Oriental hornet

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Oriental hornet
Oriental hornet (Vespa orientalis).jpg
on Cyprus
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Vespidae
Genus: Vespa
Species:
V. orientalis
Binomial name
Vespa orientalis
Linnaeus, 1771
Vespa orientalis Distribution Map2.svg
Distribution of Vespa orientalis [2] [3] [4]

The Oriental hornet (Vespa orientalis) is a social insect species of the family Vespidae. It can be found in Southwest Asia, Northeast Africa, the island of Madagascar (but no reports have been made of its presence on the island for many years), the Middle East, Central Asia, and parts of Southern Europe. [2] Oriental hornets have also been found in a few isolated locations such as Mexico and Chile due to human introduction. [5] [6] The Oriental hornet lives in seasonal colonies consisting of caste system dominated by a queen. [2] The hornet builds its nests underground and communicates using sound vibrations. [7] The hornet has a yellow stripe on its cuticle (exoskeleton), which can absorb sunlight to generate a small electrical potential, and this might help supply energy for digging. [8] The adult hornet eats nectar and fruits and scavenges for insects and animal proteins to feed to its young. [9] Because they are scavengers, the hornets may also serve as a transmitter of disease following consumption of infected plants. [10] The hornets are a primary pest to honey bees, attacking bee colonies to obtain honey and animal proteins. [11] The sting of an Oriental hornet can be quite painful to humans and some humans are allergic to stings. [10]

Contents

Taxonomy and phylogeny

The Oriental hornet (V. orientalis) belongs to the family Vespidae, which consists of wasps, hornets, and yellowjackets. It is a member of the genus Vespa , which constitutes true hornets. [2] V. orientalis has unique adaptations to arid climates, which has made it difficult to assess its phylogenetic relationship to other species of the genus Vespa, based on morphological data alone. [12] Thus, only recently has the use of molecular phylogenetics given evidence that V. orientalis is most closely related to Vespa affinis and Vespa mocsaryana . [12] While a history exists of recognizing subspecies within many of the hornets, including V. orientalis, the most recent taxonomic revision of the genus treats all subspecific names in the genus Vespa as synonyms, effectively relegating them to no more than informal names for regional color forms. [13]

Description and identification

The adult hornet has two pairs of wings and a body measuring between 25 and 35 mm (0.98 and 1.38 in) long. [2] Drones and workers are smaller in size than the queen. V. orientalis is a reddish-brown color and has distinctive thick yellow bands on the abdomen and yellow patches on the head between the eyes. It has very strong jaws and will bite if provoked. [2] Females (workers and the queen) have an ovipositor, which is a specialized organ shaped like a tube that is used for laying eggs. The ovipositor extends from the end of the abdomen and is also used as a stinger. [2] Males (drones) can be distinguished from workers by the number of segments on their antenna. Drones have 13 segments, while workers only have 12. [14] The Oriental hornet looks similar to the European hornet (V. crabro) and should not be confused with the Asian giant hornet (V. mandarinia) of East Asia. [14]

Distribution

Oriental hornets can be found in Central Asia, southwestern Asia from Armenia and Turkey to India and Nepal, throughout the Middle East, in Northeast Africa, in some Afrotropical countries such as Ethiopia and Somalia, and in parts of Southern Europe: Croatia, Bosnia, Montenegro, North Macedonia, Albania, Greece, Spain, Bulgaria, Cyprus, Malta, Sicily and the southern half of peninsular Italy. [2] [15] [16] Oriental hornets have been introduced by humans into additional locations, including Chile, [6] Madagascar, Mexico, and Xinjiang, China, as well as the occasional introduction via fruit into Belgium and the United Kingdom. [5] The Oriental hornet is the only member of the genus Vespa that can be found in desert climates such as those in North Africa, the Middle East, and parts of southwestern Asia. [14]

Nests

The Oriental hornet typically lives in nests that it digs underground. [8] On the roof base of each comb cell one or more mineral granules of polycrystalline material is attached. [17] A nest contains multiple combs in which the colony lives. [7] While nests are most commonly found under ground, some paper nests are constructed in protective hollows such as inside hollow trees, in shipping containers, parked vehicles, and aircraft. [2] To construct the paper nests, the workers strip the bark from twigs, tree branches, and shrubs to collect fiber. [2]

Colonies

Workers at the nest entrance are fanning their wings to cool down the nest on a hot midday. Vespa orientalis nest 1.jpg
Workers at the nest entrance are fanning their wings to cool down the nest on a hot midday.

