Asian giant hornet
The Asian giant hornet ("Vespa mandarinia"), including the color form referred to as the Japanese giant hornet, 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 with a few more additional sightings in 2020, and nests found in 2021, prompting concern that it could become an invasive species. 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 rotted pine roots. It feeds primarily on larger insects, colonies of other eusocial insects, tree sap, and honey from honey bee colonies. The hornet has a body length of , a wingspan around , and a stinger long, which injects a large amount of potent venom. Although the scientific literature and official government sources continue to refer to this species by its established common name, the mainstream media have taken to using the nickname "murder hornet". The species has been around  since the Miocene as indicated by fossils found in Shanwang formation. Taxonomy and phylogeny. "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". 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. Division of the genus into subgenera has been attempted in the past, but has been abandoned, due to the anatomical similarity among species and because behavioral similarity is not associated with phylogeny. As of 2012, three subspecies were recognized: "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. 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. Description. 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. The thorax is dark brown, with two grey wings varying in span from . 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 long and delivers a potent venom that in cases of multiple hornets stinging simultaneously can kill a human. Queens and workers. The queens are considerably larger than workers. Queens can exceed , while workers are between . The reproductive anatomy is consistent between the two, but workers do not reproduce. Drones. Drones (males) are similar to females, and can attain in length, but lack stingers. This is a consistent feature among the Hymenoptera. Larvae. Larvae spin  a silk cocoon when they complete development and are ready to pupate. 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. Misidentifications. 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, "V. velutina" in Europe, and the European hornet ("V. crabro") in the eastern United States. The eastern cicada killer ("Sphecius speciosus") also lives in the Eastern United States . It is a large digger wasp that can reach in length, and has also been mistaken for the Asian giant hornet. Distribution. Ecological distribution. "V. mandarinia" is primarily a forest dweller. When it does live in urban landscapes, however, "V. mandarinia" is highly associated with green space. It is the most dependent upon green space of the "Vespa" species (with "V. analis" the least). Extremely urbanized areas thus provide a refuge for "V. analis", whereas "V. mandarinia" – its predator – is entirely absent. Geographic distribution. Asia. 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 have mainly been concentrated in the Vancouver area, with nests also discovered in neighboring Whatcom County, Washington, in the United States. A mitochondrial DNA analysis was performed to determine the maternal population(s) ancestral to the British Columbia and Washington introduced populations. The high dissimilarity between these two was similar to the mutual distances between each of the Chinese, Japanese, and Korean native populations suggesting the specimens collected in 2019 were from two different maternal populations, Japanese in BC and South Korean in Washington. This suggests that two simultaneous  introductions of the Asian giant hornet occurred in North America within about of one another within a few months. In April 2020, authorities in Washington 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. 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. Two assessment models of their  potential to spread from their present location on the US–Canada border suggested that they could spread northward into coastal British Columbia and Southeast Alaska, and southward as far as southern Oregon. 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. In 2020, the United States Congress considered specific legislation to eradicate "V. mandarinia" 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. British Columbia Agriculture is prepared for a "long fight" lasting years, if necessary. 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. On June 4, 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, but DNA testing was needed. USDA APHIS (Animal and Plant Health Inspection Service) did the  analysis and a few days later 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. Nesting. "V. mandarinia" nests in low mountain foothills and lowland forests. 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. Unlike other species of "Vespa", "V. mandarinia" almost exclusively inhabits subterranean nests in 1978 it was still doubted that aerial nests were possible, as Matsuura and Sakagami reported this to be unknown in Japan in 1973 and aerial nesting is still described as extremely rare in Japan, and yet all nests in the invasive range have been aerial. In a study of 31 nests, 25 were found around rotten pine roots, and another study found only 9 of 56 nests above ground. Additionally, rodents, snakes, or other burrowing animals  previously made some of the tunnels. The depth of these nests was between . The entrance at the ground surface varies in length from either horizontally, inclined, or vertically. The queens that found the nest prefer narrow cavities. Nests of "V. mandarinia" typically lack a developed envelope. During the initial stages of development, the envelope is in an inverted-bowl shape. 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. A system of one main pillar and secondary pillars connects the combs. Nests  usually have four to seven combs. 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 with 1,192 cells (no obstacles, circular) and (elliptical; wrapped around a root system). Colony cycle. The nesting cycle of "V. mandarinia" is fairly consistent with that of other eusocial insects. Six phases occur in each cycle. 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. 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. 