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Vespa mandarinia

By Zach Barth; Thomas Kearns; Elizabeth Wason

Geographic Range

Vespa mandarinia is found in eastern and southeastern parts of Asia, north of the tropics. It is most common in Japan, where it has been well studied. (Abe, et al., 1982; Matsuura and Sakagami, 1973)

Habitat

Vespa mandarinia tends to live in low mountains and/or forested areas. It is notably absent in plains and at higher elevations. The species creates subterranean nests by digging, coopting cavities that were dug by small rodents, or finding suitable spaces near rotted pine roots. The depth of their nests ranges between 6 and 60 cm. (Matsuura and Sakagami, 1973; Yamane, 1976)

Physical Description

The Asian giant hornet is so named because it is the world’s largest hornet. A queen’s body length can exceed 5 cm, with a wingspan that can exceed 7.6 cm. Males and the female workers are considerably smaller than the queens (3.5 to 3.9 cm in body length). Queens and workers differ only in size. Both queens and workers have reproductive anatomy, although workers do not reproduce. Males of this species are morphologically similar to females, but as in all hymenopterans, males lack the stinger that is evolved from the ovipositor. Coloration is the same between sexes. Vespa mandarinia has a broad head that is a light shade of orange. Its antennae are typically a medium or dark shade of brown, with yellow-orange scapes. It has a set of compound eyes that range from dark brown to black. It also has three simple eyes that are similar in color to that of the compound eye. The Asian giant hornet can be distinguished from other species by the presence of a deeply incised clypeus and a rather large genae. The mandible is relatively large and is a deep orange hue, with a black tooth that can be used for burrowing.

The thorax region has a few distinguishing features. The thorax is dark brown, with a distinctly large scutellum that has a noticeable impressed medial line. Its two sets of wings are typically grey and extend far from the body. Its forelegs are brighter with dark tarsi, while the mid- and hind legs are dark brown throughout.

The abdomen (gaster) contains bands of dark brown and black, which alternate with bands that are a similar orange-yellow hue as the head. The sixth segment is entirely yellow. The stinger region can reach up to 6 mm and the associated toxin is considerably venomous. (Elzinga, 1981; Matsuura and Sakagami, 1973; Shreeves and Field, 2008)

  • Sexual Dimorphism
  • female larger
  • Range length
    3.5 to 5.5 cm
    1.38 to 2.17 in
  • Range wingspan
    3.5 to 7.6 cm
    1.38 to 2.99 in

Development

The Asian giant hornet emerges from its egg as a soft, white, apodous larva. The larva has a yellow head that is covered with chitin. The body has three thoracic and ten abdominal segments that do not have constrictions between them. The larvae feed on a paste consisting of prey that was masticated by the workers or queen. The larvae eventually grow large enough to enter the pupal stage and undergo complete metamorphosis. After metamorphosis, they emerge as adults. The workers require about 40 days to grow from egg to adult. Queens and males develop later in the season, with males maturing faster than queens. (Ross and Matthews, 1991; Yamane, 1976)

Reproduction

Mating in Vespa mandarinia takes place during the fall. In contrast to other hornets, copulation takes place at nest entrances. Sex pheromones are also used to initiate male sexual behavior. Once a queen exits the nest, a male will seize her in mid-air, causing both of them to fall to the ground. Copulation continues for 8 to 45 seconds. Following copulation, the queen finds a place to overwinter. (Richards, 1971; Ross and Matthews, 1991; Wilson, 2005)

After overwintering, a fertilized queen starts her colony in the spring. Once the queen has established a rudimentary nest, she produces an initial clutch of up to 40 eggs, which becomes the first generation of workers. Toward the end of summer, the queen produces thousands of additional eggs. Fertilized eggs become the queens of the next season, and unfertilized eggs become the males of the next season. The new queens and males are cared for by the workers, whereas the first generation of workers were cared for by the queen.

