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Polistes dominula

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Polistes dominula

European Paper Wasp
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Vespidae
Genus: Polistes
Species: P. dominula
Binomial name
Polistes dominula
(Christ, 1791)
Synonyms

Polistes gallicus auct.

Polistes dominulus

The European paper wasp[1] (Polistes dominula often misspelled as dominulus) is one of the more common and well-known species of social wasps in Europe. The dominant females (queens) are the principal egg layers, while the subordinate female (auxiliaries) primarily forage and do not lay eggs. This hierarchy is not permanent; when the queen is removed from the nest, the second-most dominant female assumes the role of the queen. Dominance in females is indicated in the coloration of the clypeus (face), whereas dominance in males is shown in the coloration of the abdomen. Polistes dominula wasps have a lek-based mating system. Unlike most social insects, 35% of Polistes dominula wasps in a colony are unrelated. It is considered an invasive species in Canada and the United States.

Nomenclature

Polistes dominula is frequently referred to as Polistes gallicus, which is a separate species. It was originally described in 1791 by Johann Ludwig Christ as Vespa dominula. The specific epithet dominula, is a noun meaning "little mistress",[2] and following the International Code of Zoological Nomenclature, species epithets which are nouns do not change when a species is placed in a different genus. Authors unaware that dominula was a noun have misspelled the species name as dominulus for decades. Polistes dominula is often referred to as the European Paper Wasp because of its native distribution and its nests, which are constructed from paper and saliva.[3]

Life Cycle

Nests are begun by overwintering founding queens, or foundresses, which spend about a month in the spring constructing a nest and provisioning offspring, the first of which will become daughter workers in the growing colony. One or more foundresses will begin the colonies in the spring. If there are multiple foundresses, the one who lays the most eggs will be the dominant queen. The remaining foundresses will be subordinate and do work to help the colony. [4]

Males are produced later, and when they start to appear, a few daughters may mate and leave their nest, to become foundresses the next season. The switch from production of workers to production of future foundresses (gynes) is not utterly abrupt, as has been considered the case for other species of Polistes.

The colony disperses in the late summer, with only males and future foundresses produced instead of workers, and individuals frequently cluster in groups (called a hibernaculum) to over-winter. Hibernation does not usually take place on former nest sites.


Dominance Hierarchy

Caste System

Hierarchies in social insects serve two functions:

  1. allows a single reproducing individual to emerge and
  2. enables the progressive exclusion of non-reproducing individuals from the nest space[5]

Morphologically, there are few differences between the foundress and subordinate members of the colony. However, several studies have shown that behavioral differentiation does occur,[6][7] with the role an individual female takes being determined by social interaction within the colony. Dominant females, also known as the queens, are the principal egg-layers. Queens occupy the nest, oviposition, and rarely forage. In comparison, auxiliaries, or subordinate females, primarily forage and do not lay eggs.[8] Autumnal helpers display a unique, behavioral phenotype demonstrated by only a small percentage of workers. These individuals leave their natal nest to overwinter in order to found new colonies in the spring. Compared to non-helpers, these helpers exhibit higher overall levels of activity, demonstrating “higher hourly frequencies of behavioral type”.[9] Specifically they gave “more trophallaxis, performed more attacks, and received more RDB [ritualized dominance behavior]... Non-helpers showed higher levels of trophallaxis received, and performed more RDB”.[9] Researchers have also found that the survival rate of helpers were around 14% while the survival rate of non-helpers was around 59%.[9]

These behavioral divisions are not permanent. For instance, if an alpha female is removed from a nest, then another female, usually the second-most dominant, beta female, assumes the role and behavioral profile of the removed dominant. Indeed, individuals alternate between different profiles of behavior within their own dominance rank position. Liebig et al. found that when larvae were artificially removed, the frequency of worker reproduction increased. They concluded that workers lay eggs when they perceive a decline in queen power, as demonstrated by artificially empty cells.[10]

Evidently, the interactions of females in the nest can influence which daughters become workers and which become gynes. Despite some minor physiological differences (primarily in the fat body), gyne-destined females produced late in the colony cycle can be induced to become workers if placed on nests that are at an earlier stage of colony development, and the reverse is also true. This indicates a significant degree of flexibility in the caste system of this species.

