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{{Short description|Species of wasp}} | ||
| {{Speciesbox | {{Speciesbox | ||
| | name = European horned what | |||
| | image = VespaCrabroCollezione.jpg | | image = VespaCrabroCollezione.jpg | ||
| | image_caption = Queen (top), worker (middle) and male (bottom) of the "germana" color form | | image_caption = Queen (top), worker (middle) and male (bottom) of the "germana" color form | ||
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| The '''European hornet''' (''Vespa crabro'') is the largest ] ] native to ]. It is also the only ] found in ], having been introduced |
The '''European hornet''' ('''''Vespa crabro''''') is the largest ] ] native to ]. It is also the only ] found in ], having been introduced to the United States and Canada from Europe as early as 1840.<ref name="Hornets">{{cite web |url= http://www.vespa-crabro.com/hornets.htm |title=Hornets: Gentle Giants |access-date=2014-09-30 |publisher=Dieter Kosmeier |year= 2013}}</ref><ref name="psu">{{Cite web |url=https://ento.psu.edu/extension/factsheets/european-hornet |title=European Hornet (Department of Entomology) |website=Department of Entomology (Penn State University) |language=en-us |access-date=2018-06-27}}</ref> Vespines, such as ''V. crabro'', are known for making intricate paper-like nests out of surrounding plant materials and other fibers.<ref name="nest">{{cite journal |author= Bagriaçik, N. |title=Determination of some structural features of the nest paper of ''Vespa orientalis'' Linneaus , 1771 and ''Vespa crabro'' Linneaus , 1758 (Hymenoptera: Vespinae) in Turkey |journal=Archives of Biological Sciences |year=2011 |volume=63 |issue=2 |pages=449–455 |doi=10.2298/ABS1102449B|doi-access=free }}</ref> Unlike most other vespines, reproductive suppression involves ] instead of queen pheromone control, as was previously thought. | ||
| This species stings in response to being stepped on or grabbed, but generally avoids conflict. It is also defensive of its nest and can be aggressive around food sources. European hornets are largely carnivorous and hunt large insects such as |
This species stings in response to being stepped on or grabbed, but generally avoids conflict. It is also defensive of its nest and can be aggressive around food sources. Care should be taken when they are found in these circumstances, as they may sting without warning. European hornets are largely carnivorous and hunt large insects such as ]s, ]s, large ]s, ], and ]es.<ref>{{dead link|fix-attempted=yes|date=April 2020}} goo</ref><ref> What's that bugs?</ref><ref> What's that bugs?</ref><ref> Education environnement, nature et patrimoine</ref><ref> cafe,daum</ref> They also feed on fallen fruit and other sources of sugary food.<ref name="psu" /> Mutual predation between medium-sized hornets and the ] (robber flies) is often reported. | ||
| ==Taxonomy== | ==Taxonomy== | ||
| The European hornet is a true |
The European hornet is a true hornet (genus ''Vespa''), a group characterized by eusocial species. The genus is in the subfamily ], members of which are known for chewing up their food to feed it to their young, as well as chewing up paper-like materials to make their nests. According to a recent phylogenetic study, its closest relative is '']''.<ref name="tree">{{cite journal |author1=Perrard, A. |author2=Pickett, K. M. |author3=Villemant, C. |author4=Kojima, J. |author5=Carpenter, J. |title=Phylogeny of hornets: a total evidence approach (Hymenoptera, Vespidae, Vespinae, Vespa) |journal=Journal of Hymenoptera Research |year=2013 |volume=32 |pages=1–15 |doi= 10.3897/JHR.32.4685|doi-access=free }}</ref> | ||
| ] | ] | ||
| The former subspecies of ''V. crabro'' (e.g.<ref>{{cite journal |author1=V. Dubatolov |author2=J. Kojima |author3=J. M. Carpenter |author4=A. Lvovsky |year=2003 |title=Subspecies of ''Vespa crabro'' in two different papers by Birula in 1925 |journal=Entomological Science |volume=6 |issue=2003 |pages=215–216 |doi=10.1046/j.1343-8786.2003.00037.x}}</ref><ref>{{cite journal |author1=J.M. Carpenter |author2=J. Kojima |year=1997 |title=Checklist of the species in the subfamily Vespinae (Insecta: Hymenoptera: Vespidae) |journal=Natural History Bulletin of Ibaraki University |volume=1 |issue=1997 |pages=51–92}}</ref>) are now considered to be geographic color forms; while |
The former subspecies of ''V. crabro'' (e.g.<ref>{{cite journal |author1=V. Dubatolov |author2=J. Kojima |author3=J. M. Carpenter |author4=A. Lvovsky |year=2003 |title=Subspecies of ''Vespa crabro'' in two different papers by Birula in 1925 |journal=Entomological Science |volume=6 |issue=2003 |pages=215–216 |doi=10.1046/j.1343-8786.2003.00037.x|s2cid=84172284 }}</ref><ref>{{cite journal |author1=J.M. Carpenter |author2=J. Kojima |year=1997 |title=Checklist of the species in the subfamily Vespinae (Insecta: Hymenoptera: Vespidae) |journal=Natural History Bulletin of Ibaraki University |volume=1 |issue=1997 |pages=51–92}}</ref>) are now considered to be geographic color forms; while a history exists of recognizing subspecies within many of the ''Vespa'' species, 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.<ref name="Vespa2020">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</ref> | ||
| ==Description== | ==Description== | ||
