Vespa velutina
(Asian hornet)

Pictures

Picture	Title	Caption	Copyright
Title Wild habit Caption Vespa velutina (Asian hornet); habit on flowers (Apiacea). nr. Amarante, Portugal. September 2016. Copyright ©Charles J. Sharp/via wikipedia - CC BY-SA 4.0	Wild habit	Vespa velutina (Asian hornet); habit on flowers (Apiacea). nr. Amarante, Portugal. September 2016.	©Charles J. Sharp/via wikipedia - CC BY-SA 4.0
Title Adult Caption Vespa velutina (Asian hornet); male at rest. Copyright ©Gilles San Martin/via flickr - CC BY-SA 2.0	Adult	Vespa velutina (Asian hornet); male at rest.	©Gilles San Martin/via flickr - CC BY-SA 2.0
Title Adult Caption Vespa velutina (Asian hornet); male at rest, showing facial detail. Copyright ©Gilles San Martin/via flickr - CC BY-SA 2.0	Adult	Vespa velutina (Asian hornet); male at rest, showing facial detail.	©Gilles San Martin/via flickr - CC BY-SA 2.0
Title Adult Caption Vespa velutina (Asian hornet); subspecies nigrithorax. Habit, drinking from a pond. Century Park, Shanghai, China. September 2017. Copyright ©Nature.Catcher/via flickr - CC BY-SA 4.0	Adult	Vespa velutina (Asian hornet); subspecies nigrithorax. Habit, drinking from a pond. Century Park, Shanghai, China. September 2017.	©Nature.Catcher/via flickr - CC BY-SA 4.0
Title Adult Caption Vespa velutina (Asian hornet); dorsal view. Museum set specimen, originally from Jardin des Plantes, Toulouse, France. June 2013. Copyright ©Muséum de Toulouse/Didier Descouens-2013 - CC BY-SA 4.0	Adult	Vespa velutina (Asian hornet); dorsal view. Museum set specimen, originally from Jardin des Plantes, Toulouse, France. June 2013.	©Muséum de Toulouse/Didier Descouens-2013 - CC BY-SA 4.0
Title Adult Caption Vespa velutina (Asian hornet); ventral view. Museum set specimen, originally from Jardin des Plantes, Toulouse, France. June 2013. Copyright ©Muséum de Toulouse/Didier Descouens-2013 - CC BY-SA 4.0	Adult	Vespa velutina (Asian hornet); ventral view. Museum set specimen, originally from Jardin des Plantes, Toulouse, France. June 2013.	©Muséum de Toulouse/Didier Descouens-2013 - CC BY-SA 4.0
Title Natural enemy Caption Vespa velutina (Asian hornet); natural enemy. A montage image (not to scale) of a parasitic Mermithid nematode (probably Pheromermis vasparum) and its host, Vespa velutina. (From the paper: (2015). "Can parasites halt the invader? Mermithid nematodes parasitizing the yellow-legged Asian hornet in France". PeerJ 3: e947. DOI:10.7717/peerj.947. ISSN 2167-8359) Copyright ©Quentin Rome/via wikipedia - CC BY-SA 4.0	Natural enemy	Vespa velutina (Asian hornet); natural enemy. A montage image (not to scale) of a parasitic Mermithid nematode (probably Pheromermis vasparum) and its host, Vespa velutina. (From the paper: (2015). "Can parasites halt the invader? Mermithid nematodes parasitizing the yellow-legged Asian hornet in France". PeerJ 3: e947. DOI:10.7717/peerj.947. ISSN 2167-8359)	©Quentin Rome/via wikipedia - CC BY-SA 4.0

Identity

Preferred Scientific Name

• Vespa velutina Lepeletier, 1836

Preferred Common Name

• Asian hornet

Other Scientific Names

• Vespa auraria Smith, 1852
• Vespa crabro var. immaculata Morawitz, 1889
• Vespa flavitarsa Sonan, 1929
• Vespa fruhstorferi Stadelmann, 1894
• Vespa mongolica var. divergens Pérez, 1910

International Common Names

• English: Asian black hornet; Asian hornet; yellow-legged hornet
• Spanish: avispa asiática
• French: frelon à pattes jaunes; frelon asiatique

Local Common Names

• Germany: Asiatische Hornisse
• Italy: calabrone asiatico
• Portugal: avispón asiático

Summary of Invasiveness

Vespa velutina (Hymenoptera: Vespidae) is a hornet of Asian origin which is a generalist predator of medium- to large-sized insects, and scavenger of vertebrate carrion. It has large impacts on Diptera and social hymenopterans, and in particular on honey bees (Apis spp.). It has recently been spreading in Asia (it is an invasive species in South Korea and Japan), and the subspecies V. v. nigrithorax has been accidentally introduced to Europe where it was first recorded from southern France in 2005. Since then it has been found in Spain, Portugal, Belgium, Italy, the UK, the Netherlands, Germany, the Channel Islands and the Balearic Islands. This invasive species threatens honey production and native pollinating insects. It may be introduced and transported accidentally with soil associated with plants, garden furniture and pots, timber, vegetables, camping equipment, etc.