Female Dominance

Typically, the alpha female dominates all other individuals of a colony, laying the majority of eggs and partaking in differential oophagy. The alpha female devotes much of her time to social interaction, in comparison to subordinates that are much more involved in foraging and brood care.[11] Tibbetts and Dale demonstrated that the clypeus, the yellow region above the mouth, exhibits “remarkable variation across individuals in the number, size, and shape of black spots” and this variation correlates to dominance; more dominant individuals have more black spots.[12] Zanette et al. found that the arrival at the nest correlates with the dominance hierarchy. Therefore, individuals that join the nest later are seldom dominant.[13]

Polistes dominula show distinct behavioral differences in response to face marking. Tibbetts and Lindsay used paint to alter the number of facial spots on two wasps of the same size after killing them. They put these dead wasps as guards in front of food sources and introduced a third wasp to see where this wasp would go. The third wasps chose the food source that was guarded by the wasp with less spots 39 out of the 48 times (Davies et al.). Therefore, the dominance of wasps of the same size is predicted by facial coloring, with more dominant individuals having more spots.[14] Tibbetts and Dale experimented to determine whether social costs maintain the honesty of facial signals. They altered the facial coloring of wasps and then put them together to battle. The winner is clearly identified by mounting the loser while the loser lowers its antennae. While the manipulation of coloring did not affect who won the battle for dominance, it did significantly affect the behavior after the battle. Losers who were painted with more spots experienced six times more aggression than controls who had not been painted. Tibbetts and Dale concluded that the honesty of facial coloring is explained by social costs that are imposed when wasps do not signal honestly.[15]

Male Dominance

Male abdominal spots correlate with dominance. Smaller, elliptically shaped spots reflect a more dominant male who is preferred by females and wins competitions with other males. This is in contrast to males with larger, irregularly shaped abdominal spots who are generally subordinate and less sexual successful. [16]

Abdominal Behavior

Abdomen rubbing occurs during the egg stage on multiple-female colonies more than on solitary colonies. Alpha females perform abdomen stroking more than subordinate females. After an alpha female is removed, subordinates increase their abdomen stroking rate. Dani et al. suggest that the substance released by abdomen stroking has two potential functions: (1) to repress future ovarian development in the brood or (2) to inform the brood of which adult female is the dominant.[17]

Relatedness

Polistes dominula is a social insect which lives in colonies. In most social insects, colonies are composed of related individuals. It is usually assumed that social insects help close relatives. However, in Polistes dominula 35% of the nest mates are unrelated.[4][12] In many other species, unrelated individuals only help the queen when no other options are present. In this case, subordinates work for unrelated queens even when other options may be present. No other social insect submits to unrelated queens in this way. This seemingly unfavorable behavior parallels some vertebrate systems. It is thought that this unrelated assistance is evidence of altruism in Polistes dominula. [4]

It was found that the majority of nests had one or more females that were unrelated, especially in the winter before nests are formed and workers born. The nests tend to form from foundresses of different nests from previous years. The foundresses are also found over-wintering with other wasp species, showing why unrelated wasps are found during winter. However, after winter when the nests are starting to be formed, there is an increase in relatedness in the nests, which could result from foundresses searching for more related sisters, instead of unrelated wasps. After winter, as wasps leave their winter areas and return to their nests, there is an increase in relatedness in the early nest phase. In later stages of the nest, there are more unrelated wasps, which could be because new wasps join established nests. [18]

Still, Polistes dominula females are able to distinguish between nestmates and non-nestmates. Triads of wasps show more discrimination and aggression toward non-nestmates than dyads of wasps. In triads, there is increased aggression because a defense of the nest can be shared with other nestmates, but for an individual the cost of the aggression is greater than the benefit of the defense. [19]

Behavior

Foundress Behavior

Pratte observed that in the first 12 days of the nesting period, 75% of foundresses left their original nest, traveling to around three nests before settling permanently. Pratte suggested that foundresses visit multiple nests before choosing to determine the nest with the highest reproductive payoff.[8] Foundresses choose nest sites by weighing the benefit of an expanded colony with the cost of predation risk.[20] Zanette et al. demonstrated that multiple-foundress nests have a higher chance of survival compared to single-foundress nests, and in general, foundresses found nests with those they hibernate with in the same aggregation.[13]