Oriental hornets live in seasonal colonies that are formed every year in the spring by a single queen that mated during the previous fall. [2] During the fall, the queen of an established colony lays eggs which will develop into new queens and drones. After mating, the drones die off, while the newly fertilized queens seek hideouts in which to hibernate for the winter. [2] The newly founded colony grows throughout the spring and summer until the population and activity of the colony peaks in the late summer and early fall. [7] The peak size of the colony is several thousand individuals and a colony typically contains three to six combs, each containing 600-900 individual cells. [2]

Behavior

V. orientalis is a type of social wasp. Individuals live collectively in colonies with one queen and thousands of workers. Social wasps are unusual in their practice of altruism in which nonreproductive individuals work for the benefit of the colony. This occurs because all individuals in a colony are closely related.

Social structure

Within a colony, the caste system is dominated by the queen that is the only reproductive female in the colony. Subordinate to the queen are the workers (daughters) and drones (sons). [2] Like the queen, drones solely serve reproductive roles. The workers are responsible for the rest of the labor. Workers specialize in the performance of different tasks. Workers are responsible for foraging for food, providing shelter, defending the colony, and caring for the colony's brood, which consist of the queen's offspring. [2]

Communication

Oriental hornets communicate through sound vibrations. The three main types of vibrations used to communicate are taps to the queen, awakening taps, and larval hunger signals. [7] When workers tap while facing the queen, three effects are noted: the queen starts to search the combs for vacant cells in which to lay her eggs in, the workers go back to performing their typical duties, and hunger signals by larvae cease immediately and are not resumed for at least 30 minutes. [7] The main purpose of these taps seem to be to encourage the queen to lay more eggs. Awakening taps by workers cause a general intensification of activities in the colony. [7] The effects are minimal during the day. At night, however, the vibrations wake the whole colony, which causes the larvae commence their hunger signals and the workers to go forage for food to feed the larvae. [7] Larval hunger signals produce no detectable changes in larval activity during the daytime. The workers, however, pay more attention and give more food to the cells that are in the vicinity of where the vibrations originated. [7] At night, the larvae emulate each other's hunger signals and awake the whole nest. [7]

Kin selection and altruism

V. orientalis hornets live in colonies in which the workers are all daughters of the queen, so the workers are all sisters. Social wasps are haplodiploid, which means that males are all haploid and develop from unfertilized eggs, while females are diploid and develop from eggs fertilized by drones. Queens commonly mate with only one drone. Each drone has only one set of chromosomes to pass on to its offspring. Thus, sister workers that share paternity are unusually closely related to each other. Each received 100% its father's genes and 50% of its mother's genes, so each is on average 75% related to its sisters and only 50% related to the queen. Thus, a worker is benefited by looking after the queen and her colony to best ensure the survival of its genes. An individual acting in the interests of others and not just itself it is known as altruism.

Interactions with other species

Diet

Oriental hornets capture other insects such as grasshoppers, flies, honey bees, and vespids with which they feed the colony's brood. [2] They will also collect other animal proteins for their young such as pieces of fresh or spoiled meat and fish. [9] The adults eat carbohydrates such as nectar, honeydew, and fruits. [9]

Conflict with bees

The best place for hornets to find a combination of animal proteins (bees or larvae) and carbohydrates (honey) are bee hives. [9] Oriental hornets have been known to cause serious damage to bee colonies. [9] They are the primary pest that attacks honey bee colonies in many countries. [11]

