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The Norwegian wasp (''Dolichovespula norwegica'') is a species of
eusocial Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping gen ...
wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder ...
. It is common in
Scandinavia Scandinavia; Sámi languages: /. ( ) is a subregion in Northern Europe, with strong historical, cultural, and linguistic ties between its constituent peoples. In English usage, ''Scandinavia'' most commonly refers to Denmark, Norway, and S ...
and can also be found in
Scotland Scotland (, ) is a country that is part of the United Kingdom. Covering the northern third of the island of Great Britain, mainland Scotland has a border with England to the southeast and is otherwise surrounded by the Atlantic Ocean to ...
and other areas in
Britain Britain most often refers to: * The United Kingdom, a sovereign state in Europe comprising the island of Great Britain, the north-eastern part of the island of Ireland and many smaller islands * Great Britain, the largest island in the United King ...
and
Ireland Ireland ( ; ga, Éire ; Ulster-Scots: ) is an island in the North Atlantic Ocean, in north-western Europe. It is separated from Great Britain to its east by the North Channel, the Irish Sea, and St George's Channel. Ireland is the s ...
. Often known for being a tree wasp, it nests in low branches and bushes and feeds on insects. It also obtains nectar from
blueberry Blueberries are a widely distributed and widespread group of perennial flowering plants with blue or purple berries. They are classified in the section ''Cyanococcus'' within the genus '' Vaccinium''. ''Vaccinium'' also includes cranberries ...
and
snowberry ''Symphoricarpos'', commonly known as the snowberry, waxberry, or ghostberry, is a small genus of about 15 species of deciduous shrubs in the honeysuckle family, Caprifoliaceae. With the exception of the Chinese coralberry, '' S. sinensis'', whi ...
flowers.''Dolichovespula norwegica'' (Norwegian wasp). 2009. http://www.record-lrc.co.uk/c1.aspx?Mod=Article&ArticleID=Dolichoveespula_norwegica ''Dolichovespula norwegica'' (Norwegian wasp), Record 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.Bristol Wasp Control Experts .''https://pestbristol.co.uk/british-wasps/'' (Norwegian wasp),Bristol Pest Control


Taxonomy and phylogeny

''D. norwegica'' is placed in the family
Vespidae The Vespidae are a large (nearly 5000 species), diverse, cosmopolitan family of wasps, including nearly all the known eusocial wasps (such as ''Polistes fuscatus'', ''Vespa orientalis'', and ''Vespula germanica'') and many solitary wasps. Each ...
and the genus ''
Dolichovespula ''Dolichovespula'' is a small genus of social wasps distributed widely throughout the Northern Hemisphere. The yellow and black members of the genus are known by the common name yellowjackets in North America, such as '' Dolichovespula norwegica' ...
''. Based on recent studies on
mitochondria A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is used ...
l
genes In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
, ''Dolichovespula'' and ''Vespula'' are
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gr ...
, meaning they descended from common ancestors. Two species groups, ''maculata'' and ''norwegica'' form the ''Dolichovespula'' clade. While the ''maculata'' has physical attributes such as pronotal
striae Stretch marks, also known as striae () or striae distensae, are a form of scarring on the skin with an off-color hue. Over time they may diminish, but will not disappear completely. Striae are caused by tearing of the dermis during periods of r ...
, emarginated apices of the seventh metasomal
sternum The sternum or breastbone is a long flat bone located in the central part of the chest. It connects to the ribs via cartilage and forms the front of the rib cage, thus helping to protect the heart, lungs, and major blood vessels from injury. Sha ...
in males, and aedeagal medial lobes, ''D. norwegica'' females have long oculomalar space and lateroanterior clypeal angles with less prominent semicircular projections. In the past, ''D. norwegica'' and ''D. albida'', the Arctic
yellowjacket Yellowjacket or yellowjacket is the common name in North America for predatory social wasps of the genus, genera ''Vespula'' and ''Dolichovespula''. Members of these genera are known simply as "wasps" in other English-speaking countries. Most of ...
from northern North America, were considered to be the same species, but studies in 2011 of the male genitalia show that they are not
conspecific Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organis ...
. Often, male genitalia are used to characterise the
Nearctic The Nearctic realm is one of the eight biogeographic realms constituting the Earth's land surface. The Nearctic realm covers most of North America, including Greenland, Central Florida, and the highlands of Mexico. The parts of North America ...
and
Palearctic The Palearctic or Palaearctic is the largest of the eight biogeographic realms of the Earth. It stretches across all of Eurasia north of the foothills of the Himalayas, and North Africa. The realm consists of several bioregions: the Euro-Sib ...
forms of ''D. norwegica''.