Taxonomic Tree

• Domain: Eukaryota
•     Kingdom: Metazoa
•         Phylum: Arthropoda
•             Subphylum: Uniramia
•                 Class: Insecta
•                     Order: Hymenoptera
•                         Family: Vespidae
•                             Genus: Vespa
•                                 Species: Vespa velutina

Notes on Taxonomy and Nomenclature

Vespa velutina is one of the 22 currently recognized Vespa species (Archer, 2012; Perrard et al. 2013). It includes 12 colour forms (among which are auraria Smith, 1852, nigrithorax du Buysson, 1905 and pruthii Sonan, 1929). These forms were long considered as subspecies (Vecht, 1957) but came to be treated as synonyms of the nominal form according to Carpenter and Kojima (1997). In 1991, Archer removed auraria and pruthii to establish a separate species V. auraria, because of its apparent sympatry with V. velutina nigrithorax in different localities, but this was rejected by Nguyen et al. (2006) because the colour form auraria intergraded with nigrithorax in northern Vietnam and no other morphological character could be found to distinguish them. Further molecular studies confirmed the synonymy of auraria and pruthii with V. velutina (Perrard 2012). Archer (2012) provided a key supporting the identification of V. velutina colour forms and their distribution. It is nigrithorax that is invasive in Europe.

Description

V. velutina is highly variable in colour and 10 subspecies have been identified (Vecht 1957; Carpenter and Kojima 1997; Nguyen et al., 2006). The subspecies nigrithorax du Buysson (1905) is invasive in Europe, where it varies in size from 17 to 32 mm (Rome and Villemant, 2018).

C. Villemant (Muséum National d’Histoire Naturelle, Paris, France, personal communication, 2013) describes Vespa velutina nigrithorax as follows:

Female: Head and thorax finely punctate, with fine silky sparsely scattered erect hairs, abdomen chagrined. Body dark brown with face and mouthparts orange (except dark teeth); antennae brown dorsally, orange ventrally. Dorsal face of abdominal segments brown with clearer apical margins: a thin yellow band on the first segment and a thin orange band on the second and third segments; fourth abdominal segment almost entirely orange with a median basal triangular black marking, generally not visible when the hornet is alive; fifth and sixth abdominal segments more or less orange-brown. Ventral part of abdomen brown, the triangular apex often lighter; second and third segments ventrally yellow with a median basal black marking. Legs brown except yellow tarsi; wings brownish hyaline. There are no distinct morphological differences between the sexual and sterile (worker) females. Some workers are smaller (notably in spring) but in autumn many workers are as big as the future queens. Inside the colony, the laying queen can be recognized by her distended abdomen and, at the end of the season, damaged wings. Male: very similar to females in size and colour, antennae longer. Ventral face of abdomen brown, apex truncated with a pair of yellow spots.

A key to the Vespinae can be found in Archer (2012);  Martin (1995) and Nguyen and Carpenter (2002) also provide keys to the Asian species of hornets in the genera Vespa and Provespa.

As for the nests of V. velutina, they are usually larger than those of other European species (a particularly large nest was found in Aquitaine, France that was 1 m tall and 80 cm across -- Rome et al., 2015). Colonies often relocate from a primary nest initiated by the queen to a larger secondary nest (Matsuura and Yamane, 1990). Primary nests have basal entrances, while in secondary nests the entrance is typically located laterally, unlike that of V. crabro which is usually basal (Perrard et al., 2009; F. Muller, Museum National d'Histoire Naturelle, Paris, France, personal communication, 2013).

Distribution

Of the 22 Vespa hornet species from Asia, only a few have extended their geographical range to include the Philippines and New Guinea. Only two species are native to Europe: the European hornet, Vespa crabro Linnaeus (1758) and the oriental hornet Vespa orientalis Linnaeus (1771) (Matsuura and Yamane, 1990). V. crabro is found throughout Europe whereas V. orientalis is restricted to Bulgaria, Greece, southern Italy and eastern North Africa (Carpenter and Kojima, 1997).

V. velutina is widespread in Asia, from north-eastern India throughout southern and central China as far as Taiwan and as far south as Indonesia (Archer 1994) and is recorded in the following countries: Afghanistan, Bhutan, China (including Hong Kong), India, Indonesia (except Kalimantan and Irian Jaya), Japan, Laos, Malaysia, Myanmar, Nepal, Pakistan, South Korea, Taiwan, Thailand and Vietnam (Archer, 2012). Its presence in South Korea results from an introduction in 2003 and it has become invasive there (Choi et al., 2012; Jung et al., 2007). It was reported from the nearby Japanese Tsushima Island in 2012 (Kishi and Goka, 2017) and, in 2015, on Kyushu Island (Minoshima et al., (2015).