| The eyes of ''V. crabro'' are deeply indented and shaped like a "C". Its wings are reddish-orange, while the ] abdomen is striped with brown and yellow. It has hair on the thorax and abdomen, although the European hornet is not as hairy as most ]s.<ref>{{cite web |first=E. |last=Day |date=12 March 2015 |id=2911-1422 (ENTO-123NP) |publisher=Virginia Cooperative Extension |url=http://pubs.ext.vt.edu/2911/2911-1422/2911-1422.html |title=European Hornet}}</ref> Due to this coloration and abdomen pattern, ''V. crabro'' is often mistaken for the ]. Typical mass size for the European Hornet is {{val|477.5|59.9|u=mg}}.<ref name="size">{{cite journal |author1=Kovac, H. |author2=Stabentheiner, A. |title=Does size matter? – Thermoregulation of 'heavyweight' and 'lightweight' wasps (Vespa crabro and Vespula sp.) |journal=Biology Open |year=2012 |volume=1 |pages=848–856 |doi= 10.1242/bio.20121156 |pmc=3498068 |pmid=23162695}}</ref> Workers average around {{convert|25|mm|1|abbr=on}} in length, while the larger queens can reach up to {{convert|35|mm|1|abbr=on}} |
The eyes of ''V. crabro'' are deeply indented and shaped like a "C". Its wings are reddish-orange, while the ] abdomen is striped with brown and yellow. It has hair on the thorax and abdomen, although the European hornet is not as hairy as most ]s.<ref>{{cite web |first=E. |last=Day |date=12 March 2015 |id=2911-1422 (ENTO-123NP) |publisher=Virginia Cooperative Extension |url=http://pubs.ext.vt.edu/2911/2911-1422/2911-1422.html |title=European Hornet}}</ref> Due to this coloration and abdomen pattern, ''V. crabro'' is often mistaken for the ]. Typical mass size for the European Hornet is {{val|477.5|59.9|u=mg}}.<ref name="size">{{cite journal |author1=Kovac, H. |author2=Stabentheiner, A. |title=Does size matter? – Thermoregulation of 'heavyweight' and 'lightweight' wasps (Vespa crabro and Vespula sp.) |journal=Biology Open |year=2012 |volume=1 |issue=9 |pages=848–856 |doi= 10.1242/bio.20121156 |pmc=3498068 |pmid=23162695}}</ref> Workers average around {{convert|25|mm|1|abbr=on}} in length, while the larger queens can reach up to {{convert|35|mm|1|abbr=on}}.<ref name="psu" /> This is significantly larger than most common wasps (such as '']''), but smaller than the Asian giant hornet. Females are typically larger than males in both size and mass. However, male abdomens have seven segments, whereas female abdomens have six.<ref name="Hornets"/> There is a ] ], two ] ganglia, and five ] ganglia.<ref name="Anatomy-Catalogue" /> Only females possess a stinger, it is a modified egg laying device (ovipositor): males cannot sting. The antennae of males are slightly longer, with 13 segments compared to twelve segments in females.<ref name="Hornets"/> | ||
| ==Nests== | ==Nests== | ||
| Individuals typically live in paper nests, which consist of a pedicle (a paper comb on the inside), an envelope, and a single entry hole on the outside. Materials such as twigs, branches, and other available plant resources are broken up, chewed, and shaped into a nest by the workers. These pieces are not uniform in shape, but are glued together very closely. Because social wasps generally prefer to build nests in the dark, envelopes are commonly found surrounding the nests to make them dark if the colony could not locate a dark crevice in which to build.<ref name="nest"/> | Individuals typically live in paper nests, which consist of a pedicle (a paper comb on the inside), an envelope, and a single entry hole on the outside. Materials such as twigs, branches, and other available plant resources are broken up, chewed, and shaped into a nest by the workers. These pieces are not uniform in shape, but are glued together very closely. Because social wasps generally prefer to build nests in the dark, envelopes are commonly found surrounding the nests to make them dark if the colony could not locate a dark crevice in which to build.<ref name="nest"/> The ] '']'' is ] with the European hornet, living much of its life in the detritus of the nest.<ref name = ukb>{{cite web | url = https://www.ukbeetles.co.uk/velleius-dilatatus | title = Quedius dilatatus (Fabricius, 1787) Hornet rove beetle | access-date = 19 February 2023 | publisher = UK Beetles}}</ref> | ||
| ] | ] | ||
| ===Construction=== | ===Construction=== | ||
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| ===Physical and chemical composition=== | ===Physical and chemical composition=== | ||
| Minerals such as ], ], and ] are commonly found in the soil and they, too, become part of the comb walls. The average dry weight of the nest is about {{convert|80.87|g}}. Cells in the paper comb are typically {{convert|8|-|9|mm}} long and {{convert|4|-|5|mm}} in diameter. Analysis of the composition of nests in northern Turkey revealed oxygen, carbon, and nitrogen as the main elements, while trace amounts of ], ], iron, and ] were found, although none of ], ], or ], providing evidence that European hornets use the surrounding soil as a resource in building their nests. The ratio of fibrous material to actual saliva affects the nest's ability to absorb water, and thus how well its inside stays dry. In the nests studied in Turkey, fiber content was 23%, with 77% hornet saliva. This combination resulted in optimal water absorption capacity.<ref name="nest"/> | Minerals such as ], ], and ] are commonly found in the soil and they, too, become part of the comb walls. The average dry weight of the nest is about {{convert|80.87|g}}. Cells in the paper comb are typically {{convert|8|-|9|mm}} long and {{convert|4|-|5|mm}} in diameter. Analysis of the composition of nests in northern Turkey revealed oxygen, carbon, and nitrogen as the main elements, while trace amounts of ], ], iron, and ] were found, although none of ], ], or ], providing evidence that European hornets use the surrounding soil as a resource in building their nests. The ratio of fibrous material to actual saliva affects the nest's ability to absorb water, and thus how well its inside stays dry. In the nests studied in ], fiber content was 23%, with 77% hornet saliva. This combination resulted in optimal water absorption capacity.<ref name="nest"/> | ||