Vespa mandarinia has a eusocial mating system. This type of mating system involves an advanced level of colonial existence. Cooperative care for the young, overlapping generations, and the existence of reproductive and non-reproductive classes are hallmarks of eusociality, all of which are exhibited by V. mandarinia. Cooperative care for the young is observed primarily in the workers that help raise the larvae, although the queen raises the initial workers. Overlapping generations occur during the autumn, when new queens and males begin to mate while the original colony-founding queen is still alive. Vespa mandarinia colonies include diploid queens (reproductive females), workers (non-reproductive females), and haploid reproductive males. (Matsuura and Sakagami, 1973; Ross and Matthews, 1991; Takahashi, 2004; Yamane, 1976)

  • Breeding season
    Mating occurs in the fall, and eggs are laid from spring to fall.
  • Range number of offspring
    40 to 25000
  • Average number of offspring
    10000

Vespa mandarinia exhibits a great deal of parental investment. This hornet species builds nests underground, in tree hollows, or in urban buildings, which workers aggressively defend. The young are fed by adults that have chewed their prey, typically bees, into a fine paste. This paste is then regurgitated and fed to the young. The paste is highly nutritious for the growing larvae. (Lee, 2010; Matsuura and Sakagami, 1973; Ross and Matthews, 1991; Shreeves and Field, 2008)

  • Parental Investment
  • female parental care
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • female
  • pre-independence
    • provisioning
      • female
    • protecting
      • female

Lifespan/Longevity

Workers live from spring until winter. Males typically die shortly after mating in the fall. Queens live for up to one year; they build colonies in the spring after their mating season, while the current adult queens die in the winter. (Matsuura and Sakagami, 1973)

Behavior

Vespa mandarinia is a eusocial species; thus, its social structure involves a complex colonial existence, which includes cooperative care of the young, overlapping generations, and the existence of reproductive and non-reproductive classes.

The species builds subterranean nests in cavities that it digs itself or that have already been dug by small rodents. These nests also can be found near rotted pine roots, in tree hollows, and even in urban structures.

Vespa mandarinia is unique in that it is the only eusocial wasp species to stage group attacks against beehives and other eusocial wasp nests. These attacks do not usually take place until the end of the summer, more than halfway through a colony’s lifespan. These group attacks can be divided into three stages. The first, termed the hunting phase, involves solitary workers waiting near the entrance to a hunted hive. These workers capture prey in flight using their mandibles. The prey are bitten to death, and body parts are removed from the mesosoma. Attacking hornets return to their nest and feed the prey to their larvae. The hunting phase can continue indefinitely, and progression to the next phase depends on the distance between the V. mandarinia nest and the prey colony.

The second phase is termed the slaughter phase. This phase involves 2 to 50 V. mandarinia hornets. The attacking hornets focus on a single hive or nest that has been marked with a chemical secreted by a V. mandarinia worker. The attackers usually stay near the hive entrance and kill counter-attacking prey. The dead bodies of the prey species are ignored during the slaughter phase. Vespa mandarinia casualties are uncommon during attacks on Apis mellifera hives, but up to half of the attackers may be killed in an encounter with another Vespa species. Once engaged in this part of the attack, the attacking hornets will not stop, even if disturbed. In a particularly long slaughter phase, attacking V. mandarinia occasionally will starve to death. The length of the slaughter phase is variable and depends on both the number of attacking hornets and the intensity of the defense.

Once the defense has ceased, the occupation phase begins. Vespa mandarinia becomes territorial, and guards will threaten other animals that come near the occupied hive by audibly clicking their mandibles. Workers often will spend the night in an occupied hive instead of in their own colony’s nest.

Japanese honeybees (unlike some other bee species) have evolved a defense against the aggression exhibited by V. mandarinia. The honeybees can detect the pheromone that V. mandarinia hornets use to mark beehives for future attacks. When they detect this pheromone, the honeybees respond by increasing the number of defenders stationed at the nest entrance. More than 500 honeybees can mob an individual hornet scout, surrounding it faster than the hornet can bite and kill them. The bees form a ball around the hornet, then vibrate their wing muscles in the same way that they warm their nest in the winter. This coordinated behavior raises the temperature and the CO2 concentration to lethal levels around the trapped hornet. The hornet cannot survive, but the bees can withstand the conditions. Thus, the honeybees kill the hornet before it can report the location of the beehive to its nestmates. (Lee, 2010; Matsuura and Sakagami, 1973; Ono, et al., 1995; Ross and Matthews, 1991; Taylor, et al., 2011)

Home Range

Vespa mandarinia workers often fly 1 to 2 km from their nest. They can travel up to 8 km from their nest.