In 2005 the first confirmed report came from the southwest of France (Haxaire et al., 2006; Villemant et al., 2006a,b), and the species has now spread throughout almost the whole of France (Rome, 2019); in 2010 it was reported for the first time in Spain (López et al., 2011) and in 2011 in Portugal (Grosso-Silva and Maia, 2012). It has also spread to Belgium (Rome et al., 2013; Q. Rome, Museum National d'Histoire Naturelle, Paris, France, personal communication, 2019), Italy (Federazione Apicoltori Italiani, 2013), the UK (in 2016 -- NNSS undated; with further confirmed sightings in 2018 -- DEFRA, 2018), the Netherlands (in 2017 -- Smit et al., 2018), the Channel Islands (States of Guernsey Government, 2016), the Balearic Islands (Leza et al., 2018) and Germany (Q. Rome, Museum National d'Histoire Naturelle, Paris, France, personal communication, 2019).

Distribution Table

The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

History of Introduction and Spread

The presence in France of V. velutina was first confirmed in publications in 2006, following the discovery in 2005 of several solitary queens and a first colony belonging to the subspecies nigrithorax du Buysson (1905) in the Lot-et-Garonne department (Haxaire et al. 2006; Villemant et al., 2006a,b). This subspecies is found in India and China (Chauzat and Martin, 2009); hibernating founding female(s) might have arrived in Lot-et-Garonne a year earlier in terracotta bonsai pots from China. Subsequent genetic studies showed that the hornets in France originated from eastern China (Jiangsu/Zhejiang) (Arca et al., 2015).

V. velutina has adapted very well to the southwestern French climate and populations soared quickly to the point that eradication was no longer possible. By the end of 2006, it was recorded in eleven departments in the southwest of France, while in 2007, it spread across 20 southwestern departments (Rortais et al., 2010). By 2010 it had been reported from much of western France and a few departments further east (Muséum National d’Histoire Naturelle, 2012). It is now found throughout most of France (Rome, 2019).

In 2010 V. velutina was reported for the first time in Spain, in Guipuzcoa and Navarra provinces in the north of the country near the border with France (López et al., 2011). It was reported from Portugal in 2011 (Rome et al., 2013), Belgium in 2011 (Rome et al., 2013), Italy in 2013 (Federazione Apicoltori Italiani, 2013) and England (UK) in 2016 (Budge et al., 2017) with nine confirmed sightings in 2018 (up to November, DEFRA, 2018). It has also been reported in Germany (Q. Rome, Museum National d'Histoire Naturelle, Paris, France, personal communication, 2019).

In Asia, V. velutina was found in Japan on Tsushima Island in 2012 and on Kyushu Island in 2015. Molecular studies by Takashima et al. (2018) indicate that specimens from each island came from the same origin.

Introductions

Risk of Introduction

In Asia, V. velutina is found in climate ranges close to those found in the south of Europe. It was thought that only unusually cold winters could stop its spread further north and/or reduce the population build-up in the southwestern areas of France (Villemant et al. 2006b), but modelling showed that in fact only dry summers could limit expansion in southern Europe (Villemant et al., 2011). The same study showed that the range could potentially include most of the southern part of Europe and possibly other areas around the world. V. velutina can apparently survive long distance transport (as seen in the French incursion with specimens from China) so its potential for introduction and spread in countries with similar climates to the south-western part of France is extremely high. In France, it spread to cover 120,000 km² within 3 years, demonstrating that it can colonize large areas in a very short period of time if the climatic conditions are favourable. In the UK it was considered to be very likely to enter and become established, and likely to spread rapidly with moderate impacts if and when it does arrive (Marris et al., 2011); first sightings were recorded in 2016 (Budge et al., 2017).

Habitat

V. velutina commonly build two nests in a season (approximately 70% of nests relocated in summer in France -- Rome et al., 2015). Primary nests are often found in or on man-made structures (77% in France -- Franklin et al., 2017), while secondary nests are more likely to be found on natural structures (78% in France -- Franklin et al., 2017), such as in tree canopies well above the ground, or occasionally in undisturbed and sheltered areas in buildings (under stairwells, in abandoned barns, chicken coops or sections of buildings, etc.) or in bramble bushes, and very rarely underground (Martin, 1995; Rome et al., 2009; Franklin et al., 2017). See the Habitat table for the types of habitat in which the species is found.

Habitat List

Biology and Ecology

Reproductive Biology

All the hornets have an annual life cycle. Nests in temperate regions are founded in spring by a single queen after the over-wintering hibernation period. In Vespa velutina, the queen (also known as a foundress) quickly builds a small embryo nest in which to rear the first batch of workers (females) in an enclosed and protected place (a wall cavity, tree hollow, shed, porch etc.). Nests are built from wood removed from dead trees, shrubs, posts, etc. These nests are only small -- they consist of 30-40 cells and are initially the size of a lemon. It appears that all nests are only ever used once and are destroyed by birds or the weather, although the same nest-sites can be used year after year. It takes 30-50 days for the first batch of workers to emerge in sub-tropical and temperate regions (Dong and Wang, 1989; Choi et al., 2012). If the site does not allow for colony expansion, relocation is triggered and the colony relocates and builds a secondary nest up to 200 m from the primary nest (Matsuura and Yamane, 1990)