Sting

Antibacterial effect of venom

The venom of V. orientalis was tested on Gram-positive bacteria, Staphylococcus aureus and Bacillus subtilis, and Gram-negative bacteria, Escherichia coli and Klebsiella pneumoniae. Venom extract was shown to be effective in inhibiting the growth of Gram-positive bacteria. The venom is therefore regarded as a potential therapeutic agent. [18]

Physiology

Solar energy harvesting

Unlike most wasps which are most active during the early morning, V. orientalis is unique in showing a peak of activity during the middle of the day. [8] Oriental hornets dig their nests underground by picking up soil in their mandibles, flying a short distance, dropping the soil, and returning to the nest to continue digging. [8] The hornet's digging is correlated with insolation (solar energy). The more insolation, the more active the hornet. [8] This daytime digging behavior of V. orientalis may be possible because of the ability of its cuticle to harvest solar energy. [8]

Cuticle

The cuticle of the Oriental hornet contains yellow pigment that protects the cuticle from harmful UV radiation from the sun. [8] The brown segments of the hornet's body contain melanin, which serves a similar purpose. The brown cuticle is composed of about 30 layers, the thickness of which increases from interior to exterior. [8] The brown melanin pigment is found in these layers. The yellow cuticle is composed of 15 layers that contain yellow pigment. The yellow stripe contains xanthopterin in barrel-shaped granules. [8]

Electric potential

Enzyme activity in the layer of yellow xanthopterin granules was demonstrated to be higher in hornets kept in dark environments and lower in those exposed to ultraviolet (UV) light. [8] Measuring the electric voltage between the hypocuticle and the exocuticle of the yellow stripe shows a negative electric potential at the hypocuticle with respect to the positive exocuticle. [8] If the yellow stripe is exposed to light, the potential difference between light and dark conditions increases. In dark conditions, the stripe decreases in electric potential. [8] Pterins have a role in photoreception and phototransduction of near-UV light to blue light and pterins may play a role in photosynthesis. [8] Some of the same researchers speculate that this is the energy for the heat organ [19] (see #Thermoregulatory organ below).

Also the silk has shown to have similar electric properties when exposed to light. [20]

Thermoregulatory organ

There is a thermoregulatory organ in the thorax of V. orientalis. [19] It maintains at 6 °C (11 °F) higher than the rest of the body, and 9 °C (16 °F) above ambient. [19] (Specifically it is located in the dorsal part of the thorax, inward from the meso-scutal plate of the prothorax. [19] This organ is near the median notal suture. [19] It is 1 millimetre (3⁄64 in) in diameter and butterfly-shaped. [19] The uppermost, hindmost side abuts the base of the forewings.) [19] The authors speculate that this heat generation is powered by the #Electric potential (above) that some of the same authors have previously discovered and investigated. [19]

Research and experiments

Israeli Space Agency

Face of an Oriental hornet, photo taken in Masada National Park, Israel Oriental Wasp - Face.jpg
Face of an Oriental hornet, photo taken in Masada National Park, Israel

The Israeli Space Agency Investigation About Hornets [21] (ISAIAH) was a project from Tel-Aviv University initiated in 1984 to explore the effects of near-zero gravity on oriental hornets, their development, and their nest-building instincts. The experiments were funded by the Israel Space Agency with the goal of discovering ways to prevent astronauts from suffering headaches, nausea, and vomiting during the missions. The sample of 230 Oriental hornets, flight hardware, and measuring instruments were all packed onto the Space Shuttle Endeavour, mission STS-47, in 1992. [21]

During the launch, 202 hornets died as a result of a malfunction in the water system that caused an abnormal increase in humidity. The surviving hornets lost their sense of direction, and unlike the control unit hornets, were unable to climb on the walls or stay in clusters. Instead, they stayed motionless and apart from each other. Roughly 3 to 4 days after returning to earth, the hornets started climbing on the walls again and building a nest. The surviving hornets lived for an average of 23 days, compared to an average survival of 43 days for the control group hornets. [22]

Related Research Articles

<span class="mw-page-title-main">Hornet</span> Genus of eusocial wasp

Hornets are the largest of the eusocial wasps, and are similar in appearance to their close relatives yellowjackets. Some species can reach up to 5.5 cm (2.2 in) in length. They are distinguished from other vespine wasps by the relatively large top margin of the head. Worldwide, 22 species of Vespa are recognized. Most species only occur in the tropics of Asia, though the European hornet, is widely distributed throughout Europe, Russia, North America, and north-eastern Asia. Wasps native to North America in the genus Dolichovespula are commonly referred to as hornets, but are actually yellowjackets.

