Parachartergus colobopterus | |
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Species: | Parachartergus colobopterus |
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Parachartergus colobopterus (Licht., 1796) | |
Parachartergus colobopterus is an epiponine social wasp belonging to the subfamily Polistinae. This species can be found through Central and South America and is unique because its colonies contain multiple queens. However, relatedness among nest mates remains relatively high as a result of cyclical oligogyny, which is a system where the number of queens varies over time. [1] Because workers and queens do not demonstrate any significant morphological differentiation, individuals of this species are totipotent, capable of differentiation into any caste, because caste is not genetically determined. Relatedness and conflict therefore play a major role in determining the dominance hierarchy and behavioral patterns of this wasp species, especially the behavior of worker policing. [2] Another unique characteristic of this wasp is that it is generally a docile species: when the nest is continuously provoked, the colony members will leave the nest instead of mounting an attack. [3]
P. colobopterus is an epiponine wasp – one of several tribes of Polistinae within the family Vespidae. There are 23 other genera and 200 species of epiponine wasps. [4] This species was classified by Martin Lichtenstein in 1796. [5]
Like other Neotropical wasps, workers are not morphologically different from queens. This means that workers and queens cannot be identified visually, but can be identified by the presence of mature eggs in their ovaries, as seen after dissection. Males are similar in appearance to females, but can be identified by their testes. [1]
Colonies of P. colobopterus can be found on tree trunks and large branches. These nests grow vertically, so that the youngest combs are at the bottom of the nest. [1] The envelope of the nest is not attached to the combs; instead, it is attached directly to the substrate. [2]
P. colobopterus is a Neotropical wasp, with a range stretching through Central and South America. This wasp build nests on tree trunks or branches that are in or near forested areas. [4]
When a colony relocates to a new nesting site, the workers and queens will follow a chemical trail laid by scouts that leads them as they swarm to the new site. Once the colony has arrived at the new site, workers will begin to construct the nest and the queen will begin to lay eggs, which are cared for by the workers. Colonies may be initiated at any time of the year, and may persist for several years or only for several generations of broods. Each colony has multiple queens. When there are only one or two queens left, young females initiate dominance displays in order to determine which females will become the colony's new queens. [1] This multiple queen nesting structure is also seen in another South American wasp, Polybia emaciata .
P. colobopterus, like other wasps, is a social insect. This wasp demonstrates social behaviors such as division of labor, alarm pheromones, trial pheromones, partitioning of tasks, and extended persistence of colonies. [2] The individual wasps that make up a colony are divided into reproductive castes, where queens are able to mate and freely reproduce while workers care for young. Unlike other wasp species, P. colobopterus has multiple queens. It can be difficult to explain the occurrence of sociality in populations where individuals are not necessarily directly related, such as this wasp. However, relatedness remains high enough that sociality is a beneficial strategy. [1] P. colobopterus demonstrates that organisms do not necessarily need to have extreme differentiation between castes in order to maintain a successful, highly social system.
P. colobopterus colonies have multiple queens and demonstrate a unique dominance hierarchy. New queens are produced periodically as part of a strategy called cyclical oligogyny. [1] The patterns of conflict and relatedness in this wasp can call be traced back to this system of cyclical oligogyny. Queen number varies, fluctuating from high to low over the course of a colony's lifecycle. [2] When queens age and the number of queens has been reduced to only one or two wasps, new queens will be produced by the colony. This increases the relatedness of the colony as a whole by increasing the relatedness between the queens. [6] In some cases, too many females will attempt to be new queens and workers will then suppress some new queens. [1] Since there are no morphological differences between queens and workers, any workers may vie with each other for the position of queen. [2]
Five types of interactions between P. colobopterus wasps have been identified. Wasps may antennate, or touch each other with their antennae. They also demonstrate feeding behavior, where a wasp bearing food will offer the food to another wasp. Wasps may use their mandibles to request a feeding by biting the mandibles of another wasp; additionally, they may use their mandibles to groom another wasp by carefully moving them the mandibles over their nest mate’s body. Finally, wasps may use their mandibles to attack each other by biting another wasp’s body. Together, these interactions form the basis of the behavior of P. colobopterus. Research has demonstrated that queens rarely engage in interactions with workers except to request a feeding, and do not appear to regulate worker behavior. [7]
Like all other hymenopterans, P. colobopterus is haplodiploid. This means that unfertilized eggs will produce males, while fertilized eggs will produce females. Workers are therefore able to produce their own male offspring even though they have not been inseminated. Since each worker would be more closely related to her son than the male offspring of any other workers, it may be expected that the production of males would be a major source of conflict. [6] However, workers are more closely related to the queens than to each other, meaning that they are therefore more closely related to the male offspring of the queens than the male offspring of other workers. The best strategy for workers is to suppress the reproduction of other workers and allow the queens to produce males.