After this the queen's duties gradually become confined to egg laying as the workers take over the duties of foraging and nest building. As the number of workers increases, the nest undergoes a rapid period of expansion before the queen switches to laying eggs which become sexual males and females (future queens, known as gynes). By the end of summer, the colony can reach a maximum size of more than 1000 adult workers (average in a French study by Rome et al. 2015 was 436 workers present between late October and early November) and hundreds to thousands of sexuals (i.e. potential queens and males) (Choi et al., 2012; Archer, 2012). In France, Villemant et al. (2011) found an average of 350 gynes (future queens) vs 900 males. When the new sexuals start to emerge, the females feed on larval regurgitations to build up their fat bodies, after which they leave the nest and mate, usually up in the tree canopy. The mother queen lives for about one year. After mating the males die. The workers probably die of old age in the tropics and not starvation due to the onset of winter as they do in temperate regions. The fertilised queens undergo a period of dormancy in temperate climates and hibernate during winter. They can hibernate alone or in clusters of 2 to 3 under the bark of trees or under stones. The mortality rate of overwintering queens is not known. Because of the speed of spread of V. velutina in France (around 78 km per year -- Robinet et al., 2017), it is assumed that fertile queens migrate after winter hibernation.

Nests can be initiated at any time of the year in tropical regions but are only built yearly in spring in temperate climates; they are left empty during winter. There is a positive correlation between the nest size and the number of queens raised (Chauzat and Martin 2009). If the queen dies before sexual production, some of the workers will lay unfertilized (haploid) eggs which will develop into males but the colony will usually perish as any new queens do not lay eggs until some weeks after fertilization. Queenless V. velutina nests have been reported (F. Muller, Museum National d'Histoire Naturelle, Paris, France, personal communication, 2013).

Nutrition

Kuo and Yeh (1990) observed that V. velutina was better at catching prey (e.g. flies and honey bees: for further detail on types of prey, see ‘Impact on Biodiversity’ below) as it is much faster and more agile than other hornet species. V. velutina has been reported to limit colony development of European honey bees in Asia by the persistent predation of adult bees (Shah and Shah, 1991). In fact, it is one of the most adept hornets at catching honey bees on the wing: other species land on the hive and grab bees that try and attack the hornet, whereas V. velutina hovers in front of the hive at a distance of 30–40 cm, and swoops down trying repeatedly to catch foragers. Once a bee has been caught it may be taken straight to the nest, or is carried off to a tree branch where the head, wings, legs and abdomen are removed (C. Villemant, Museum National d'Histoire Naturelle, Paris, France, personal communication, 2013) and a meatball made of the flight muscle which is fed to larvae back in the nest. The attempts by hornets to attack honey bee colonies are numerous and frequent, particularly at the end of the season (September to December) when the production of new queens makes high demands on hornet workers (Mollet and Torre 2006). Adult V. velutina do not consume meat themselves (although they may ingest meat juices), but they feed the larvae in the colony on insect prey and meat from mammal and bird carcasses, as well as fish and meat from fishmonger’s and butcher’s stalls. Adult V. velutina feed on sweet carbohydrates, such as nectar, ripe fruit and tree sap, and on special regurgitations from the larvae (Matsuura and Yamane, 1990).

Environmental Requirements

A key characteristic of hornets’ success is their resilience to environmental change and their capacity to overcome difficulties (Chauzat and Martin, 2009; Villemant et al., 2011; Barbet-Massin et al., 2013). Natural population control may occur due to lack of prey, bad weather at key stages of the life cycle, limited nest sites and usurpation (queen fighting) (Martin, 1992).

One of the key factors that allows hornets to be highly successful predators is their ability to thermoregulate their nests to a constant temperature of around 30°C even if ambient temperatures are lower (Spradberry, 1973; Martin, 1990). When temperatures reach maximum levels in summer months, hornets ventilate their nest. Workers regurgitate water from their mandibles onto the nest (sometimes soaking the nest wall), and evaporate that water through the vibration of their wings to cool down the colony (Perrard et al. 2009).

Climate

Notes on Natural Enemies

The nests of V. velutina have some natural predators in Aquitaine (France) -- in the period of pre-winter decline of the colony, green woodpeckers (Picus viridis), jays (Garrulus glandarius) and tits (Paridae) are often seen pillaging nests and eating the remaining larvae -- but these are unable to attack large active colonies. There have been rare records of more specialist enemies such as the European honey buzzard (Pernis apivorus) and the European bee-eater (Merops apiaster), but these have a negligible impact on the population (F. Muller, Museum National d'Histoire Naturelle, Paris, France, personal communication, 2013). It is not yet known if there are other hornet predators (Mollet and Torre, 2006).

Like other wasps, V. velutina is susceptible to various diseases and parasites, including Conops vesicularis (a parasitic fly), and various nematodes and entomopathogenic fungi (Turchi and Derijard, 2018).