Communication and Perception

Vespa mandarinia uses visual cues for flight navigation. Communication through scent marks and pheromones is also known to occur. Workers apply secretions to the colonies of honeybees and wasps to recruit nestmates for a group attack. A “royal court” has been observed in V. mandarinia. This “royal court” is formed by workers that cluster around a queen, licking and biting her, ingesting pheromones. Sex pheromones also are used to initiate male sexual behavior. Some acoustic communication exists in this species. Hornet larvae make an audible sound by scraping their mandibles along the cell wall, and this is believed to indicate hunger. Adult hornets also click their mandibles as a warning to organisms that come too close to their nest or a nest that they have taken over. (Ross and Matthews, 1991; Taylor, et al., 2011; Toh and Okamura, 2003)

Food Habits

Vespa mandarinia is a primarily insectivorous omnivore. Workers prey mainly on beetles (Coleoptera), but they also will hunt hornworms (Manduca), mantids (Mantodea), and eusocial wasp and bee species.

Vespa mandarinia is the only eusocial wasp species to attack beehives and other eusocial wasp nests. Colonies of Apis mellifera, Apis cerana, and all sympatric Vespa and Vespula species are targeted by V. mandarinia for group attacks. The bee or wasp prey collected in these attacks are chewed into a ball, which is taken to the V. mandarinia nest and fed to the larvae. After a successful attack of a beehive or other nest, the pupae of the prey species are removed from their cocoons. The bodies of larvae and pupae in the occupied hive or nest are chewed into balls and fed to V. mandarinia larvae. The most mature pupae are preferred, and larvae are taken only when no pupae remain. Some of the adult victims and dead attackers also are fed to the V. mandarinia larvae. Workers will retrieve food from an occupied nest for a period of several days to two weeks. Mass attacks are only observed in the autumn, when hundreds of new hornet queens and males are being reared in the nest. These new hornets require large amounts of protein, and the necessity of feeding them may prompt V. mandarinia nestmates to attack beehives.

Vespa mandarinia frequently eats tree sap, especially in oaks (Quercus). Fruits with soft pericarps and high sugar levels are also eaten. Vespa mandarinia is the only Vespa species to frequently engage in extranidal trophallaxis, or feeding outside the nest via regurgitation. Adult workers cannot digest solid foods; thus, they subsist on sap and larval saliva. All solid food is fed to larvae by trophallaxis. Extranidal trophallaxis often occurs among workers at sap sources. Most often, the flow of food is unidirectional. Reciprocal trophallaxis occurs among starving workers. These individuals will tightly embrace, hanging from a branch by a single leg, or lying or rolling on the ground. They alternately beg for and receive food, and they may maintain this embrace for over ten minutes. (Abe, et al., 1995; Matsuura and Sakagami, 1973; Ono, et al., 1995; Ross and Matthews, 1991; Taylor, et al., 2011)

  • Animal Foods
  • insects
  • Plant Foods
  • fruit
  • sap or other plant fluids

Predation

Predators of Vespa mandarinia appear to be rare; however, V. mandarinia nests are attacked by conspecific colonies, and honey buzzards may prey on this hornet.

Anti-predator adaptations include aposematic clicking with their mandibles and stinging. (Lack, 1946; Matsuura and Sakagami, 1973; Schmidt, et al., 1986)

Ecosystem Roles

The Asian giant hornet dominates insect tree sap communities during the day, affecting the foraging behaviors of other species. Less dominant species must either wait for V. mandarinia to leave the patch or look for a separate patch elsewhere. Vespa mandarinia occupies the highest rank in the arthropod food web within its geographic range. During the later stages of the nesting period, V. mandarinia is the most perilous natural enemy of sympatric Vespa species at the colony level. In some years, more than half of the colonies of Polistes testaceicolor, Vespa simillima, and Vespula flaviceps may be exterminated within a particular area due to V. mandarinia group attacks.

Predators of V. mandarinia may include Pernis apivorus (honey buzzard). Vespa mandarinia also is a host to Xenos moutoni, a strepsipteran endoparasite. (Lack, 1946; Maeta, et al., 1998; Ross and Matthews, 1991; Yoshimoto, 2009)

Commensal/Parasitic Species
  • Xenos moutoni

Economic Importance for Humans: Positive

A compound derived from Vespa mandarinia larval saliva (called Vespa Amino Acid Mixture) is sold as a nutritional supplement. Mice that were administered VAAM have demonstrated increased swimming endurance, along with decreased lactate and increased glucose concentrations in their blood after swimming. These results suggest that VAAM inhibits muscle catabolism during endurance exercise and could be used to improve athletic performance. However, at the time of this writing, no peer-reviewed scientific evidence exists to suggest that this effect occurs in humans. (Abe, et al., 1995)

Economic Importance for Humans: Negative

Vespa mandarinia can deliver a venomous sting that is harmful to humans. In Japan, V. mandarinia causes an estimated 30 to 50 deaths each year.