| ==Distribution== | ==Distribution== | ||
| The European hornet ''V. crabro'' originated in Eurasia. Nests historically ranged from ] to the ]. However, ] reported that ''V. crabro'' was introduced to North America in the mid-19th century, where it is now well established. In 2010, they were found as far south as ]. The few nests in Guatemala were thought to be recent, accidental introductions, as they were the first to be documented.<ref>{{cite journal |last1=Landolt |first1=P. J. |last2=Sierra |first2=J. M. |last3=Unruh |first3=T. R. |last4=Zack |first4=R. S. |title=A new species of ''Vespula'', and the first record of ''Vespa crabro'' L.(Hymenoptera: Vespidae) from Guatemala, Central America| journal=Zootaxa |year=2010 |issue=1 |volume=2629 |pages=61–68 |doi=10.11646/zootaxa.2629.1.4|doi-access=free }}</ref> | |||
| ==Life cycle== | ==Life cycle== | ||
| ] | ] | ||
| ''V. crabro'' prefers to build nests in dark places, usually hollow tree trunks. After the site has been chosen, the queen lays eggs in the combs inside the nest. The workers dispose of any eggs that are not laid by their queen; this behavior is called worker policing. Based on laboratory data, the average rate of egg-laying is 2.31 eggs per day. However, in this same nest, cell construction rate was only 1.63 cells per day.<ref>{{cite journal |last=Archer |first=M.E. |title=The queen colony phase of vespine wasps (Hymenoptera, Vespidae)| journal=Insectes Sociaux|year=2010 |volume=57 |pages=133–145 |doi= 10.1007/s00040-009-0063-8}}</ref> | ''V. crabro'' prefers to build nests in dark places, usually hollow tree trunks. After the site has been chosen, the queen lays eggs in the combs inside the nest. The workers dispose of any eggs that are not laid by their queen; this behavior is called worker policing. Based on laboratory data, the average rate of egg-laying is 2.31 eggs per day. However, in this same nest, cell construction rate was only 1.63 cells per day.<ref>{{cite journal |last=Archer |first=M.E. |title=The queen colony phase of vespine wasps (Hymenoptera, Vespidae)| journal=Insectes Sociaux|year=2010 |volume=57 |issue=2 |pages=133–145 |doi= 10.1007/s00040-009-0063-8|s2cid=26973034 }}</ref> | ||
| ''V. crabro'' colonies seasonally change strategies of obtaining food for both the larvae and adults. In April, when the queen normally lays her eggs, the workers actively go out and forage. Later, typically around the fall season, the foraging workers switch to scavengers. Instead of putting forth the effort to catch food sources, the workers try to take what is more easily available. For example, European hornets have been seen hovering around garbage cans and picnic areas in the fall.<ref name="rearing">{{cite journal |author1=Hoffmann, W.R.E. |author2=Neumann, P. |author3=Schmolz, E. |title=Technique for rearing the European hornet (''Vespa crabro'') through an entire colony life cycle in captivity| journal=Insectes Sociaux| year=2000 | volume=47 |issue=4 |pages=351–3 |doi= 10.1007/PL00001729}}</ref> | ''V. crabro'' colonies seasonally change strategies of obtaining food for both the larvae and adults. In April, when the queen normally lays her eggs, the workers actively go out and forage. Later, typically around the fall season, the foraging workers switch to scavengers. Instead of putting forth the effort to catch food sources, the workers try to take what is more easily available. For example, European hornets have been seen hovering around garbage cans and picnic areas in the fall.<ref name="rearing">{{cite journal |author1=Hoffmann, W.R.E. |author2=Neumann, P. |author3=Schmolz, E. |title=Technique for rearing the European hornet (''Vespa crabro'') through an entire colony life cycle in captivity| journal=Insectes Sociaux| year=2000 | volume=47 |issue=4 |pages=351–3 |doi= 10.1007/PL00001729|s2cid=17378825 }}</ref> | ||
| ==Worker policing== | ==Worker policing== | ||
| Very few workers lay eggs in healthy ''V. crabro'' colonies. While only queens mate and produce fertilized female eggs, workers are capable of laying ] male eggs. Workers are generally more closely related to male offspring of other workers than to male offspring of the queen.<ref name="social">{{cite journal |author=Hamilton, W. D. |title=The genetical evolution of social behaviour. I |journal=Journal of Theoretical Biology |year=1964 |volume=7 |issue=1 |pages=1–16 |doi= 10.1016/0022-5193(64)90038-4 |pmid=5875341}}</ref> Workers would have reproductive benefits from laying male eggs, but do not do so.<ref name="police">{{cite journal | |