In order to maximize their individual fitness, workers will try to reduce the reproduction of other wasps. This is known as worker policing. [6] As previously mentioned, workers will attempt to suppress other workers. Worker policing also plays a large role in modulating the number of queens per colony. As cyclical oligogyny progresses, new queens will attempt to assert their dominance after the death of the old queens. Any worker could be morphologically able to function as a new queen, but the number of new queens is restricted by workers as they aggressively interact with potential new queens. [2] This suppresses the development of the female’s ovaries, preventing her from becoming queen. [8]
In many other wasp species, an individual's caste is determined morphologically; that is, workers are differentiated from queens and are physically unable to carry out the role of a queen. However, such morphological differences do not exist in P. colobopterus, and workers are therefore not physically restricted from reproducing. [1] The consequence is that adult females compete with each other for queenhood. According to kin selection theory, since workers are able to reproduce, it would be advantageous to them to reproduce instead of rearing the offspring of the queens. However, it also benefits each worker to limit the reproduction of other workers. Therefore, if there are multiple active queens in a colony, workers will collectively suppress females. [2]
Data by Strassman et al. indicates that reproduction in P. colobopterus reaches a maximum during the rainy season. At these times, there is a high number of new queens and a trend of colony foundation. Colonies have been known to persist for an extended period of time – up to 4.5 years. Mortality has not been shown to be associated with stage in the queen cycle, nor with season. The overall mortality rate of P. colobopterus is not consistent with other social insects, since it has a decreased mortality rate early in the history of a colony; this is likely due to the fact that it is founded by a swarm, not just an individual queen. Since the success of the new colony is not completely dependent on the survival of the queen, this increases the colony’s initial chance of survival. [4]
Nests represent a highly concentrated supply of resources, which makes them desirable to predators. P. colobopterus wasps have a nonaggressive response when their nest is disturbed. After the initial provocation, wasps leave the nest and begin vigorous displays on its outer envelope. If the provocation continues, females will abandon the nest entirely, leaving the offspring behind. This unique behavior is particularly interesting because the females possess stings that could be used in colony defense. However, field observations indicate females will only sting in self-defense if directly attacked. There have been several theories proposed for the defense response of P. colobopterus, but it seems most likely that this response is due to the lack of vertebrate predation. Furthermore, the nests of these wasps are camouflaged by plant matter, which may further reduce the likelihood of vertebrate predation and negate the need for aggressive defense. [3]
P. colobopterus exhibits a novel defensive behavior against vertebrates. When a predator comes near, instead of flying off and stinging the predator, the workers of this species bend their gaster around and actually spray venom at the predator, something that is likely to be effective particularly when it hits the eyes of the predator. [9]
Polistes annularis is a species of paper wasp found throughout the eastern half of the United States. This species of red paper wasp is known for its large size and its red-and-black coloration and is variably referred to as a ringed paper wasp or jack Spaniard wasp. It builds its nest under overhangs near bodies of water that minimize the amount of sunlight penetration. It clusters its nests together in large aggregations, and consumes nectar and other insects. Its principal predator is the ant, although birds are also known to prey on it. Unlike other wasps, P. annularis is relatively robust in winter conditions, and has also been observed to store honey in advance of hibernation. This species has also been used as a model species to demonstrate the ability to use microsatellite markers in maternity assignment of social insects.