Vespa mandarinia also is a major pest of apiaries in Japan. Apis mellifera is the most common apicultural species, and it has not evolved defenses against V. mandarinia. Attacks on an A. mellifera hive by 20 to 30 Asian giant hornets usually will result in the deaths of 5,000 to 25,000 bees within 1 to 6 hours. Tens of thousands of A. mellifera hives are damaged each year by V. mandarinia. (Matsuura, 1988; Yanagawa, et al., 2007)

  • Negative Impacts
  • injures humans
  • crop pest

Conservation Status

Currently, there are no efforts to conserve this species.

Contributors

Zach Barth (author), The College of New Jersey, Thomas Kearns (author), The College of New Jersey, Elizabeth Wason (author, editor), Animal Diversity Web Staff, Keith Pecor (editor), The College of New Jersey.

Glossary

Palearctic

living in the northern part of the Old World. In otherwords, Europe and Asia and northern Africa.

World Map

acoustic

uses sound to communicate

agricultural

living in landscapes dominated by human agriculture.

aposematic

having coloration that serves a protective function for the animal, usually used to refer to animals with colors that warn predators of their toxicity. For example: animals with bright red or yellow coloration are often toxic or distasteful.

carnivore

an animal that mainly eats meat

chemical

uses smells or other chemicals to communicate

colonial

used loosely to describe any group of organisms living together or in close proximity to each other - for example nesting shorebirds that live in large colonies. More specifically refers to a group of organisms in which members act as specialized subunits (a continuous, modular society) - as in clonal organisms.

delayed fertilization

a substantial delay (longer than the minimum time required for sperm to travel to the egg) takes place between copulation and fertilization, used to describe female sperm storage.

diurnal
  1. active during the day, 2. lasting for one day.
drug

a substance used for the diagnosis, cure, mitigation, treatment, or prevention of disease

eusocial

the condition in which individuals in a group display each of the following three traits: cooperative care of young; some individuals in the group give up reproduction and specialize in care of young; overlap of at least two generations of life stages capable of contributing to colony labor

female parental care

parental care is carried out by females

fertilization

union of egg and spermatozoan

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

fossorial

Referring to a burrowing life-style or behavior, specialized for digging or burrowing.

frugivore

an animal that mainly eats fruit

herbivore

An animal that eats mainly plants or parts of plants.

hibernation

the state that some animals enter during winter in which normal physiological processes are significantly reduced, thus lowering the animal's energy requirements. The act or condition of passing winter in a torpid or resting state, typically involving the abandonment of homoiothermy in mammals.

insectivore

An animal that eats mainly insects or spiders.

internal fertilization

fertilization takes place within the female's body

metamorphosis

A large change in the shape or structure of an animal that happens as the animal grows. In insects, "incomplete metamorphosis" is when young animals are similar to adults and change gradually into the adult form, and "complete metamorphosis" is when there is a profound change between larval and adult forms. Butterflies have complete metamorphosis, grasshoppers have incomplete metamorphosis.

mountains

This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.

native range

the area in which the animal is naturally found, the region in which it is endemic.

omnivore

an animal that mainly eats all kinds of things, including plants and animals

oriental

found in the oriental region of the world. In other words, India and southeast Asia.

World Map

oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

pheromones

chemicals released into air or water that are detected by and responded to by other animals of the same species

scent marks

communicates by producing scents from special gland(s) and placing them on a surface whether others can smell or taste them

seasonal breeding

breeding is confined to a particular season

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

sperm-storing

mature spermatozoa are stored by females following copulation. Male sperm storage also occurs, as sperm are retained in the male epididymes (in mammals) for a period that can, in some cases, extend over several weeks or more, but here we use the term to refer only to sperm storage by females.

tactile

uses touch to communicate

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

terrestrial

Living on the ground.

territorial

defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement

venomous

an animal which has an organ capable of injecting a poisonous substance into a wound (for example, scorpions, jellyfish, and rattlesnakes).

visual

uses sight to communicate

References

Abe, T., A. Niwa, A. Niwa. 1982. Purification and properties of a presynaptically acting neurotoxin, mandaratoxin, from hornet (Vespa mandarinia). Biochemistry, 21/7: 1693-1697.