Very few workers lay eggs in healthy ''V. crabro'' colonies. While only queens mate and produce fertilized female eggs, workers are capable of laying ] male eggs. Workers are generally more closely related to male offspring of other workers than to male offspring of the queen.<ref name="social">{{cite journal |author=Hamilton, W. D. |title=The genetical evolution of social behaviour. I |journal=Journal of Theoretical Biology |year=1964 |volume=7 |issue=1 |pages=1–16 |doi= 10.1016/0022-5193(64)90038-4 |pmid=5875341|bibcode=1964JThBi...7....1H }}</ref> Workers would have reproductive benefits from laying male eggs, but do not do so.<ref name="police">{{cite journal |last1=Foster |first1=K.R. |last2=Gulliver |first2=J. |last3=Ratnieks |first3=F.L.W. |title=Worker policing in the European hornet ''Vespa crabro'' |journal=Insectes Sociaux |year=2002 |volume=49 |issue=1 |pages=41–44 |doi=10.1007/s00040-002-8277-z|s2cid=45182961 }}</ref> This was originally thought to be caused by ] control by the queen, but new evidence has shown that this is not the case. Workers enforce sterility on one another in a strategy known as ]. Workers either physically destroy worker-laid eggs or discriminate against those workers that attempt to lay eggs. Ensuring that only the queen reproduces is thought to be likely to benefit the organization and productivity of the colony as a whole, by reducing conflicts.<ref name="police" /> | ||
| ==Alarm behavior== | ==Alarm behavior== | ||
| Social hymenopteran species typically communicate with each other through behaviors or pheromones. In the European hornet, a typical alarm dance is performed outside of the nest and consists of consistent buzzing, darting in and out of the nest, and attacking or approaching the target of the alarm pheromone. The alarm pheromone is stored in, and secreted from, internal venom sacs. 2-Methyl-3-butene-2-ol is the main pheromone component |
Social hymenopteran species typically communicate with each other through behaviors or pheromones. In the European hornet, a typical alarm dance is performed outside of the nest and consists of consistent buzzing, darting in and out of the nest, and attacking or approaching the target of the alarm pheromone. The alarm pheromone is stored in, and secreted from, internal venom sacs. 2-Methyl-3-butene-2-ol is the main pheromone component that causes ''V. crabro'' to express this defensive behavior. Other pentenols and pentanols are contained within these venomous sacs, but their primary purpose is likely not to warn fellow hornets that danger is nearby, because these chemicals do not induce alarm behavior.<ref name="alarm">{{cite journal |last=Veith |first=H.J. |title=2-Methyl-3-butene-2-ol, a major component of the alarm pheromone of the hornet ''Vespa crabro'' |journal=Naturwissenschaften |year=1984 |volume=71 |issue=6 |pages=328–329 |doi=10.1007/BF00396622 |bibcode=1984NW.....71..328V|s2cid=1629303 }}</ref> | ||
| ==Kleptoparasitism== | ==Kleptoparasitism== | ||
| European hornets have been observed to steal prey from ]s, which can be classified as an example of ]. This behavior was first documented in 2011 against a yellow garden spider, '']''. A European hornet flew into the spider’s web and appeared entangled. The hornet cut free a captured insect (possibly a ] wasp), wrapped in silk, from the spider’s web. The spider did not attack or interfere with ''V. crabro'' while it was stealing its prey.<ref name="steal"/> This behavior follows the pattern of most vespines' changing their foraging techniques from hunting to scavenging, especially once the |
European hornets have been observed to steal prey from ]s, which can be classified as an example of ]. This behavior was first documented in 2011 against a yellow garden spider, '']''. A European hornet flew into the spider’s web and appeared entangled. The hornet cut free a captured insect (possibly a ] wasp), wrapped in silk, from the spider’s web. The spider did not attack or interfere with ''V. crabro'' while it was stealing its prey.<ref name="steal">{{cite journal |author=Davis, M.| title=A hornet (''Vespa crabro'') steals prey from a spider (''Argiope aurantia'') |journal=Southeastern Naturalist |year=2011 |volume=10 |issue=1 |pages=191–192 |doi=10.1656/058.010.0119| s2cid=86068392 }}</ref> This behavior follows the pattern of most vespines' changing their foraging techniques from hunting to scavenging, especially once the autumn season begins. | ||
| ''V. crabro ''has also been observed attacking '']'' nests.<ref>{{cite journal | |
''V. crabro ''has also been observed attacking '']'' nests.<ref>{{cite journal |last1=Rusina |first1=L. Yu. |first2=E. S. |last2=Orlova |title=The relationship between phenotypic variability in future foundresses of ''Polistes Nimpha'' (Christ) (Hymenoptera, Vespidae, Polistinae) and infestation of their larvae by the mite ''Sphexicozela Connivens'' Mahunka (Acari, Astigmata, Winterschmidtiidae) |journal=Entomological Review |volume=91 |issue=6 |pages=685–691 |year=2011 |doi=10.1134/S0013873811060017 |s2cid=1999452 }} | ||
| </ref> | </ref> | ||
| ==Relationship with humans== | ==Relationship with humans== | ||
| ] | ] | ||
| ===Endangered species and legal protection=== | ===Endangered species and legal protection=== | ||