<span class="mw-page-title-main">Vespidae</span> Family of insects

The Vespidae are a large, diverse, cosmopolitan family of wasps, including nearly all the known eusocial wasps and many solitary wasps. Each social wasp colony includes a queen and a number of female workers with varying degrees of sterility relative to the queen. In temperate social species, colonies usually last only one year, dying at the onset of winter. New queens and males (drones) are produced towards the end of the summer, and after mating, the queens hibernate over winter in cracks or other sheltered locations. The nests of most species are constructed out of mud, but polistines and vespines use plant fibers, chewed to form a sort of paper. Many species are pollen vectors contributing to the pollination of several plants, being potential or even effective pollinators, while others are notable predators of pest insect species.

<i>Dolichovespula maculata</i> Species of wasp

Dolichovespula maculata is a species of wasp in the genus Dolichovespula and a member of the eusocial, cosmopolitan family Vespidae. It is known by many colloquial names, primarily bald-faced hornet, but also including bald-faced aerial yellowjacket, bald-faced wasp, bald hornet, white-faced hornet, blackjacket, white-tailed hornet, spruce wasp, and bull wasp. Technically a species of yellowjacket wasp, it is not one of the true hornets, which are in the genus Vespa. Colonies contain 400 to 700 workers, the largest recorded colony size in its genus, Dolichovespula. It builds a characteristic large hanging paper nest up to 58 cm (23 in) in length. Workers aggressively defend their nest by repeatedly stinging invaders.

<span class="mw-page-title-main">Asian giant hornet</span> Predatory hornet, largest in the world

The Asian giant hornet or northern giant hornet, 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. However, by the end of the season in November 2022, there were no confirmed sightings in North America at all, suggesting the wasps may have been eradicated in that region.

<span class="mw-page-title-main">European hornet</span> Species of wasp

The European hornet is the largest eusocial wasp native to Europe. It is also the only true hornet found in North America, having been introduced to the United States and Canada from Europe as early as 1840. Vespines, such as V. crabro, are known for making intricate paper-like nests out of surrounding plant materials and other fibers. Unlike most other vespines, reproductive suppression involves worker policing instead of queen pheromone control, as was previously thought.

<span class="mw-page-title-main">Median wasp</span> Species of wasp

The median wasp is a species of social wasp of the family Vespidae found throughout Europe and Asia. It builds aerial paper nests often in shrubs or trees, and occasionally under the eaves of buildings. It is most common to see this wasp between May and October during its 3.3 month colony cycle. Behaviours of this wasp include nest defense, curling which is believed to function in brood incubation, and gastral vibration which is involved in larval feeding. The median wasp has a halplodiploid sex determination system that results in a high level of relatedness within the colony. This species is not usually aggressive but will sting if they feel their nest is threatened. Most foraging in the nest is done by the workers once the first ones reach adulthood. These workers forage for insects, nectar, and wood for nest construction in temperatures as low as 7 °C (45 °F). The median wasp is known to be occasionally affected by the fungus Cordyceps sphecocephala and the Cricket paralysis virus.

<i>Provespa</i> Genus of wasps

Provespa is a small genus of Vespidae, made up of nocturnal wasps from Southeast Asia, sometimes referred to as "night wasps" or "night hornets", though they are not true hornets. They are the only nocturnal members of the subfamily Vespinae, and also the only vespines where new colonies are formed by swarming. They tend to build their nests from fibrous plant material, making them a uniform greyish brown colour which is often difficult to locate.