Description and identification

The Norwegian wasp has eye-catching black and yellow colour patterns over its body. Starting from the face, it has a long malar space and is separated by a black bar that runs vertically. The sides of its
thorax The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the c ...
contain small black hairs that project outwards, and the rear has distinct yellow spots. One can often see red colouring on the front of the
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the to ...
. ''Dolichovespula'' is derived from the Greek word, ''dolichos'', which means long. Its genus name matches with physical characteristics.Spradbery, Philip. 1973., ''Wasps'', Sidgwick and Jackson. pp. 8, 245. 0.283.97938.0. Accessed on 25 September 2014. The typical length of an individual of this species is 11– 18 mm long. Compared to ''D. maculata'', whose adults range from 2–3 cm, queens are generally around 1.7 cm and workers are smaller, around 1.3 cm Ings, Tom
"British Social Wasps"
, ''BWARS (Bees, Wasps and Ants Recording Society)'', Retrieved on 24 September 2014. p. 3


Nests

''D. norwegica'' nests have a loose, coarse-woven texture, which suggests that the species is
terrestrial Terrestrial refers to things related to land or the planet Earth. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on or near the ground, as opposed to ...
. One such nest was found, spherical in size, about 10.5 cm long, 10.5 cm at its widest point, and 15 cm off the ground. This nest was fastened from the top of a
willow Willows, also called sallows and osiers, from the genus ''Salix'', comprise around 400 speciesMabberley, D.J. 1997. The Plant Book, Cambridge University Press #2: Cambridge. of typically deciduous trees and shrubs, found primarily on moist so ...
branch and was further stabilised by a side branch attachment. The nest consisted of 11 envelope layers with a few extra layers on the edge of the top comb. The texture of each layer was rough with some holes from weaving. Some fibres were fine and grey, while some were coarse and straw-coloured. Occasionally, black- and rust-coloured fibers are seen, as well. The most common source of nest fibres is weathered wood. On the outer layer of the nest, leaves of broadleaf plants were also observed. The observed nest had three combs and 357 cells. From close observations, the worker cells in comb one were an average of 5.17 mm, while the reproductive cells in combs two and three had an average of 6.33 mm. Nests of ''D. norwegica'' in
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
were reported to have a maximum of 1,400 cells. Overall, nests of ''D. norwegica'' wasps are commonly found on tree branches, bushes, walls of houses, and even cavities in the ground. This wasp species apparently prefers to build nests in
rural In general, a rural area or a countryside is a geographic area that is located outside towns and cities. Typical rural areas have a low population density and small settlements. Agricultural areas and areas with forestry typically are de ...
areas.


Distribution and habitat

Norwegian wasps typically construct their nests on tree branches or bushes, often selecting areas that are closed rather than exposed, such as moors. Despite its common name, ''D. norwegica'' is also found throughout England, Ireland, and commonly in Scotland. Since nests are of a coarse, woven texture with a few holes, they are detracted from thermoregulation. Furthermore, nests are also less durable, so must be built in protected areas.


Colony cycle

''D. norwegica'' queens and males leave their home colonies around late summer, when the queens are fertilised and pass the winter by finding an
overwintering Overwintering is the process by which some organisms pass through or wait out the winter season, or pass through that period of the year when "winter" conditions (cold or sub-zero temperatures, ice, snow, limited food supplies) make normal act ...
site. Before spring arrives, ''D. norwegica'' males die. Upon arrival of spring (around mid-April or early May), the queens emerge and feed, with the goal of finding a new nest location. First, the queen starts building her nest, and rears workers in the first cells (average diameter of 4.5 mm). The workers then replace the queen in the nest and assume brood-rearing activities. Later cells, where queens and males are reared, on average are 5.5 mm in diameter. By June, nests have about 50 workers, and by the end of July, about 300 workers. After this period, however, the number of workers declines rapidly. The largest number of workers in a colony ever reported was 363, while other colonies had around 150 workers. A typical colony produced 1471 wasps (measure from 14 colonies with large cells), and of that total, 43% were queens and 57% were drones. The colony cycle of ''D. norwegica'' is short; this species has a tendency to start in the spring. Their strategy is known as summer-advantage strategy, which is distinctive of the ''Dolichovespula'' genus. Such a strategy is implemented and used to take advantage of the short, yet favorable conditions of the summer. Most colonies die out after mid-August and a few last until early September. The typical number of days a colony exists is 95–115.


