Vespula germanica

European wasp
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Hymenoptera
Suborder:	Apocrita
Family:	Vespidae
Genus:	Vespula
Species:	V. germanica
Binomial name
Vespula germanica (Fabricius, 1793)

Vespula germanica (European wasp, German wasp, or German yellowjacket) is a species of wasp found in much of the Northern Hemisphere, native to Europe, Northern Africa, and temperate Asia. It has been introduced and is well-established in many other places, including North America, South America (Argentina and Chile), Australia, and New Zealand. German wasps are part of the family Vespidae and are sometimes mistakenly referred to as paper wasps because they build grey paper nests, although strictly speaking, paper wasps are part of the subfamily Polistinae. In North America, they are also known as yellowjackets.

Taxonomy and phylogeny

Vespula germanica belongs to the genus Vespula, which includes various species of social wasps that are found throughout the Northern Hemisphere. In North America, these wasps are most commonly known as yellowjackets, but this name also applies to species within the sister genus Dolichovespula. Members of the Vespula genus are often confused with other genera, especially the paper wasp Polistes dominula. Colonies of V. germanica share many characteristics with those of V. vulgaris and V. pensylvanica, so they are very often studied together.^[1]

Description and identification

V. germanica

The German wasp is about 13 mm (0.5 in) long, has a mass of 74.1 ± 9.6 mg,^[2] and has typical wasp colours of black and yellow. It is very similar to the common wasp (V. vulgaris), but unlike the common wasp, has three tiny black dots on the clypeus. German wasps also have black dots on their abdomens, while the common wasp's analogous markings are fused with the black rings above them, forming a different pattern. However, the identification of this species might be difficult because this black mark on its clypeus can sometimes appear broken, once again making it look extremely similar to V. germanica.

The three dots on the German wasp's face distinguish it from the common wasp.

Distribution and habitat

V. germanica originated in Europe, Asia, and North Africa and can now be found in sections of almost every continent. It has established populations in North America, South America, South Africa, New Zealand, and Australia. German yellowjackets are known to be especially successful and destructive invaders of new territories. These wasps are polyphagous predators which feed on native arthropods, and because they are able to outcompete many other animals for food, they have caused considerable harm to the indigenous wildlife of areas which they have invaded. V. germanica disperses at a rate of no more than 1000 m per year, so their rapid dispersal is most likely being aided by accidental human transport of hibernating queens. This normally leads to a bottleneck effect during the establishment of new populations, which explains the significant genetic differentiation often found among geographically distant populations.^[3]

German wasp rasping wood for building its nest

The nest is made from chewed plant fibres, mixed with saliva. The majority of nests are found in the soil below ground. A significant portion of nests are found in artificial structures such as attics, and a small portion are found above ground.^[4]

Colony cycle

A single queen initiates a nest in the spring by constructing an embryonic nest, which contains a series of hexagonal cells. These cells are used to house one wasp through the immature stages of life: egg, larval instars, and pupae. The colony grows rapidly during the summer with a huge increase in worker numbers and nest size. By the end of summer, the rate of growth slows considerably and more males are produced than workers, and the focus is shifted from building small cells to building cells that are 30-40% larger. These new cells house the new queens and males. In the fall, the new queens begin hibernation, while the old queens die off and the colony collapses. In some cases, the nests may survive through the winter and reach the next season. If this occurs, the nest will become polygynous and reach a much larger size than in the previous year.^[4]

V. germanica nests are strongly affected by climate. Average colonies in Australia contain over 9500 small workers and 3600 large queen and male cells. By the end of May, colonies in Australia have 15,000 wasps emerging from small cells and 2500 wasps emerging from the large cells. This is up to 80% more than colonies observed in England where climate conditions are much less favorable. Additionally, a significant number of nests in Australia are able to survive the winter, in comparison to England where none of the nests survive. This demonstrates that V. germanica nests can reach a substantial size in tolerable climate conditions.^[4]

Behavior

Worker specialization

High variation in the sequence and diversity of tasks performed byV. germanica workers suggests limited temporal polyethism exists within the species. Temporal polyethism is a mechanism in which workers specialize on tasks in a sequential order throughout their lives, instead of performing numerous tasks concurrently. The typical order of tasks in V. germanica is nest work, pulp foraging, carbohydrate foraging and protein foraging. Only 61% of the wasps observed perform more than two of these tasks during their lifetimes. Additionally, about 40% of the wasps specialize on only one task per day, but these individuals varied with respect to age and sequence of task performance. The schedule of temporal polyethism found in V. germanica is highly accelerated when compared to other Hymenoptera, which is likely related to worker lifespan. Vespula workers may not live long enough to benefit from a strict system of worker specialization in which workers repeatedly perform one task.^[5]

Mating behavior

Vespula germanica queens are typically polyandrous. The queens mate with a moderate number of males, usually between one and seven, with no optimal number of mates. In addition, mating events are independent of each other, which discredits the hypothesis that queens terminate mating behavior after mating with a male that possesses an optimal set of attributes.^[6]