Abe, T., Y. Takiguchi, M. Tamura, J. Shimura, K. Yamazaki. 1995. Effects of Vespa Amino Acid Mixture (VAAM) isolated from hornet larval saliva and modified VMM nutrients on endurance exercise in swimming mice: improvement in performance and changes of blood lactate and glucose. Japanese Journal of Physical Fitness and Sports Medicine, 44: 225-238.

Elzinga, R. 1981. Fundamentals of Entomology. Prentice-Hall inc.: Prentice-Hall.

Lack, D. 1946. Competition for food by birds of prey. Journal of Animal Ecology, 15/2: 123-129.

Lee, J. 2010. Notes on Polistes testaceicolor and Vespa mandarinia (Hymenoptera, Vespidae) in Hong Kong, and a key to all Vespa species known from the SAR. Hong Kong Entomological Bulletin, 2/2: 31-36. Accessed April 22, 2013 at http://hkentsoc.org/bulletin/HKEB2(2)_vespa_Lee.pdf.

Maeta, Y., K. Takahashi, N. Shimada. 1998. Host body size as a factor determining the egg complement of Strepsiptera, an insect parasite. International Journal of Insect Morphology and Embryology, 27/1: 27-37.

Matsuura, M. 1988. Ecological study on vespine wasps (Hymenoptera: Vespidae) attacking honeybee colonies. I. Seasonal changes in the frequency of visits to apiaries by vespine wasps and damage inflicted, especially in the absence of artificial protection. Applied Entomology and Zoology, 23/4: 428-440.

Matsuura, M., S. Sakagami. 1973. A bionomic sketch of the giant hornet, Vespa mandarinia, a serious pest for Japanese apiculture. Journal of the Faculty of Science Hokkaido University Series VI: Zoology, 19/1: 125-162. Accessed October 24, 2012 at http://133.87.26.249/dspace/bitstream/2115/27557/1/19%281%29_P125-162.pdf.

Ono, M., T. Igarashi, E. Ohno, M. Sasaki. 1995. Unusual thermal defence by a honeybee against mass attack by hornets. Nature, 377: 334-336.

Richards, O. 1971. The biology of social wasps (Hymenoptera, Vespida). Biological Reviews, 46/4: 483-528.

Ross, K., R. Matthews. 1991. The Social Biology of Wasps. Ithaca, NY: Cornell University Press..

Schmidt, J., S. Yamane, M. Matsuura, C. Starr. 1986. Hornet venoms: Lethalities and lethal capacities. Toxicon, 24/9: 950-954.

Shreeves, G., J. Field. 2008. Parental care and sexual size dimorphism in wasps and bees. Behavioral Ecology and Sociobiology, 62/5: 843-852.

Takahashi, J. 2004. Mating structure and male production in the giant hornet Vespa mandarinia. Applied Entomology and Zoology, 39/2: 343-349.

Taylor, B., E. Nordheim, T. Schueller, R. Jeanne. 2011. Recruitment in swarm-founding wasps: Polybia occidentalis does not actively scent-mark carbohydrate food sources. Psyche: A Journal of Entomology, 2011: 1-7.

Toh, Y., J. Okamura. 2003. Foraging navigation of hornets studied in natural habitats and laboratory experiments. Zoological Science, 20/3: 311-324.

Wilson, E. 2005. Eusociality: origin and consequences. Proceedings of the National Academy of Sciences, 102/38: 3367-3371.

Yamane, S. 1976. Morphological and taxonomic studies on vespine larvae, with reference to the phylogeny of the subfamily Vespinae (Hymenoptera: Vespidae). Insecta Matsumurana, New Series 8: 1-45.

Yanagawa, Y., K. Morita, T. Sugiura, Y. Okada. 2007. Cutaneous hemorrhage or necrosis findings after Vespa mandarinia (wasp) stings may predict the occurrence of multiple organ injury: A case report and review of literature. Clinical Toxicology, 45/7: 803-807.

Yoshimoto, J. 2009. Interspecific variation in competitor avoidance and foraging success in sap-attracted insects. European Journal of Entomology, 106/4: 529-533.

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