| Unwarranted fear of ''V. crabro'' has often led to the destruction of nests. This has led to the decline of the species, which is often locally threatened or even endangered. European hornets benefit from legal protection in some countries, notably ], where killing a European hornet or nest has been illegal since January 1, 1987, with a fine up to €50,000.<ref>{{cite web |url=http://www.vespa-crabro.de/hornets3.htm |title=Hornets are worthy of protection!}}</ref> | Unwarranted fear of ''V. crabro'' has often led to the destruction of nests. This has led to the decline of the species, which is often locally threatened or even endangered. European hornets benefit from legal protection in some countries, notably ], where killing a European hornet or nest has been illegal since January 1, 1987, with a fine up to €50,000.<ref name="Hornets are worthy of protection!">{{cite web |url=http://www.vespa-crabro.de/hornets3.htm |title=Hornets are worthy of protection!}}</ref> However, the highest reported fine levied in Germany for killing of wasps was €45, and experts report that fines are rarely imposed.<ref>{{cite web |url=https://medium.com/@sophanemwise/the-wasp-killing-myth-in-germany-2ef995c50451 |title=The Wasp-Killing Myth in Germany|date=25 August 2018}}</ref><ref>{{cite web |url=https://correctiv.org/faktencheck/2018/07/24/nein-wer-eine-einzige-wespe-toetet-muss-nicht-50000-euro-bussgeld-zahlen |title=Nein – Wer eine einzige Wespe tötet muss nicht 50.000 Euro ($51,942.25 USD) Bußgeld zahlen | ||
| |date=24 July 2018 | |||
| }}</ref> | |||
| ===Problems associated=== | ===Problems associated=== | ||
| European hornets |
European hornets hunt many species of insects to feed their larvae. While many of these insects are considered to be garden pests, European hornets also prey on ]s (''Apis mellifera''). However, they do not typically pose a threat to healthy western honey bee colonies, unlike the ] or ], which are invasive in some parts of the world.<ref>{{cite web|last1=Rortais |first1=A |last2=Villemant |first2=CL |last3=Gargominy |first3=OL |last4=Rome |first4=QU |last5=Haxaire |first5=JE |last6=Papachristoforou |first6=AL |last7=Arnold |first7=GÉ. |title=A new enemy of honeybees in Europe: The Asian hornet Vespa velutina. Atlas of Biodiversity Risks–from Europe to globe, from stories to maps. |publisher=Sofia & Moscow: Pensoft. |date=2010 |url = https://www.researchgate.net/publication/224842243}}</ref> Some ]s in Europe may keep hornets and their nests in specially designed boxes hanging on trees, on the premise that the protection they provide against ]s is worth the occasional bee they may prey upon.<ref name="Hornets are worthy of protection!"/> European hornets tend to ] branches, which results in dead branches.<ref>{{cite web |id=ENTFACT-600 |first1=D. |last1=Johnson |first2=L. |last2=Townsend |title=European Hornet in Kentucky |date=May 2016 |publisher=University of Kentucky |url=https://entomology.ca.uky.edu/ef600}}</ref> | ||
| ===Stings: case study=== | ===Stings: case study=== | ||
| Most cases of stings from ''V. crabro'' do not require medical attention, but rarely can be serious. A documented case requiring treatment displayed symptoms including tingling at the site of the sting, |
Most cases of stings from ''V. crabro'' do not require medical attention, but rarely can be serious. A documented case requiring treatment displayed symptoms including tingling at the site of the sting, headaches, and shortness of breath. In the hospital, the victim was found to have a fast, irregular heartbeat with a ] of 111/63. A subsequent EKG demonstrated ] with a rapid ventricular response. ''V. crabro'' venom contains ]s such as ], ], ] and noradrenalineneurotoxin apamin, as well as enzymes ] and ], and proteins ] and ]. These compounds have been shown to cause episodes of ] in smaller animals. The mechanism of the described attack is still undetermined, but the victim was possibly abnormally susceptible to vespine stings. Currently, the two most effective treatments for reactions are ] or ]. The victim in this case study was given an oral dose of propafenone (150 mg) and his atrial fibrillation resolved.<ref name="sting">{{cite journal |author1=Okutucu, S. |author2=Şabanov, C. |author3=Abdulhayoğlu, E. |author4=Aksu, N. M. |author5=Erbil, B. |author6=Aytemir, K. |author7=Özkutlu, H. |title=A rare cause of atrial fibrillation: a European hornet sting |journal=] |year=2011 |volume=11 |issue=6 |pages=559–560 |doi=10.5152/akd.2011.144|pmid=21827996 |doi-access=free }}</ref> | ||
| ==Gallery== | ==Gallery== | ||
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| ==See also== | ==See also== | ||
| *] | *] | ||
| *]s | |||
| *] | *] | ||
| {{Clear}} | {{Clear}} | ||
| ==Notes== | ==Notes== | ||
| {{Reflist|30em |
{{Reflist|30em|refs= | ||
| <ref name="Anatomy-Catalogue">{{cite book|url=https://books.google.com/books?id=OftGAQAAMAAJ&pg=PA41|publisher=]|location=Red Lion Court, ], ]|date=1902|volume=2|edition=2nd|title=Descriptive and Illustrated Catalogue of the Physiological Series of Comparative Anatomy contained in The Museum of the Royal College of Surgeons of England|pages=41–42}}</ref> | |||
| }} | |||
| ==External links== | ==External links== | ||
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| {{Taxonbar|from=Q30258}} | {{Taxonbar|from=Q30258}} | ||
| {{Authority control}} | |||
| ] | ] | ||
| ] | ] | ||
| ] | ] | ||
| ] | ] | ||
| ] | ] | ||
| ] | ] | ||
Latest revision as of 21:31, 1 December 2024
Species of wasp
| European hornet | |
|---|---|
| |
| Queen (top), worker (middle) and male (bottom) of the "germana" color form | |
Scientific classification 
| |
| Domain: | Eukaryota |
| Kingdom: | Animalia |
| Phylum: | Arthropoda |
| Class: | Insecta |
| Order: | Hymenoptera |
| Family: | Vespidae |
| Genus: | Vespa |
| Species: | V. crabro |
| Binomial name | |
| Vespa crabro Linnaeus, 1758 | |
The European hornet (Vespa crabro) is the largest eusocial wasp native to Europe. It is also the only true hornet (genus Vespa) 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.