<span class="mw-page-title-main">Eastern yellowjacket</span> Species of insect

The eastern yellow jacket or eastern yellowjacket is a wasp found in eastern North America. Although most of their nests are subterranean, they are often considered a pest due to their nesting in recreational areas and buildings. This yellow jacket is a social insect, living in colonies of hundreds to thousands of individuals. Along with their subfamily, Vespinae, this species demonstrates supportive parental care for offspring, separation of reproductive and sterile castes, and overlapping generations. They aggressively defend their hives from threats and are known to inflict painful stings.

<i>Apis cerana</i> Species of insect

Apis cerana, the eastern honey bee, Asiatic honey bee or Asian honey bee, is a species of honey bee native to South, Southeast and East Asia. This species is the sister species of Apis koschevnikovi and both are in the same subgenus as the western (European) honey bee, Apis mellifera. A. cerana is known to live sympatrically along with Apis koschevnikovi within the same geographic location. Apis cerana colonies are known for building nests consisting of multiple combs in cavities containing a small entrance, presumably for defense against invasion by individuals of another nest. The diet of this honey bee species consists mostly of pollen and nectar, or honey. Moreover, Apis cerana is known for its highly social behavior, reflective of its classification as a type of honey bee.

<i>Dolichovespula saxonica</i> Species of wasp

Dolichovespula saxonica, also known as the Saxon wasp, is a common social wasp found in the Palearctic region, specifically in large parts of Europe and in northern and central Asia. Although originally from continental Europe, D. saxonica has since colonised Britain, mainly in the south and east, but has been recorded as far north as East Lothian, Scotland. Most of their nests are above ground in trees and bushes, but they can also be found in buildings. Due to the proliferation of nests in urban areas and near residential homes, D. saxonica can be a pest for people. As a result, many human interventions are in place to remove Saxon nests. D. saxonica has been found to use chemical signaling in a lot of behaviours, such as alarm calls, fertility cues, and chemical trails.

<i>Dolichovespula norwegica</i> Species of wasp

The Norwegian wasp is a species of eusocial wasp. It is common in Scandinavia and can also be found in Scotland and other areas in Britain and Ireland. Often known for being a tree wasp, it nests in low branches and bushes and feeds on insects. It also obtains nectar from blueberry and snowberry flowers. Although D. norwegica is rarely considered a pest in the past, a few cases of pest problems relating to them have been reported. The species is not endangered.

<i>Dolichovespula sylvestris</i> Species of wasp

The tree wasp is a species of eusocial wasp in the family Vespidae, found in the temperate regions of Eurasia, particularly in western Europe. Despite being called the tree wasp, it builds both aerial and underground paper nests, and can be found in rural and urban habitats. D. sylvestris is a medium-sized wasp that has yellow and black stripes and a black dot in the center of its clypeus. It is most common to see this wasp between May and September during its 3.5 month colony cycle.

<i>Dolichovespula adulterina</i> Species of wasp

Dolichovespula adulterina is a species of parasitic social wasp found in the Palearctic region. D. adulterina feeds on a variety of foods, including insects, spiders, arthropods, meat, molluscs, fruit, nectar, and larval secretions. D. adulterina was formerly considered to be synonymous with D. arctica from the Holarctic region, but more recent research indicates that D. arctica is a separate species.

<i>Vespula rufa</i> Species of wasp

Vespula rufa, commonly known as the red wasp, is a social wasp species belonging to the genus Vespula. It is found in northern and central Europe, parts of Asia, and northern parts of North America. Vespula rufa is characterised by red-brown markings and body segmentation, with the appearance varying amongst the different roles of individuals in the species. These wasps build small nests in dry banks underground that are not far below the surface. The colony cycle begins in the fall. Vespula rufa feed on live insects. One interesting fact about Vespula rufa is that the queen policing occurs in the species, and that worker policing occurs at much lower rates than other species in the genus Vespula. There are predators and parasites of the species. The species goes through a series of events before leaving the nest.