Polistes exclamans, the Guinea paper wasp, is a social wasp and is part of the family Vespidae of the order Hymenoptera. It is found throughout the United States, Mexico, the Bahamas, Jamaica and parts of Canada. Due to solitary nest founding by queens, P. exclamans has extended its range in the past few decades and now covers the eastern half of the United States, as well as part of the north. This expansion is typically attributed to changing global climate and temperatures. P. exclamans has three specific castes, including males, workers, and queens, but the dominance hierarchy is further distinguished by age. The older the wasp is, the higher it is in ranking within the colony. In most P. exclamans nests, there is one queen who lays all the eggs in the colony. The physiological similarities between the worker and queen castes have led to experiments attempting to distinguish the characteristics of these two castes and how they are determined, though males have easily identifiable physiological characteristics. Since P. exclamans live in relatively small, open combed nests, they are often subject to predators and parasites, such as Chalcoela iphitalis, Elasmus polistis, and birds. P. exclamans have defense and recognition strategies that help protect against these predators and parasites.
The Central American paper wasp is a nocturnal eusocial wasp. It is famous for its swarm based emigration behavior, and is native to the lowlands of Central and northern South America. This species has developed special night vision adaptations to facilitate their night-time swarming and foraging behavior and has important medicinal properties for the Pankararú people of Brazil.
Apoica flavissima is a paper wasp found primarily in South America. The species is distinguishable by its light coloring, unique single comb nests, and nocturnal nature. A notable feature of this species is the size dimorphism between queens and workers. Unlike most Vespidae wasps, Apocia flavissima queens are smaller than their worker counterparts which results in unique intraspecies relationships.
Polybia occidentalis, commonly known as camoati, is a swarm-founding advanced eusocial wasp. Swarm-founding means that a swarm of these wasps find a nesting site and build the nest together. This species can be found in Central and South America. P. occidentalis preys on nectar, insects, and carbohydrate sources, while birds and ants prey on and parasitize them. P. occidentalis workers bite each other to communicate the time to start working.
Polistes bellicosus is a social paper wasp from the order Hymenoptera typically found within Texas, namely the Houston area. Like other paper wasps, Polistes bellicosus build nests by manipulating exposed fibers into paper to create cells. P. bellicosus often rebuild their nests at least once per colony season due to predation.
Synoeca cyanea, commonly known as the marimbondo-tatu in Brazil, is a swarm-founding eusocial wasp. Native to Brazil and Argentina, S. cyanea is one of the largest and most aggressive species of social wasps and is feared in many rural areas. It begins its colony cycle in the early spring and continues until nest abandonment. Throughout its life, S. cyanea forage sugary substances and animal carcasses for food and wood pulp for its nest. S. cyanea is also known for its strong venom, which is enough to cause haemolytic activity.
Angiopolybia pallens is a species of social wasp predominantly found in South America. The wasp is generally seen in Brazilian rainforests. This species was discovered by Lepeletier in 1836. It typically feeds on nectar and carrion. In fact much of its feeding behavior and impact on humans is centered on feeding on animal carcasses. The wasp species displays a caste differentiation that can be seen by difference in ovarian development. Additionally they have a unique colony establishment procedure. It begins with a few individuals from the nest leaving to find a good site and then the rest of the colony follows using specific communication signals that are further discussed in this article.
Agelaia pallipes is a species of social paper wasp found from Costa Rica to Brazil, Argentina and Paraguay. A. pallipes is ground-nesting and is one of the most aggressive wasps in South America. This species is a predator of other insects, including flies, moths, and ground crickets, as well as baby birds.
Protonectarina sylveirae, commonly referred to as the Brazilian wasp, is a neotropical swarm-founding wasp species that ranges widely across South America. This species relies heavily on the consumption of animal protein rather than nectar. P. sylveirae preys heavily on agricultural pests to coffee crops, keeping pest populations low.
Leipomeles dorsata is a neotropical paper wasp that is found across Central America and northern South America. It is a eusocial wasp with little differentiation between reproducing and non-reproducing females. In fact, workers can become temporary reproductives if the main reproductives are killed, allowing reproduction to continue until the main reproductive population recovers. The colony cycles through different ratios of main reproductive females and subordinate reproductive females, starting with few or no primary reproducing females, and increasing until there are only main reproductives.