Population levels are quite low in Asia compared to Europe, most probably due to competition with many other Vespa species for food resources and nesting sites (Matsuura, 1984). In France, extremely high densities of nests have been reported: 8 nests per km² in mixed land usage to 23 nests per km² in an urban area (Franklin et al., 2017).

Means of Movement and Dispersal

The most likely introduction pathway for V. velutina is accidental introduction via trading activities. The fact that the fertilized queens can survive long periods of time hibernating makes it an ideal candidate for long distance transport in shipments of goods, for example in containers, in pots, in building material, under the bark of trees, in decorative material or wooden items or in cars, boxes, trucks, farming equipment, etc.

Only one mated queen is needed to start a new colony and initiate further spread of the species. Arca et al. (2015) showed that the French population may have originated from a single multi-mated queen.

In France, V. velutina spread to cover 120,000 km² within 3 years, demonstrating that it can colonize large areas in a very short period of time if the climatic conditions are favourable.

Pathway Causes

Pathway Vectors

Impact Summary

Economic Impact

In parts of Asia (Kashmir and China), V. velutina is considered an enemy of honey bees. This species is able to destroy up to 30% of a colony of the Asian honey bee Apis cerana. Hornet workers attack the honey bee guards one by one, before robbing their brood nest in order to feed their own larvae. The hawking area in front of the hives is territorial -- other hornets that intrude are quickly expelled. In Yunnan Province, China, hornet predation levels are highest in the morning and afternoon, which corresponds with the daily rhythm of honey bee flights (Tan et al., 2005 and 2007). Attack by V. velutina can lead to colony death (Qun, 2001; Tan et al., 2005); if a honey bee colony becomes sufficiently deprived of workers, V. velutina will then enter the hive, feed on the honey and remove the brood.

The introduction into Europe of V. velutina is certainly detrimental to beekeepers. In France, the data is still patchy, with some areas experiencing greater losses than others. There is little data available on actual colony losses, but the figure of 30% destroyed or weakened is generally referred to (Monceau et al., 2014). Colonies rarely fail while being attacked, but are more likely to die off during the winter due to too few bees in the winter cluster, lack of stores due to ‘foraging paralysis’ (foragers do not risk leaving the hive when under attack from hornets) or a combination of both of these (Requier et al., 2019). As in Asia, very weak colonies with little hive activity and guard behaviour may be entered by V. velutina and robbed of honey, pollen, larvae and pupae (Arca et al., 2014). Even though some honey bees from Asia and Cyprus have developed strategies to defend themselves against other species of hornets (Ono et al., 1995; Papachristoforou et al., 2007), western European honey bees may not be able to fend off this intruder.

There is little economic data available on potential losses to bee keepers and/or to the bee/honey industries. This is partly because it is difficult to disentangle the causes of colony losses, which may be due to weather, insecticides, disease, parasites, etc. A recent study (Ferreira-Golpe et al., 2018) found that Spanish beekeepers used 20% of the value of their production in combating V. velutina, using a variety of strategies.

V. velutina can cause damage to fruit (Fédération Départementale des Groupements de Défense contre les Organismes Nuisibles de Loire-Atlantique, 2015; Abeilles et Fleurs, 2017).

Environmental Impact

Impact on Habitats

Studies are needed to assess the impact at an ecological scale of the introduction of a new predator such as V. velutina on the balance of the whole ecosystem.

Hornets use plant material for nest-building -- they gnaw at branches to extract material which they chew up to build their "paper" nests. There is no obvious adverse impact on vegetation from this activity.

Impact on Biodiversity

Hornets are well known for their attacks on other hymenopteran species, especially honey bees. Studies by Muller et al. (2010, 2013) in France demonstrate that V. velutina preys on a range of insects and the carcasses of mammals and birds. The prey spectrum consisted of 59% hymenopteran species [of which bees (Apidae) represented over 35%], 32% dipterans, and 9% others (orders Hemiptera, Orthoptera, Lepidoptera, Mecoptera, Trichopetra, Coleoptera, Heteroptera, Neuroptera, Dermaptera, and Blattaria). Prey collected in different environments varied greatly, which corroborates earlier studies on the adaptability of this species (Matsuura and Yamane, 1990). Previous work also reported that the main prey of V. velutina was Brachycera (Diptera) and social hymenopteran species including bumblebees and honeybees (Vecht, 1957; Williams, 1988; Abrol, 1994). Perrard et al. (2009) studied the activity and behaviour of colonies established in the Southwest of France and determined that V. velutina can also attack halictids.

Choi et al. (2012) found that the arrival of V. velutina had disturbed the relative abundance of the 6 previously known species of Vespa in Busan city, Korea. V. velutina now accounts for 37% of the Vespa population and has caused a 20% drop in V. simillima and a 10% drop in V. mandarinia.