V. germanica workers are unable to mate, so are incapable of producing diploid offspring. In some nests, however, the workers produce male offspring. Worker reproduction has been documented in other Vespula wasp colonies, but usually only after the death of the queen. However, workers of V. germanica have been found reproducing in the presence of a queen.^[6]

Foraging behavior

Vespula germanica are known to be opportunistic predators and scavengers. They are extremely efficient at both hunting for small, live food sources as well as collecting from large stationary sources.^[7] It is believed that the flexibility in V. germanica’s foraging behavior is a key factor in their ability to rapidly colonize new areas in a variety of ecological environments. As scavengers, V. germanica are forced to make numerous trips between the location of the food source and the nest, where the larvae are kept and fed.^[8] This calls for V. germanica foragers to be very adept at relocating previously explored food sources.

Cognitive plasticity

The ability to relocate previously discovered food sources is only one example of the diverse cognitive mechanisms at play within V. germanica’s behaviors. V. germanica foragers continue to visit a feeder after food removal, but are able to rapidly remove associations that no longer provide a reward. The amount of time that the foragers continue to search these sites depends on the number of times they had visited it in the past. The choices made by V. germanica take into account both current and past experiences. These abilities imply that these wasps have important learning and memory capacities which enable them to memorize various resource characteristics, including the route to resources, as well as specific spatial location with respect to local landmarks. In addition, V. germanica wasps have been shown to have sensorimotor learning capacities which allow them to associate visual stimuli with certain motor responses.^[8] This reported high cognitive plasticity enables V. germanica wasps to inhabit a variety of regions. Its flexibility in nesting and diet habits in conjunction with its foraging capabilities may help explain the success with which V. germanica has invaded so many different ecological areas.^[7]

Kin selection

Genetic relatedness within colonies

V. germanica queens are typically polyandrous.^[9] Because the queen mates with multiple males, the workers are more closely related to the queen’s sons than to sons of other workers.^[6] These asymmetries in relatedness are believed to be a factor leading to worker policing within colonies.^[9]

Male skew

Male reproductive skew within V. germanica indicates that males do not contribute equally to the production of offspring when compared to females. The level of skew observed was higher among males in nests with queens that mated with multiple males. Reproductive skew may be linked to patterns of sex allocation, but sex ratio data for Vespula colonies suggest that a split-sex ratio is not produced. This contradicts the theory that the reproductive skew seen in V. germanica is part of an evolutionary strategy of males due to asymmetries in relatedness.^[6]

Worker policing

Worker policing is selected for when it benefits the colony. These benefits are achieved through various measures, such as increasing colony efficiency and establishing a female-based sex allocation ratio. Worker policing is especially seen in colonies where the queens mate with multiple males. This is because, on average, the workers are more closely related to sons of the queen than to those of other workers. In the past, V. germanica was thought to be an exception to the general pattern that high mating frequency led to low levels of worker reproduction. However, recent studies have indicated that while worker reproduction is occurring at a considerable rate, worker policing keeps these worker-laid eggs from reaching adulthood. This is demonstrated by the results of a study carried out in Belgium, which showed that while an estimated 58.4% of male eggs were laid by workers, only 0.44% of the adult males were the workers’ sons. These numbers indicate the efficiency with which worker policing is carried out. Policing by aggression, which prevents workers from laying eggs in the first place, leads to increases in colony productivity, while policing by eating the eggs of workers is more likely to result in sex-ratio benefits.^[9]

Diet

V. germanica wasps have a very diverse diet. They are known to eat carrion, live arthropods, fruit,^[8] honeydew, and processed human food and garbage.^[10] They are opportunistic scavengers and hunters, and are able to obtain food from a variety of different sources.^[7] For example, many species of Vespula wasps have been observed feeding on dead honey bees (Apis mellifera) found outside the beehive entrance in the late summer.^[11] This flexibility in diet is very beneficial since V. germanica often must compete for resources with the native biota in the areas it invades.^[12]

Pest status

This species is considered a pest in most areas outside its native range, though its long residency in North America is such that it is not treated with any level of urgency there, in contrast to areas such as South America, where the introduction is more recent, and the impacts far more dramatic, prompting a greater degree of concern over control measures.^[13]

Workers at the nest entrance

Along with the closely related common wasp and two species of Polistes, the German wasp is considered to be a pest in New Zealand. It was probably introduced in the late 19th century, but did not appear in large numbers until around 1940.^[14] Wasp numbers reach their greatest densities in beech forest of the South Island, due to the abundance of honeydew produced by the beech scale insect in this type of forest. It has a serious effect on the forest ecology, since less honeydew remains available for the native birds. German wasps were, however, quickly succeeded in much of the South Island and its beech forests by the introduction of the common wasp in the 1970s.^[15][16]

Adult foraging on flowers

References

External links

• Differences between Yellowjackets and Hornets NOTE: This link is broken.
• Photographs of Yellowjackets (and other insects) in flight
• Yellowjackets - Center for Invasive Species Research