This species stings in response to being stepped on or grabbed, but generally avoids conflict. It is also defensive of its nest and can be aggressive around food sources. Care should be taken when they are found in these circumstances, as they may sting without warning. European hornets are largely carnivorous and hunt large insects such as beetles, wasps, large moths, dragonflies, and mantises. They also feed on fallen fruit and other sources of sugary food. Mutual predation between medium-sized hornets and the Asilidae (robber flies) is often reported.
Taxonomy
The European hornet is a true hornet (genus Vespa), a group characterized by eusocial species. The genus is in the subfamily Vespinae, members of which are known for chewing up their food to feed it to their young, as well as chewing up paper-like materials to make their nests. According to a recent phylogenetic study, its closest relative is Vespa dybowskii.
The former subspecies of V. crabro (e.g.) are now considered to be geographic color forms; while a history exists of recognizing subspecies within many of the Vespa species, 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.
Description
The eyes of V. crabro are deeply indented and shaped like a "C". Its wings are reddish-orange, while the petiolate abdomen is striped with brown and yellow. It has hair on the thorax and abdomen, although the European hornet is not as hairy as most bees. Due to this coloration and abdomen pattern, V. crabro is often mistaken for the Asian giant hornet. Typical mass size for the European Hornet is 477.5±59.9 mg. Workers average around 25 mm (1.0 in) in length, while the larger queens can reach up to 35 mm (1.4 in). This is significantly larger than most common wasps (such as Vespula vulgaris), but smaller than the Asian giant hornet. Females are typically larger than males in both size and mass. However, male abdomens have seven segments, whereas female abdomens have six. There is a cerebral ganglion, two thoracic ganglia, and five abdominal ganglia. Only females possess a stinger, it is a modified egg laying device (ovipositor): males cannot sting. The antennae of males are slightly longer, with 13 segments compared to twelve segments in females.
Nests
Individuals typically live in paper nests, which consist of a pedicle (a paper comb on the inside), an envelope, and a single entry hole on the outside. Materials such as twigs, branches, and other available plant resources are broken up, chewed, and shaped into a nest by the workers. These pieces are not uniform in shape, but are glued together very closely. Because social wasps generally prefer to build nests in the dark, envelopes are commonly found surrounding the nests to make them dark if the colony could not locate a dark crevice in which to build. The rove beetle Velleius dilatatus is commensal with the European hornet, living much of its life in the detritus of the nest.
Construction
The nest is composed of a paper-pulp mixture created by female workers chewing up dead bark, trees, or plant matter from nearby surroundings and mixing it with their saliva. To build the actual comb, saliva is used as a cement to piece together organic and inorganic materials that are readily available to the colony. This cement not only holds together the comb, but also protects the comb from being damaged by water. It provides a protective barrier to help protect the colony from wind or other harsh weather conditions. Available resources, location, and amount of mastication affect the final nest's appearance, so much variation is seen among the nests of V. crabro.
Physical and chemical composition
Minerals such as titanium, iron, and zirconium are commonly found in the soil and they, too, become part of the comb walls. The average dry weight of the nest is about 80.87 grams (2.853 oz). Cells in the paper comb are typically 8–9 millimetres (0.31–0.35 in) long and 4–5 millimetres (0.16–0.20 in) in diameter. Analysis of the composition of nests in northern Turkey revealed oxygen, carbon, and nitrogen as the main elements, while trace amounts of silicon, calcium, iron, and potassium were found, although none of aluminium, magnesium, or sodium, providing evidence that European hornets use the surrounding soil as a resource in building their nests. The ratio of fibrous material to actual saliva affects the nest's ability to absorb water, and thus how well its inside stays dry. In the nests studied in Turkey, fiber content was 23%, with 77% hornet saliva. This combination resulted in optimal water absorption capacity.
Distribution
The European hornet V. crabro originated in Eurasia. Nests historically ranged from Japan to the United Kingdom. However, Saussure reported that V. crabro was introduced to North America in the mid-19th century, where it is now well established. In 2010, they were found as far south as Guatemala. The few nests in Guatemala were thought to be recent, accidental introductions, as they were the first to be documented.