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, 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. Sting. The stinger of the Asian giant hornet is about long, which is about longer than that of a honeybee. Venom. That stinger injects an especially potent venom that contains, like many bee and wasp venoms, a cytolytic peptide (specifically, a mastoparan) that can damage tissue by  stimulating phospholipase action, in addition to its own phospholipase. Masato Ono, an entomologist at Tamagawa University, described the sensation of being stung as feeling "like a hot nail being driven into my leg". 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. The venom contains a neurotoxin called mandaratoxin, a single-chain polypeptide with a molecular weight around 20 kDa. 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 of 4.0mg/kg. (In comparison, the deadliest wasp venom (at least to laboratory mice) by weight belongs to "V. luctuosa" at 1.6mg/kg.) The potency of the "V. mandarinia" sting is due, rather, to the relatively large amount of venom injected. Immunogenicity. 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. 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. Since 2001, the yearly human death toll caused by stings of bees, wasps, and hornets in Japan has been ranging between 12 and 26. Since this number also includes deaths caused by bees, wasps, and other  hornet species, the number of deaths caused by Asian giant hornets is likely to be much lower. Advice in China is that people stung more than 10 times should seek medical help, and require emergency treatment for more than 30 stings. The stings can cause kidney failure. In 2013, stings by Asian giant hornets killed 41 people and injured more than 1,600 people in Shaanxi, China. One source claims that most people who sought medical attention for a sting entered anaphylaxis before or during the medical session, and that deaths from envenomation are primarily related to anaphylactic shock or cardiac  arrest. Either way, certainly some venom fractions are immunogenic and some are cardiotoxic to mammals. Deaths have occurred as a result of multiple organ failure, typically after a large number of stings. Sting victims who experience life-threatening organ failure commonly exhibit signs of skin hemorrhaging and necrosis, though in patients without organ failure, these symptoms are very rare. The two likely reasons for skin hemorrhaging and necrosis are an inability to effectively neutralize the venom, or unusually potent venom toxicity for that set of stings. In either case, for a small number of victims, these stings lead to multiple  organ injury. While not all such victims displayed lesions or necrosis, a strong correlation existed between the number of stings and the severity of injury. Those who died, on average, were stung 59 times (with a standard error of 12), while those who survived suffered only, on average, 28 stings (with a standard error of four, and a ). Parasites. 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. 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. 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. 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. 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). 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. Diet. The Asian giant hornet is intensely predatory; it hunts medium- to large-sized insects, such as bees, other hornet species, 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. This hornet often attacks both colonies of other "Vespa" species ("V. simillima" being the usual prey species) and honey bee hives to obtain the adults, pupae, and larvae as food for their own larvae; sometimes, they also 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. 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 in a single day, at speeds up to . 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. 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. Native honey bees. 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, Kakugo virus infection, though, may provide an extrinsic defence. 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. 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. This raises the temperature in the ball to the critical  temperature of . In addition, the exertions of the honey bees raise the level of carbon dioxide (CO) in the ball. At that concentration of CO, they can tolerate up to , but the hornet cannot survive the combination of high temperature and high carbon dioxide level. 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. 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. Another defence used by "Apis cerana" is speeding up dramatically when returning to the colony, to avoid midair attacks. Diet in North America. The WSDA 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 be beef from a hamburger. Pollination. "V. mandarinia" is among the diurnal pollinators of the obligate plant parasite, the plant "Mitrastemon yamamotoi". Extermination methods. As of 1973, six different methods were used to control hornets in Japan; these methods decrease damage done by "V. mandarinia". Beating. 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. 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 . V. mandarinia" travels up to away from the nest. For the rare nest that is up in a tree, wrapping the tree in plastic and vacuuming the hornets out is used. 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. 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. 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. 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. 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. Human consumption. In some Japanese mountain villages, the nests are excavated and the larvae are considered a delicacy when fried. 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. Economic impact. If "V. mandarinia" reaches all suitable habitat in North America, control costs in the United States will be over US$113.7 million/year (possibly significantly higher). 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. New York, Massachusetts, Pennsylvania, Connecticut, North Carolina, New Jersey, and Virginia would be most severely affected. By region, New England would be worst hit, and to a lesser degree the entire northeast and the entirety of eastern North America. 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. Alfalfa/other hays, apples, grapes, tobacco, cotton, and blueberries would be the crops most severely  affected.