Comb Arrangement

The nest, consisting of a single comb, is the heart of the colony, “where food is stored and the immature brood reared, as well as the central spatial reference where the majority of individuals spend most of their time”.[21] Baracchi et al. found that Polistes dominula do not occupy the comb in random distribution. Each wasp spends the majority of its time on the comb in a relatively small area, approximately 12% of the comb. This small use of space is the norm regardless of the number of wasps on the nest. However, this area could occasionally cover up to 50% of the comb. Dominant females occupy a smaller area than do subordinate foundresses and workers. Superimposition rates are low, which suggests that wasps limit each other’s spaces. Workers prefer to overlap areas with other workers, while foundresses prefer to overlap areas with other foundresses. Around 70% of workers are active and occupy a small area of the comb, while the rest do not have particular fidelity areas and spend most of their time away from the nest or remaining motionless behind the nest. Alpha females are affected by cell content, resting more frequently on capped cells and avoiding empty ones.[21]

Cooperation

Cooperation provides survival benefits; “Multiple-foundress colonies are more likely to survive to produce offspring than are single-foundress colonies, and individual foundresses in multiple-foundress groups are less likely to disappear before worker emergence than foundresses nesting alone. Therefore, association provides substantial productivity and survival benefits for cooperating foundresses”.[22] Tibbetts and Reeve suggest that cooperation provides survival benefits only if individual foundresses on a multi-foundress nest have a greater chance of survival than individuals who found nests alone or the foundress’ contribution to the nest is preserved even after she leaves.[22]

Queen Loss

Strassmann et al. suggest that queen loss is a crisis for a colony of social insects. This situation may cause “increased aggression and work inefficiencies, and the new queen, if the colony can produce one, could be slow to develop mature eggs”.[23] Strassmann et al. observed, “replacement queens did not mate in the first 12 days after queen removal and few had mature eggs in their ovaries, though after a month most had mated and had developed ovaries".[23] Nest growth decreases with colonies that lose their queen, which can be expected if increased dominance behavior interfered with other essential behaviors or if the new queen is not competent at egg laying. Strasmmann et al. found that among queenless colonies of P. dominula, individuals demonstrated an increased level of chewing and climbing, but not of lunging and biting. These are all dominance behaviors. There is no difference of foraging behaviors between colonies with and without queens. Strassmann et al. concluded, “the high cost of replacing the queen may indicate that workers are kept reproductively suppressed enough not to be a threat to the existing queen".[23]

Nest Adoption

Adoption is a result of three situations: when (1) queens lose their nests and ‘make the best of a bad situation,’ (2) workers leave multiple-foundress nests, and (3) subordinates employ a ‘sit-and-wait’ strategy, waiting for nests to be orphaned. Nests are orphaned when the adult wasps die while taking care of their nest, leaving an immature brood. Orphaned nests allow a new wasp to gain status as queen without a fight.[24] Starks observed that females who adopt nests are less cooperative and expend less energy than those who found nests. “Spring foundresses sometimes initiate colonies alone, form associations with other females, usurp established conspecific colonies, or even adopt abandoned nests”.[25] Females would adopt an abandoned nest if they lost their nest due to predation or other damage or if they waited to adopt an orphaned nest rather than found their own. Starks found that females engaging in the sit-and-wait reproductive strategy adopted the most mature nest rather than the nest with an increased probability of containing kin (from the same population). Stark concludes that females prioritize the quality of the nest over rescuing possible kin from another abandoned nest. Females demonstrate a preference for mature nests and nests with a large proportion of fourth and fifth instar larvae. When a female adopts an orphaned nest, she will destroy the existing eggs and instar larvae, but allow older larvae and pupae to complete development. Stark concludes that adopting a nest maximizes the potential of “possessing a mature pre-worker phase colony without expanding excessive energy during the nest founding period of cooperating in the construction of the nest”.[25]

Of note, once a nest is established it may be used by multiple generations and in multiple seasons. Nests aboandonded during the hibernation cycle are almost always re-populated the following spring. With each succession, the nest continues to grow in size. A few have been recorded at 8 inches in diameter. It has also been noted that multiple nests may be located in the same area within inches of each other.[26]