Behavior


Mating

''D. norwegica'', as is common among social wasps,
mating In biology, mating is the pairing of either opposite- sex or hermaphroditic organisms for the purposes of sexual reproduction. ''Fertilization'' is the fusion of two gametes. ''Copulation'' is the union of the sex organs of two sexually reprod ...
commonly occurs during the warm season. Males tend to occupy a dense area on trees and shrubs, and perform
nuptial A wedding is a ceremony where two people are united in marriage. Wedding traditions and customs vary greatly between cultures, ethnic groups, religions, countries, and social classes. Most wedding ceremonies involve an exchange of marriage vo ...
flights. They swarm in groups. ''D. norwegica'' males and a few workers have been found with on a rocky summit in Scotland in groups.


Kin selection

As in related wasps, the queen mating frequency and sperm use both influence paternity, which is important for influencing colony kin structure and reproductive tendencies of the colony. In ''D. norwegica'' colonies, male paternity is low compared to ''Vespula''. This leads to worker-queen conflict over male production. Of five species of the genus ''Dolichovespula'' studied, ''D. norwegica'' has the second-lowest value for effective paternity (1.08). Worker-worker relatedness is high among ''D. norwegica'' with a value of 0.71.


Genetic relatedness within colonies

Since the effective mating frequency of ''D. norwegica'' queens is low, worker-worker
relatedness The coefficient of relationship is a measure of the degree of consanguinity (or biological relationship) between two individuals. The term coefficient of relationship was defined by Sewall Wright in 1922, and was derived from his definition of th ...
is higher than worker–queen relatedness. Genetically speaking, this means that each worker is more related to other workers' sons than the queen's sons, creating worker-queen conflict over reproduction. In one colony of ''D. norwegica'', two matrilines were found, suggesting that the nest was once taken over by another queen. This is intriguing because it is the first case in ''Dolichovespula''.


Worker–queen conflict

Several examples of worker-queen conflict exist among members of ''D. norwegica''. Few workers were detected to have full
ovary The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. ...
activation and were producing males. Ovary activation is most likely due to the absence of the queen. Workers may kill off their queens so they can reproduce; no queens were found on collection of 12 of 14 ''D. norwegica'' nests.
Queens Queens is a borough of New York City, coextensive with Queens County, in the U.S. state of New York. Located on Long Island, it is the largest New York City borough by area. It is bordered by the borough of Brooklyn at the western tip of Long ...
are often recognizable due to their heavily worn wings.
Matricide Matricide is the act of killing one's own mother. Known or suspected matricides * Amastrine, Amastris, queen of Heraclea, was drowned by her two sons in 284 BC. * Cleopatra III of Egypt was assassinated in 101 BC by order of her son, Pto ...
is common after workers are reared and queens have laid their eggs in annual colonies with low paternity. Despite possibilities of matricide and ovary activations, workers produce very few males.
Worker policing Worker policing is a behavior seen in colonies of social hymenopterans (ants, bees, and wasps) whereby worker females eat or remove eggs that have been laid by other workers rather than those laid by a queen. Worker policing ensures that the o ...
over eggs is also seen where workers choose to keep queen-laid eggs and remove worker-laid eggs. Workers are unlikely to reproduce because it is too costly; worker reproduction can reduce colony productivity and/or obstruct reproduction of females in the colony. Furthermore, very few workers have active ovaries.


Worker policing

Three general forms of egg policing are found to conclusively contribute to reproduction ratios of ''D. norwegica''. All three forms target worker-laid eggs over queen-laid eggs. Two forms that are probably most influential of policing are worker policing and selfish policing. Despite being more related to other workers' sons, worker-laid eggs were consumed by other workers. About two-thirds of those workers were reproductive workers. Occasionally, these workers replaced the egg with one of their own (in 31% of the cases were due to self policing). Workers never removed their own eggs, and if a nonreproductive worker removed a worker-laid egg, then the queen would replace it with her own egg. Furthermore, an important part of worker policing among ''D. norwegica'' is specialization. Evidence for specialization in policing was found when observed number of policing workers was lower than the estimate true number of policing workers. An estimated average of 14 workers specialised in policing (a fourth of the total work force).