The African swarm-founding wasp, Polybioides tabidus, is a social paper wasp from the order Hymenoptera that is typically found in Central Africa. This wasp is unique in that it exhibits cyclical oligogyny, meaning queen number varies with colony cycle. After several generations of production of workers and future queens, a subset of many workers and queens leave the original colony to begin a new one. The new colony does not produce new queens until current queens from the old colony have died. P. tabidus has been observed to display both predator and scavenger behavior, depending on the food sources available.
Protopolybia exigua is a species of vespid wasp found in South America and Southern Brazil. These neotropical wasps, of the tribe Epiponini, form large colonies with multiple queens per colony. P. exigua are small wasps that find nourishment from nectar and prey on arthropods. Their nests are disc-shaped and hang from the undersides of leaves and tree branches. This particular species of wasp can be hard to study because they frequently abandon their nests. P. exigua continuously seek refuge from phorid fly attacks and thus often flee infested nests to build new ones. The wasps' most common predators are ants and the parasitoid phorid flies from the Phoridae family.
Polybia sericea is a social, tropical wasp of the family Vespidae that can be found in South America. It founds its colonies by swarming migrations, and feeds on nectar and arthropods.
Polybia emaciata is a Neotropical swarm founding wasp that is mainly found in South America. This eusocial species has a unique colony structure in which multiple queens are present. Workers and queens are not morphologically distinct, but the high amount of relatedness maintained in the colony ensures that workers police each other. Polybia emaciata relies on a diet of liquid nectars which is supplemented with prey insects, particularly flies. It is well known for being one of only three wasp species in the genus Polybia that uses mud as the primary material for building its nest. Polybia emaciata is one of the least aggressive wasp species in the genus Polybia, and is known for its distinctive defensive behavior.
Synoeca surinama is a Neotropical swarm-founding wasp of the tribe Epiponini. It is known for its metallic blue and black appearance and painful sting. S. surinama builds nests on tree trunks and can be found in tropical climates of South America. When preparing to swarm, there are a number of pre-swarming behaviors that members of S. surinama colonies partake in, such as buzzing runs and occasional brood cannibalism. In S. surinama, social environmental conditions determine the caste ranks of individuals in the developing brood. Unlike less primitive Hymenoptera species, S. surinama display little morphological variation between egg laying queens and workers. S. surinama wasps visit flowering plants and are considered pollinators. When these wasps sting, the stinger is left in the victim and the wasp ultimately dies.
Parachartergus fraternus is a neotropical, swarm founding, polistine wasp species that is distributed throughout Central and South America. They live in nests in second growth tropical dry forests, near pasture fields, roadside areas, and urban areas as well. These wasps eat insects, such as caterpillars of Lepidoptera. They also drink nectar, honeydew, and water. The workers capture their prey during foraging. They also use venom to paralyze their prey in order to consume it later. P. fraternus wasps are not very aggressive and they do not attack when the nest is approached.
Synoeca septentrionalis is one of five species of wasps in the genus Synoeca. It is a swarm-founding wasp that is also eusocial, exhibiting complicated nest structure and defense mechanisms and a colony cycle including a pre-emergence phase and a post-emergence phase. It is typically found in areas from Central to South America. This wasp is one of the larger species of paper wasps and exhibits multiple morphological adaptations as a result of this. Synoeca septentrionalis is known for possessing a very painful sting.
Agelaia multipicta is a swarm-founding, highly eusocial wasp that lives in Mexico, Argentina, Trinidad and southern Brazil. It nests in natural cavities such as hollow trees and aggressively defends the nest from ants, who are brood predators. The workers and queens are morphologically distinguished by ovarian development as well as external features such as a larger petiole and gaster in the queen. Like other carrion-eating (necrophagous) wasp species, A. multipicta plays a scavenging role in the ecosystem. Agelaia multipicta was described by the Irish entomologist Alexander Henry Haliday in 1836.
Metapolybia cingulata is a species of social paper wasp known for having queens who exhibit usurpation behaviors and also for having flexible behavior groups. It is difficult to tell what a specific wasp's role is just upon observation. Therefore, dissections are usually performed in order to determine the sex, and thus the wasp's role in the colony. The regulating behavior, or nest building, most closely resembles that of primitive Polistes models where a queen solicits liquid from a fellow worker and then takes pulp to initiate building a cell without leaving the nest. In nest building, the amount of water present in the nest as well as the activity of the builders of the nests determines the activity of the colony.