinia specimens introduced to western N. America during or prior to 2019 are derived from different maternal lineages."
  4. ^ p. 4, "The mitochondrial genome of ... the Canadian V. mandarinia was most genetically similar to the Japanese V. mandarinia used in this study."
  5. ^ p. 4, "The mitochondrial genome of the specimen collected from Blaine, WA shared 99.5% sequence homology to the specimen characterized from South Korea,"

Further information