Queen policing

The last form is queen policing of eggs. About 32% of policing over worker-laid eggs was done by the queen (roughly one-third). Two cases are known where queen-laid eggs were eaten; in one, if a reproductive worker eats the egg, then it will replace it with its own egg, and if a nonreproductive worker eats the egg, then the queen always lays the replacement. These situations were less common and only occurred 8% of the time. Queen policing is popular in
colonies In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the '' metropolitan state'' ...
that are small and of low paternity. Relatively smaller colonies are better because the queen can monitor and control her workers.


Interaction with other species


Diet

''D. norwegica'' larvae commonly feed on
flies Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advanced m ...
, other
insects Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs of j ...
, and
spiders Spiders (order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species di ...
that are brought to them and chewed up by the adult wasps to a paste form. The adult individuals feed on nectar from''
Angelica sylvestris ''Angelica sylvestris'' or wild angelica is a species of flowering plant, native to Europe and central Asia. An annual or short-lived perennial growing to a maximum of , it has erect purplish stems and rounded umbels of minuscule white or pale p ...
'', ''Heracleum spondylium'', and ''Chamerion angsutifolium''. They generally prefer a sweeter taste. Larvae of ''D. norwegica'' have a single tooth, used to feed on the paste. Once it feeds, the larva expels a sweet liquid, which the adult wasp consumes. ''D. norwegica'' also enjoys flowers of umbellifers such as the wild parsnip.


Parasites

Norwegian wasps are known to be a host for ''
Dolichovespula adulterina ''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 ...
'', which is also a known social
parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson h ...
of '' D. saxonica''.Edwards R. (1980). ''Social Wasps: Their Biology and Control''. Rentokil Ltd., East Grinstead. However, ''Dlichovespula adulterina'' also uses ''D. norwegica'' as another host. One suggestion for this
inquiline In zoology, an inquiline (from Latin ''inquilinus'', "lodger" or "tenant") is an animal that lives commensally in the nest, burrow, or dwelling place of an animal of another species. For example, some organisms such as insects may live in the h ...
interaction between ''D. adulterina'' and ''D. norwegica'' is that ''D. adulterina'' has been found to cohabitate with ''D. norwegica''. The two species were found to occupy the same space in
northern Norway Northern Norway ( nb, Nord-Norge, , nn, Nord-Noreg; se, Davvi-Norga) is a geographical region of Norway, consisting of the two northernmost counties Nordland and Troms og Finnmark, in total about 35% of the Norwegian mainland. Some of the lar ...
, where no ''D. saxonica'' wasp were to be found nearby. Furthermore, in a few nests of ''D. norwegica'', ''D. norwegica'' workers lived with a queen from ''D. adulterina''. Other accounts report as many as 46 queens and 24 males of ''D. adulterina'' in a nest of ''D. norwegica''. Another case observed two unemerged males of ''D. norwegica'' and one unemerged male of ''D. adulterina'' from the nest. Also, cases have been observed of ''D. norwegica'' displaying parasitic behavior on ''D. sylvestris''. Moreover, ''D. norwegica'' shows
aggression Aggression is overt or covert, often harmful, social interaction with the intention of inflicting damage or other harm upon another individual; although it can be channeled into creative and practical outlets for some. It may occur either reacti ...
towards individuals of ''D. sylvestris''. Evidence of damaged legs and wings of ''D. sylvestris'' were found.


Human importance

''D. norwegica'' wasps previously were rarely considered to be
pests PESTS was an anonymous American activist group formed in 1986 to critique racism, tokenism, and exclusion in the art world. PESTS produced newsletters, posters, and other print material highlighting examples of discrimination in gallery represent ...
. They build their nests far from human activity, and rarely visited buildings, picnics, and fruits. However, if their nests were built on a hedge, low eaves, or close to humans, removing the nests was needed. Although they are less aggressive than other wasps in ''Vespula'', individuals of ''D. norwegica'' protect their nest with their stingers. More recently, more cases of pest problems relating to ''D. norwegica'' have been reported. Although they can be annoying in some cases, their predation activity on other arthropods makes them valuable allies of farmers. Their role in the pollination of wild plants is also of great importance for the protection of biodiversity.


References

{{Taxonbar, from=Q1410407 Vespidae Hymenoptera of Europe Insects described in 1781