Social Impact

Concerning its potential danger to man, V. velutina is not considered to be more aggressive than the European hornet, Vespa crabro (Haro and Blanc-Brisset, 2009; Haro et al., 2010); the medical literature indicates that in comparison with other hornet species, it is not a major health threat in Asia (Haro et al., 2010). Although V. velutina aggressiveness towards humans is widely recognised among the inhabitants of Java (Vecht, 1957) and Taiwan (Matsuura, 1973; Ho et al., 1999), its aggressiveness observed in France by Perrard et al. (2009) was low.

There are several newspaper reports of people being stung to death by V. velutina, but many of them have not been confirmed by a scientific or medical authority (C. Villemant, Muséum National d’Histoire Naturelle, Paris, France, personal communication, 2013, referring to Schwartz et al., 2012). A review of data from French Poison Control Centres between 2004 and 2011 showed only a few envenomations clearly linked to V. velutina (Haro et al. 2010; Schwartz et al 2012). Moreover, the increase of the V. velutina population in southwestern France is not correlated with an increase in the number of hymenopteran stings (Haro et al., 2010), which are mainly due to honey bees and ordinary wasps. There is however some concern about the risk of fruit pickers being stung while harvesting fruit (Fédération Départementale des Groupements de Défense contre les Organismes Nuisibles de Loire-Atlantique, 2015; Abeilles et Fleurs, 2017).

Like the native European hornet, V. velutina can be dangerous for man by inducing a life-threatening allergic reaction or after multiple stings. However, severe attacks only occur when colonies are disturbed, and because the majority of V. velutina nests hang very high in trees, such accidents remain rare (Rome et al., 2011), although with increasing densities of nests in urban areas, humans are likely to accidentally disturb nests more frequently.

Risk and Impact Factors

• Proved invasive outside its native range
• Has a broad native range
• Abundant in its native range
• Highly adaptable to different environments
• Is a habitat generalist
• Capable of securing and ingesting a wide range of food
• Benefits from human association (i.e. it is a human commensal)

• Conflict
• Negatively impacts agriculture
• Negatively impacts animal/plant collections

• Herbivory/grazing/browsing
• Predation

• Highly likely to be transported internationally accidentally
• Difficult/costly to control

Diagnosis

Molecular techniques to diagnose V. velutina are in development for use in the lab and field (Stainton et al., 2018).

Similarities to Other Species/Conditions

V. velutina is very easy to recognize since it is the only hornet or wasp that has an entirely dark brown-black body with a thin yellow stripe on the dorsal border of the 1^st abdominal tergite, an orange-yellow band towards the end of the abdomen (4th tergite), and yellow legs (C. Villemant, Muséum National d’Histoire Naturelle, Paris, France, personal communication, 2013). Being very dark, individuals stand out as dark spots in front of beehives or their own nests. V. velutina is also smaller than the two other species found in Europe and/or North America (V. crabro and V. orientalis). It also tends to build larger nests. In Java, a nest was found that was 75 cm high, containing around 12,000 brood cells in 11 combs (Vecht, 1957). In France, the largest nest dissected by Rome et al. (2015) had 11 combs and an estimated 13,547 cells.

Prevention and Control

Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

So far, countries that have been invaded by wasp species, such as New Zealand, Australia and the islands of Hawaii, have failed to control their populations despite many attempts (Beggs et al., 2011). These have included the introduction of parasitic nematodes, poison baits and entomopathogenic fungi (Martin, 2004; Beggs et al., 2011). Measures to reduce the expansion of V. velutina in Europe are reviewed by Turchi and Derijard (2018) but most efforts are concerned with the physical destruction of individual nests; although it is often difficult to spot these as they can remain hidden until leaf fall in autumn.

Current recommendations in France are aimed at bee keepers and apiarists in order to protect their bee colonies.

In Asia, the Eastern honey bee Apis cerana has developed two defensive behaviours against predation by hornets: a warning system for returning workers by shimmering their wings in unison (Tan et al., 2005) and a thermo-balling system whereby intruders are suffocated and heated to death (up to 45°C) by a ball of bees (Ono, 1995; Tan et al., 2005; Papachristoforou et al., 2007). In Kashmir, a colony of A. cerana can kill 10 hornets a day, whereas the European honey bee manages on average to only kill one (Abrol, 2006; Villemant, 2008). European honey bees can form balls but these are much less efficient at killing the hornet than A. cerana balls (Tan et al., 2005 and 2007). They might be able, however, to adapt with time.

Public Awareness

In the UK, beekeepers are encouraged to be alert for the presence of V. velutina, and anyone seeing it is asked to report the sighting to www.nonnativespecies.org/alerts/asianhornet, or alertnonnative@ceh.ac.uk (NNSS, undated), or use the Asian Hornet Watch app.

Eradication

Eradication of V. velutina from France is now impossible as it has become too well established over a large area (Villemant et al., 2011; Rome et al., 2009, 2012, 2013; Chauzat and Martin, 2009).

Physical/mechanical control

In India, control strategies such as killing hornet queens in early spring, destroying hornet nests and swatting hornets at hive entrances have been advocated. Unfortunately, none of these have been reported to be effective, although mass trapping in jars half-filled with fermented honey-water drove numbers of hornets hawking at the apiary from 10-25 per hive to 0-3 (Shah and Shah, 1991). In France similar methods have been tried with similar results; for example in September 2007, a beekeeper in the Bordeaux area killed about 80 hornets a day over several weeks without any decrease in the hornet pressure on his honey bee colonies (Chauzat and Martin, 2009).