Life cycle
V. crabro prefers to build nests in dark places, usually hollow tree trunks. After the site has been chosen, the queen lays eggs in the combs inside the nest. The workers dispose of any eggs that are not laid by their queen; this behavior is called worker policing. Based on laboratory data, the average rate of egg-laying is 2.31 eggs per day. However, in this same nest, cell construction rate was only 1.63 cells per day.
V. crabro colonies seasonally change strategies of obtaining food for both the larvae and adults. In April, when the queen normally lays her eggs, the workers actively go out and forage. Later, typically around the fall season, the foraging workers switch to scavengers. Instead of putting forth the effort to catch food sources, the workers try to take what is more easily available. For example, European hornets have been seen hovering around garbage cans and picnic areas in the fall.
Worker policing
Very few workers lay eggs in healthy V. crabro colonies. While only queens mate and produce fertilized female eggs, workers are capable of laying haploid male eggs. Workers are generally more closely related to male offspring of other workers than to male offspring of the queen. Workers would have reproductive benefits from laying male eggs, but do not do so. This was originally thought to be caused by pheromone control by the queen, but new evidence has shown that this is not the case. Workers enforce sterility on one another in a strategy known as worker policing. Workers either physically destroy worker-laid eggs or discriminate against those workers that attempt to lay eggs. Ensuring that only the queen reproduces is thought to be likely to benefit the organization and productivity of the colony as a whole, by reducing conflicts.
Alarm behavior
Social hymenopteran species typically communicate with each other through behaviors or pheromones. In the European hornet, a typical alarm dance is performed outside of the nest and consists of consistent buzzing, darting in and out of the nest, and attacking or approaching the target of the alarm pheromone. The alarm pheromone is stored in, and secreted from, internal venom sacs. 2-Methyl-3-butene-2-ol is the main pheromone component that causes V. crabro to express this defensive behavior. Other pentenols and pentanols are contained within these venomous sacs, but their primary purpose is likely not to warn fellow hornets that danger is nearby, because these chemicals do not induce alarm behavior.
Kleptoparasitism
European hornets have been observed to steal prey from spiders, which can be classified as an example of kleptoparasitism. This behavior was first documented in 2011 against a yellow garden spider, Argiope aurantia. A European hornet flew into the spider’s web and appeared entangled. The hornet cut free a captured insect (possibly a eumenine wasp), wrapped in silk, from the spider’s web. The spider did not attack or interfere with V. crabro while it was stealing its prey. This behavior follows the pattern of most vespines' changing their foraging techniques from hunting to scavenging, especially once the autumn season begins.
V. crabro has also been observed attacking Polistes nimpha nests.
Relationship with humans
Endangered species and legal protection
Unwarranted fear of V. crabro has often led to the destruction of nests. This has led to the decline of the species, which is often locally threatened or even endangered. European hornets benefit from legal protection in some countries, notably Germany, where killing a European hornet or nest has been illegal since January 1, 1987, with a fine up to €50,000. However, the highest reported fine levied in Germany for killing of wasps was €45, and experts report that fines are rarely imposed.
Problems associated
European hornets hunt many species of insects to feed their larvae. While many of these insects are considered to be garden pests, European hornets also prey on western honey bees (Apis mellifera). However, they do not typically pose a threat to healthy western honey bee colonies, unlike the Asian hornet or Asian giant hornet, which are invasive in some parts of the world. Some beekeepers in Europe may keep hornets and their nests in specially designed boxes hanging on trees, on the premise that the protection they provide against wax moths is worth the occasional bee they may prey upon. European hornets tend to girdle branches, which results in dead branches.
Stings: case study
Most cases of stings from V. crabro do not require medical attention, but rarely can be serious. A documented case requiring treatment displayed symptoms including tingling at the site of the sting, headaches, and shortness of breath. In the hospital, the victim was found to have a fast, irregular heartbeat with a blood pressure of 111/63. A subsequent EKG demonstrated atrial fibrillation with a rapid ventricular response. V. crabro venom contains neurotransmitters such as dopamine, serotonin, histamine and noradrenalineneurotoxin apamin, as well as enzymes phospholipase A and hyaluronidase, and proteins melittin and bradykinin. These compounds have been shown to cause episodes of tachycardia in smaller animals. The mechanism of the described attack is still undetermined, but the victim was possibly abnormally susceptible to vespine stings. Currently, the two most effective treatments for reactions are electrical cardioversion or propafenone. The victim in this case study was given an oral dose of propafenone (150 mg) and his atrial fibrillation resolved.
Gallery
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Vespa crabro face
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Female V. crabro dorsal view
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Male V. crabro ventral view
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Hornets' nest in hollow tree
- Video of European hornets on their nest
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European hornet
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See also
Notes
- ^ "Hornets: Gentle Giants". Dieter Kosmeier. 2013. Retrieved 2014-09-30.
- ^ "European Hornet (Department of Entomology)". Department of Entomology (Penn State University). Retrieved 2018-06-27.
- ^ Bagriaçik, N. (2011). "Determination of some structural features of the nest paper of Vespa orientalis Linneaus [sic], 1771 and Vespa crabro Linneaus [sic], 1758 (Hymenoptera: Vespinae) in Turkey". Archives of Biological Sciences. 63 (2): 449–455. doi:10.2298/ABS1102449B.
- モンスズメバチ(vespa crabro flavofasciata) goo
- European Hornet Kills Dragonfly What's that bugs?
- European Hornet eats Swamp Darner What's that bugs?