Mating

Polistes dominula wasps have a lek-based mating system. Males compete intensely for dominant positions on the lek, while females are scrupulous when selecting their mate. Males form aggregations on the uppermost portions of structures such as fences, walls, roof peaks, and trees. Males often fight with other males in mid-air or on the structure. Males who lose will fly away from the lek. Females fly through leks or perch near lekking areas to observe males before making choices on mates. Females use the highly conspicuous abdominal spots on males, which are highly variable in size and shape, to aid in mate choice. Males with smaller, more elliptically shaped spots are more dominant over other males and preferred by females compared to males who have larger, more irregularly shaped spots.[27]

Social insect males are often seen as mating machines, with an undiscriminating eagerness to mate. However, males encounter costs of unsuccessful mating in terms of energy investment. Therefore, Polistes dominula males are able to recognize female castes and preferentially choose reproductive females to workers, regardless of health or age. Males are able to differentiate castes through perceiving differences in chemical signals and physiological status. While males are able to discriminate between castes, they are not capable of discriminating between health as males showed a strong preference for gynes, both healthy and parasite-castrated, compared to workers. This is because males distinguish females by CHC profiles and CHC profiles of healthy and parasitized gynes are very similar. Therefore, males are not able to evaluate the true reproductive potential of the females they encounter.[28]

Cuticular Hydrocarbon Profile

Insect body surface is coated with cuticular hydrocarbons (CHC) for waterproofing. These chemicals also contribute to recognition among individuals, kin, and nest mates (Inoue-Murayama et al.). An analysis of the differences in CHC profiles between dominant and subordinate females found that while differences are not clear at the early stages of nest founding, these differences become prominent upon worker emergence.[29] The CHC profiles of the dominant female exhibit a greater proportion of “distinctive unsaturated alkenes of longer chain length compared to those of subordinates and subsequent worker offspring” (88).[30] When the queen is removed, the replacement queen’s CHC profile becomes similar to that of the original queen (Inoue-Murayama et al.). Dapporto et al. analyzed whether CHC is a signal for fertility or dominance and concluded that it is a signal for dominance because “subordinate individuals with developed ovaries still exhibited profiles that differed from those of dominant individuals” (88).[30]

Parasitoidism

Polistes dominula nests, constructed from paper and saliva, and resident brood host “parasites and parasitoid, including predacious Lepidoptera larvae, Hymenoptera, Diptera, and Strepsiptera” (257).[3] Madden et al. reported the “first well documented case of pasitoidism of the invasive population...in North America” in 2010.[3]

Distribution

The native range of Polistes dominula covers much of southern Europe and North Africa, and temperate parts of Asia as far east as China.[31] It has also been introduced to Australia, New Zealand, South Africa,[32] and North and South America.

Spread to the United States

This species was introduced into the United States in 1968 in the New Jersey Pine Barrens and spread throughout most of the country during the 1980s and '90s, in some cases partially replacing native species. Another introduction was discovered in the late 1970s in Cambridge, Newton and Somerville, Massachusetts. The first wave consisted of solitary-founding but socially-nesting individuals, then a wave of social founders (several females found a new nest together). In warmer regions, there have been reports of "supercolonizers" most of whom enlarge their natal nests in successive years, rather than dispersing.

In the United States, P. dominula nests earlier in the spring, in a wider variety of nest sites, and feeds on a larger variety of insects than native species, which feed almost exclusively on caterpillars. Most entomologists consider it to be an invasive species. This wasp can be mistaken for a yellowjacket, as it is black strongly marked with yellow, in a pattern very reminiscent of a yellowjacket, and quite different from the native North American species of Polistes.

Behaviors adaptations of the Polistes dominula are causing it to move past its native Europe and invade the United States. Polistes dominula has a more general diet. Most Polistes species in the United States feed only on caterpillars but the Polistes dominula eats many different genera of insects. Polistes dominula also nest in areas with better protection, and are then able to avoid predation that has affected many other Polistes species. The invasive Polistes dominula also nest in Northern states of the U.S. which has a similar ecology and habitat to that in Europe. This allows for a faster colonization. [33]

Polistes dominula was also compared to and found to be more productive than Polistes fuscatus, which is indigenous to the United States. Polistes dominula also produce workers about a week earlier and forage earlier in the day than Polistes fuscatus. These behaviors show why they are able to spread in the United States so quickly. [34]

References

Further Reading

External Links

  • reference photographs, biological notes
  • facial markings indicating dominance
  • "The European Paper-Wasp: A New Threat to Cavity-nesting Birds Is Coming!
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