Chemical control

Hornets have two basic dietary requirements: sugars for energy for the adults, and proteins for the nourishment of the brood: baiting (and poisoning) could therefore be an option. However, this could have adverse effects on non-target species (Beggs et al., 2011).

Biological control

Studies on indigenous species of entomopathogenic fungi in the genera Beauveria and Metarhizium from France show some potential for control of V. velutina nigrithorax (Poidatz et al., 2018).

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Distribution References

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Archer M, 2012. Vespine wasps of the world: behaviour, ecology and taxonomy of the Vespinae., UK: Siri Scientific Press. 352 pp.

Bertolino S, Lioy S, Laurino D, Manino A, Porporato M, 2016. Spread of the invasive yellow-legged hornet Vespa velutina (Hymenoptera: Vespidae) in Italy. Applied Entomology and Zoology. 51 (4), 589-597. http://link.springer.com/article/10.1007/s13355-016-0435-2 DOI:10.1007/s13355-016-0435-2

Budge G E, Hodgetts J, Jones E P, Ostojá-Starzewski J C, Hall J, Tomkies V, Semmence N, Brown M, Wakefield M, Stainton K, 2017. The invasion, provenance and diversity of Vespa velutina Lepeletier (Hymenoptera: Vespidae) in Great Britain. PLoS ONE. 12 (9), e0185172. http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0185172 DOI:10.1371/journal.pone.0185172

CABI, Undated. CABI Compendium: Status as determined by CABI editor. Wallingford, UK: CABI

CABI, Undated. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI

CABI, Undated. Compendium record. Wallingford, UK: CABI

Choi MoonBo, Martin S J, Lee JongWook, 2012. Distribution, spread, and impact of the invasive hornet Vespa velutina in South Korea. Journal of Asia-Pacific Entomology. 15 (3), 473-477. http://www.sciencedirect.com/science/article/pii/S1226861511001324 DOI:10.1016/j.aspen.2011.11.004

DEFRA, 2018. Asian hornet: UK sightings in 2018., UK: Department for Environment, Food & Rural Affairs. https://www.gov.uk/government/news/asian-hornet-uk-sightings-in-2018?utm_source=cec6c1f9-f157-4b1b-acee-779da3e783f3&utm_medium=email&utm_campaign=govuk-notifications&utm_content=immediate

EPPO, 2007. EPPO Global database (available online). Paris, France: EPPO. https://gd.eppo.int/

Federazione Apicoltori Italiani, 2013. Vespa velutina: first record in Italy. Alarm! (Vespa velutina: prima segnalazione in Italia. E' allarme!)., Rome, Italy: Federazione Apicoltori Italiani. http://www.federapi.biz/index.php?option=com_content&task=view&id=1176&Itemid=1

Goldarazena A, Heredia I P de, Romon P, Iturrondobeitia J C, Gonzalez M, Lopez S, 2015. Spread of the yellow-legged hornet Vespa velutina nigrithorax du Buysson (Hymenoptera: Vespidae) across Northern Spain. Bulletin OEPP/EPPO Bulletin. 45 (1), 133-138. http://onlinelibrary.wiley.com/journal/10.1111/(ISSN)1365-2338 DOI:10.1111/epp.12185

Grosso-Silva JM, Maia M, 2012. Vespa velutina Lepeletier, 1836 (Hymenoptera, Vespidae), new species for Portugal. In: Arquivos Entomolóxicos, 6 53-54.

Haxaire J, Bouguet J P, Tamisier J P, 2006. Vespa velutina Lepeletier, 1836, a fearsome new addition to the French fauna (Hym., Vespidae). (Vespa velutina Lepeletier, 1836, une redoutable nouveauté pour la faune de France (Hym., Vespidae).). Bulletin de la Société Entomologique de France. 111 (2), 194.

Kim JeongKyu, Choi MunBo, Moon TaeYoung, 2006. Occurrence of Vespa velutina Lepeletier from Korea, and a revised key for Korean Vespa species (Hymenoptera: Vespidae). Entomological Research. 36 (2), 112-115. http://www.blackwell-synergy.com/doi/abs/10.1111/j.1748-5967.2006.00018.x DOI:10.1111/j.1748-5967.2006.00018.x

Kishi S, Goka K, 2017. Review of the invasive yellow-legged hornet, Vespa velutina nigrithorax (Hymenoptera: Vespidae), in Japan and its possible chemical control. Applied Entomology and Zoology. 52 (3), 361-368. https://link.springer.com/article/10.1007/s13355-017-0506-z DOI:10.1007/s13355-017-0506-z