- insecte : frelon mangeant une mante religieuse Education environnement, nature et patrimoine
- 사마귀와 말벌의 사투(vespa crabro flavofasciata) cafe,daum
- Perrard, A.; Pickett, K. M.; Villemant, C.; Kojima, J.; Carpenter, J. (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.
- V. Dubatolov; J. Kojima; J. M. Carpenter; A. Lvovsky (2003). "Subspecies of Vespa crabro in two different papers by Birula in 1925". Entomological Science. 6 (2003): 215–216. doi:10.1046/j.1343-8786.2003.00037.x. S2CID 84172284.
- J.M. Carpenter; J. Kojima (1997). "Checklist of the species in the subfamily Vespinae (Insecta: Hymenoptera: Vespidae)". Natural History Bulletin of Ibaraki University. 1 (1997): 51–92.
- 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
- Day, E. (12 March 2015). "European Hornet". Virginia Cooperative Extension. 2911-1422 (ENTO-123NP).
- Kovac, H.; Stabentheiner, A. (2012). "Does size matter? – Thermoregulation of 'heavyweight' and 'lightweight' wasps (Vespa crabro and Vespula sp.)". Biology Open. 1 (9): 848–856. doi:10.1242/bio.20121156. PMC 3498068. PMID 23162695.
- Descriptive and Illustrated Catalogue of the Physiological Series of Comparative Anatomy contained in The Museum of the Royal College of Surgeons of England. Vol. 2 (2nd ed.). Red Lion Court, Fleet Street, London: Taylor & Francis. 1902. pp. 41–42.
- "Quedius dilatatus (Fabricius, 1787) Hornet rove beetle". UK Beetles. Retrieved 19 February 2023.
- Landolt, P. J.; Sierra, J. M.; Unruh, T. R.; Zack, R. S. (2010). "A new species of Vespula, and the first record of Vespa crabro L.(Hymenoptera: Vespidae) from Guatemala, Central America". Zootaxa. 2629 (1): 61–68. doi:10.11646/zootaxa.2629.1.4.
- Archer, M.E. (2010). "The queen colony phase of vespine wasps (Hymenoptera, Vespidae)". Insectes Sociaux. 57 (2): 133–145. doi:10.1007/s00040-009-0063-8. S2CID 26973034.
- Hoffmann, W.R.E.; Neumann, P.; Schmolz, E. (2000). "Technique for rearing the European hornet (Vespa crabro) through an entire colony life cycle in captivity". Insectes Sociaux. 47 (4): 351–3. doi:10.1007/PL00001729. S2CID 17378825.
- Hamilton, W. D. (1964). "The genetical evolution of social behaviour. I". Journal of Theoretical Biology. 7 (1): 1–16. Bibcode:1964JThBi...7....1H. doi:10.1016/0022-5193(64)90038-4. PMID 5875341.
- ^ Foster, K.R.; Gulliver, J.; Ratnieks, F.L.W. (2002). "Worker policing in the European hornet Vespa crabro". Insectes Sociaux. 49 (1): 41–44. doi:10.1007/s00040-002-8277-z. S2CID 45182961.
- Veith, H.J. (1984). "2-Methyl-3-butene-2-ol, a major component of the alarm pheromone of the hornet Vespa crabro". Naturwissenschaften. 71 (6): 328–329. Bibcode:1984NW.....71..328V. doi:10.1007/BF00396622. S2CID 1629303.
- Davis, M. (2011). "A hornet (Vespa crabro) steals prey from a spider (Argiope aurantia)". Southeastern Naturalist. 10 (1): 191–192. doi:10.1656/058.010.0119. S2CID 86068392.
- Rusina, L. Yu.; Orlova, E. S. (2011). "The relationship between phenotypic variability in future foundresses of Polistes Nimpha (Christ) (Hymenoptera, Vespidae, Polistinae) and infestation of their larvae by the mite Sphexicozela Connivens Mahunka (Acari, Astigmata, Winterschmidtiidae)". Entomological Review. 91 (6): 685–691. doi:10.1134/S0013873811060017. S2CID 1999452.
- ^ "Hornets are worthy of protection!".
- "The Wasp-Killing Myth in Germany". 25 August 2018.
- "Nein – Wer eine einzige Wespe tötet muss nicht 50.000 Euro ($51,942.25 USD) Bußgeld zahlen". 24 July 2018.
- Rortais, A; Villemant, CL; Gargominy, OL; Rome, QU; Haxaire, JE; Papachristoforou, AL; Arnold, GÉ. (2010). "A new enemy of honeybees in Europe: The Asian hornet Vespa velutina. Atlas of Biodiversity Risks–from Europe to globe, from stories to maps". Sofia & Moscow: Pensoft.
- Johnson, D.; Townsend, L. (May 2016). "European Hornet in Kentucky". University of Kentucky. ENTFACT-600.
- Okutucu, S.; Şabanov, C.; Abdulhayoğlu, E.; Aksu, N. M.; Erbil, B.; Aytemir, K.; Özkutlu, H. (2011). "A rare cause of atrial fibrillation: a European hornet sting". Anatolian Journal of Cardiology. 11 (6): 559–560. doi:10.5152/akd.2011.144. PMID 21827996.
External links
- European hornet protection site
- European Hornet - Penn State Entomology Department Fact Sheet
- "The European hornets and oak sap"