<i>Vespula squamosa</i> Species of wasp

Vespula squamosa, or the southern yellowjacket, is a social wasp. This species can be identified by its distinctive black and yellow patterning and orange queen. This species is typically found in eastern North America, and its territory extends as far south as Central America. Within these territories, they create enormous, multiple-comb nests. The colonies may be either annual or perennial depending on the climate, and in many perennial nests, polygyny takes place. In addition, this species uses pheromones both as a sexual attractant and an alarm signal. This species feeds on insects and animal carcasses; it does not produce honey. V. squamosa, a social insect, has developed a parasitic relationship with the species V. vidua and V. maculifrons. Due to their painful, venomous stings, the species is considered a pest.

<i>Dolichovespula arenaria</i> Species of wasp

Dolichovespula arenaria, also known as the common aerial yellowjacket, sandhills hornet, and common yellow hornet, is a species of wasp within the genus Dolichovespula widely distributed in the North American continent.

<i>Polistes japonicus</i> Species of wasp

Polistes japonicus is a eusocial paper wasp found in Japan. It was first described by Henri Louis Frédéric de Saussure in 1858. It is closely related to Polistes formosanus. This species lives in small colonies with few workers and a foundress queen. Nests of these wasps are sometimes used as a traditional medicine in Korea, China, and Japan.

<i>Vespa affinis</i> Species of hornet

Vespa affinis, the lesser banded hornet, is a common hornet in tropical and subtropical Asia.

<i>Vespa soror</i> Species of hornet

Vespa soror, also known as the southern giant hornet, is a species of hornet present in India, Northern Thailand, Laos, Northern Vietnam, and parts of South China, including Hong Kong, Guangdong, Fujian, and Hainan Island.

Van der Vecht's gland or Van der Vecht's organ is a gland which is located in an area of modified cuticle on the rearmost gastral sternite of female wasps. This gland secretes chemicals which are important in the determination and maintenance of the hierarchy of groups of eusocial wasps and are used in the defence of the nests in others. In the Asian giant hornet the Van der Vecht's gland is used to scent mark hives of honey bees to attract other members of their colony to cooperatively attack the hive; the only known case of the gland's use to scent mark a food source. In the cleptoparasitic paper wasp Polistes semenowi the female usurps the host foundress, usually Polistes dominula and uses an enlarged Van der Vecht's gland to produce large quantities of hydrocarbons and to control the host workers, and even sometimes the host foundress. The gland was discovered by, and named in honour of, the Dutch entomologist Jacobus van der Vecht.