Leza M, Miranda M Á, Colomar V, 2018. First detection of Vespa velutina nigrithorax (Hymenoptera: Vespidae) in the Balearic Islands (Western Mediterranean): a challenging study case. Biological Invasions. 20 (7), 1643-1649. https://link.springer.com/article/10.1007/s10530-017-1658-z DOI:10.1007/s10530-017-1658-z

López S, González M, Goldarazena A, 2011. Vespa velutina Lepeletier, 1836 (Hymenoptera: Vespidae): first records in Iberian Peninsula. Bulletin OEPP/EPPO Bulletin. 41 (3), 439-441. http://onlinelibrary.wiley.com/journal/10.1111/(ISSN)1365-2338 DOI:10.1111/j.1365-2338.2011.02513.x

Minoshima Y N, Yamane S, Ueno T, 2015. An invasive alien hornet, Vespa velutina nigrithorax du Buysson (Hymenoptera, Vespidae), found in Kitakyushu, Kyushu Island: a first record of the species from mainland Japan. Japanese Journal of Systematic Entomology. 21 (2), 259-261.

Muséum National d'Histoire Naturelle, 2012. Vespa velutina., Paris, France: Muséum National d'Histoire Naturelle. http://inpn.mnhn.fr/isb/espece/cd_nom/433589

Nguyen Thi Phuong Lien, Carpenter J M, 2002. Vespidae of Vietnam (Insecta: Hymenoptera) 1. Vespinae. Journal of the New York Entomological Society. 110 (2), 199-211. DOI:10.1664/0028-7199(2002)110[0199:VOVIHV]2.0.CO;2

Rome Q, 2019. The Asian Hornet, Vespa velutina. (Le Frelon asiatique, Vespa velutina)., Paris, France: Muséum national d’Histoire naturelle. http://frelonasiatique.mnhn.fr/home/

Rome Q, Dambrine L, Onate C, Muller F, Villemant C, García Pérez AL, Maia M, Carvalho Esteves P, Bruneau E, 2013. Spread of the invasive hornet Vespa velutina Lepeletier, 1836, in Europe in 2012 (Hym., Vespidae). In: Bulletin de la Société entomologique de France, 118 21-22. http://inpn.mnhn.fr/docs/Vespa_velutina/1643-1-Rome_et_al_2013_p.pdf

Smit J, Noordijk J, Zeegers T, 2018. Will the Asian hornet (Vespa velutina) settle in the Nederlands? (De opmars van de Aziatische hoornaar (Vespa velutina) naar Nederland.). Entomologische Berichten. 78 (1), 2-6. http://www.nev.nl/pages/publicaties/eb/

States of Alderney, 2019. Asian Hornet Spring Trapping Programme., Alderney, Channel Islands: States of Alderney. http://www.alderney.gov.gg/article/170204/Asian-Hornet-Spring-Trapping-Programme

States of Guernsey Government, 2016. Asian hornet identified in Alderney., https://www.gov.gg/article/155332/Asian-Hornet-identified-in-Alderney

States of Jersey, 2019. Asian Hornet sightings, identification and reporting., Jersey, Channel Islands: States of Jersey. https://www.gov.je/Environment/LandMarineWildlife/Insects/Pages/Asian-Hornet-sighting,-identification-and-reporting.aspx

Takahashi R, Okuyama H, Minoshima Y N, Takahashi J I, 2018. Complete mitochondrial DNA sequence of the alien hornet Vespa velutina (Insecta: Hymenoptera) invading Kyushu Island, Japan. Mitochondrial DNA Part B. 3 (1), 179-181. https://www.tandfonline.com/doi/full/10.1080/23802359.2018.1437823 DOI:10.1080/23802359.2018.1437823

Takeuchi T, Takahashi R, Kiyoshi T, Nakamura M, Minoshima Y N, Takahashi J, 2017. The origin and genetic diversity of the yellow-legged hornet, Vespa velutina introduced in Japan. Insectes Sociaux. 64 (3), 313-320. http://rd.springer.com/journal/40 DOI:10.1007/s00040-017-0545-z

Villemant C, Haxaire J, Streito JC, 2006. [English title not available]. (Premier bilan de l'invasion de Vespa velutina Lepeletier en France (Hymenoptera, Vespidae)). In: Bulletin de la Société entomologique de France, 111 (4) 235-238.

Villemant C, Haxaire J, Streito JC, 2006. The discovery of the Asian hornet Vespa velutina in France. (La découverte du frelon asiatique Vespa velutina, en France). In: Insectes, 143 (4) 3-7. http://www.inra.fr/opie-insectes/pdf/i143villemant-haxaire-streito.pdf

Links to Websites

Organizations

France: EPPO, European and Mediterranean Plant Protection Organization, 21 boulevard Richard Lenoir, 75011 Paris, http://www.eppo.int/

UK: National Bee Unit, The Animal and Plant Health Agency (APHA), National Agri-Food Innovation Campus, Sand Hutton, York,

Contributors

21/05/19: Updated by:

Sarah Bunker, consultant, UK

10/08/10 Original text by:

CRCNPB Australia, CRC for National Plant Biosecurity, Canberra, Australia

Distribution Maps