References

  1. ↑ International Union for Conservation of Nature and Natural Resources (2014). "The IUCN Red List of Threatened Species [Version 2014.2]". Archived from the original on June 27, 2014. Retrieved 25 September 2014.
  2. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 USAPHC Entomological Sciences Program. "Oriental hornets" (PDF). U.S. Army Public Health Command. Archived from the original (PDF) on 24 August 2014. Retrieved 24 September 2014.
  3. ↑ "Oriental hornet". Dieter Kosmeier. 13 June 2014. Retrieved 23 September 2014.
  4. ↑ Archer ME (1998). "Taxonomy, distribution and nesting biology of Vespa orientalis L. (Hym., Vespidae)". Entomologist's Monthly Magazine. 138: 45–51.
  5. 1 2 Dvorak, Libor (June 2006). "Oriental Hornet Vespa orientalis Linnaeus, 1771 found in Mexico" (PDF). Entomological Problems. 36: 80. Archived from the original (PDF) on 29 October 2013. Retrieved 24 September 2014.
  6. 1 2 Ríos, Mauro; Barrera, Roberto; Contreras, José (2020) Primer reporte del género Vespa Linnaeus (Hymenoptera: Vespidae: Vespinae) en Chile. Revista Chilena de Entomología 46:237-242. 10.35249/rche.46.2.20.14.
  7. 1 2 3 4 5 6 7 8 9 Ishay J, Motro A, GITTER S, Brown N (1974). "Rhythms in acoustical communication by the oriental hornet, vespa orientalis" (PDF). Animal Behaviour. 22 (3): 741–744. CiteSeerX   10.1.1.407.6956 . doi:10.1016/s0003-3472(74)80026-6 . Retrieved 25 September 2014.
  8. 1 2 3 4 5 6 7 8 9 10 11 12 13 Plotkin, Hod, Zaban; et al. (2010). "Solar energy harvesting in the epicuticle of the oriental hornet (Vespa orientalis)". Naturwissenschaften. 97 (12): 1067–1076. Bibcode:2010NW.....97.1067P. doi:10.1007/s00114-010-0728-1. PMID   21052618. S2CID   14022197.{{cite journal}}: CS1 maint: multiple names: authors list (link)
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  10. 1 2 Abdel-Ghany GM, Zalat SM, Abo-Ghalia AH, Semida FM (January 2009). "VARIATION OF VENOM AND THORACIC MUSCLE PROTEINS OF VESPA ORIENTALIS POPULATIONS IN RELATION TO GEOGRAPHICAL ISOLATION IN SOUTHERN SINAI PROTECTORATES, EGYPT" (PDF). Egyptian Journal of Natural Toxins. 6 (1): 16–32. Archived from the original (PDF) on 13 November 2014. Retrieved 25 September 2014.
  11. 1 2 Glaiim MK (2009). "HUNTING BEHAVIOR OF THE ORIENTAL HORNET, Vespa orientalis L., AND DEFENSE BEHAVIOR OF THE HONEY BEE, Apis mellifera L., IN IRAQ". Bull. Iraq Nat. Hist. Mus. 10 (4): 17–30.
  12. 1 2 Perrard A, Pickett KM, Villemant C, Kojima J, Carpenter J (24 April 2013). "Phylogeny of hornets: a total evidence approach (Hymenoptera, Vespidae, Vespinae, Vespa )". Journal of Hymenoptera Research. 32: 1–15. doi: 10.3897/jhr.32.4685 .
  13. ↑ A.H. Smith-Pardo, J.M. Carpenter, L. Kimsey (2020) The diversity of hornets in the genus Vespa (Hymenoptera: Vespidae; Vespinae), their importance and interceptions in the United States. Insect Systematics and Diversity 4(3) https://doi.org/10.1093/isd/ixaa006
  14. 1 2 3 The Global Biodiversity Information Facility (1 June 2013). "Vespa orientalis Linnaeus, 1771" . Retrieved 25 September 2014.
  15. ↑ Ćetković A. "A review of the European distribution of the Oriental hornet (Hymenoptera, Vespidae: Vespa orientalis)". Ekologija. 37: 1–22.
  16. ↑ Daglio A. On the Taxonomy and Distribution of the subfamily Vespinae. Lambert Academic Publishing, Beau Basin, v+49 pp.
  17. ↑ Ishay, J. S.; Joseph, Z.; Galushko, D.; Ermakov, N.; Bergman, D. J.; Barkay, Z.; Stokroos, I.; Van Der Want, J. (2005). "Perovskites in the comb roof base of hornets: Their possible function". Microscopy Research and Technique. 66 (5): 259–269. doi:10.1002/jemt.20168. PMID   15940685. S2CID   6818662.
  18. ↑ Jalaei, Jafar; Mehdi Fazeli; Hamid Rajaian; Seyed Shahram Shekarforoush (2014). "Investigation of Mating Preference for Nestmates in the Paper Wasp Polistes fuscatus (Hymenoptera: Vespidae)" (PDF). Journal of Venomous Animals and Toxins Including Tropical Diseases. 20 (22). Retrieved 29 September 2014.
  19. 1 2 3 4 5 6 7 8 Plotkin, Marian; Ermakov, Natalya Y.; Volynchik, Stanislav; Barkay, Zahava; Bergman, David J.; Ishay, Jacob S. (2005). "A thermoregulatory center in hornets: IR photography". Microscopy Research and Technique . Wiley Publishing. 68 (6): 321–328. doi:10.1002/jemt.20257. ISSN   1059-910X. PMID   16358280. S2CID   6710421.
  20. ↑ Hornet silk: thermophysical properties - ScienceDirect.com
  21. 1 2 "STS-47". NASA. Retrieved 20 November 2013.
  22. ↑ "The Hornet Experiment". IAMI. Archived from the original on 28 October 2013. Retrieved 20 November 2013.
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