Polistes carolina

One of two types of Red Paper Wasps (Polistes Carolina) is species of social wasp (subfamily Polistinae) in the family Vespidae. They are most commonly found in the eastern US from Texas through Nebraska. The wasp’s common name is due to the reddish-brown color of its head and body. Red Paper wasps are known to construct some of the largest nests of any wasp species and prefer to build their nests in protected spaces.^[4]

Taxonomy and phylogeny

The first description of Polistes carolina appears in the first volume of Carl Linnaeus' 12th edition of Systema Naturae published in 1767.^[5] In this volume he referred to the species as Vespa carolina.^[5] Ferdinand de Saussure later moved it to the genus Polistes in 1855 after Pierre Andre Latreille coined the new genus in 1802.^[6]P. carolina is within the family Vespidae, which includes nearly all of the eusocial wasps and many of the solitary wasps. It is further placed within the subfamily Polistnae (paper wasps), which is the second largest of the subfamilies within Vespidae. Polistinae contains two main behavioral groups: swarm founding, involving a large numbers of workers and several queens, and independent founding, which involve a few workers and foundresses. (P. carolina uses the latter.)^[7]

Polistes carolina has been found to be most closely related to Polistes metricus. Recent phylogenetic analysis has shown that both Polistes carolina and Polistes metricus share a common ancestor with Polistes aurifer and Polistes fuscatus^[8]

Description and identification

Typical Polistes carolina (of both sexes)^[9] are about 25–32 millimetres (0.98–1.26 in) long^[1][2] with black wings of lengths ranging from 15–25 millimetres (0.59–0.98 in). Brown stripes are occasionally present on the abdomen.^[10] P. carolina is often confused with P. perplexus due to their strikingly similar reddish brown coloring.^[9] These two species are the only species of red wasps in the eastern United States.^[10] One distinguishing feature between these two red wasp species is the greater presence of black markings on the thorax of P. perplexus.^[10] Both sexes of the two species can also be differentiated by the coarser transverse ridging of the propodeum of P. perplexus when compared with P. carolina.^[9] Additionally, female P. carolina have mostly bare malar space (the distance between the lower eye orbit and the mouth).^[11]

Sexual dimorphism

Female Polistes carolina are usually completely ferruginous (rust in color) with the possibility of black markings forming spots around their eyes, lines on the dorsal surface of the scape, narrow lateral stripes on their scutum, or an incomplete median stripe on the propodeum. Bands on sternum 2 or terga 3 and 4 can also be present.^[9] Additional very restricted yellow markings can be observed on mandibles, clypeus, inner orbit, tergum 1, the outer surface of tibiae, and tarsi.^[9] Females also have more triangular faces with shorter antennae.^[12]

Male Polistes carolina often have more developed black or brown markings such as spots on the mid femur and sterna. Yellow markings vary but have been reported on the face and sternum 1 through 4. Additionally males have more squarish faces with longer hooked antenna.^[12]

Nests

Like most paper wasps Polistes carolina construct their nests by chewing plant and wood fibers with saliva to create a paper-mache like material.^[13] When dried, their nests form an upside-down umbrella or dome shape with exposed honeycomb-like cells, opening at the bottom.^[4] Polistes carolina prefer to nest in protected spaces,^[14] such as naturally occurring locations in vegetation or the cavities of trees.^[12] They also frequently nest in man made structures, such as the underside of bridges, roofs, eaves, and wooden boxes.^[12] These nests are some of the largest of any wasp species and usually contain around 3,000 to 5,000 members.^[15]

Distribution and habitat

Polistes carolina is most commonly found in the eastern United States from Nebraska to Texas and along the Atlantic coast from New York to Florida.^[10] It has also been recorded as an adventitious species in Ontario, Canada and was introduced to Bermuda.^[9]

Polistes carolina prefer to nest in protected areas such as hollow trees and are often observed in woodlands.^[9] However, given the opportunity they will also construct their nests among humans^[14] such as the undersides of roofs.^[12]

Colony cycle

Polistes colony cycle involves four separate phases which often overlap: the founding phase, the worker phase, the reproductive phase, and the intermediate phase.^[12]

The founding phase

The founding phase begins in the spring and involves young reproductive females (called foundresses) building new nests, either alone or in conjunction with other foundresses.^[12] In field studies "Polistes carolina" were observed to have a range of 1 to 8 foundresses in surviving colonies.^[14] During the founding period many foundresses move between nests, sometimes settling at another nest (movers) and sometimes returning to their own nest (visitors).^[14] In this way the foundress continues to reassess her reproductive options.^[16] During these visits, foundresses were also observed to lay eggs in other nests.^[14] While most nests are initiated by one foundress, they are usually joined by full sisters who become subordinates during this period.^[16]

The worker phase

During the worker phase in many Polistes species, adult workers and early males are enclosed.^[12] Polistes carolina, however, lack early males during this time and instead only produce worker females.^[14] As workers emerge, they begin to assume colony tasks, such as nest maintenance, foraging, and larva care.^[12]

The reproductive phase

The reproductive phase lasts from the emergence of the first reproductives until the colony begins to decline and new reproductives disperse to form their own nests.^[12] During this time each foundress mates with a different male and lays her eggs, with the dominant foundress laying the majority of the eggs.^[14]

The intermediate phase

The time between colony decline and the founding of new colonies. During this time the initial colony begins to disperse as new reproductives search for locations to initiate their own nests.^[12] The foundresses of the colony often disappear during this time as males accumulate in the nest.^[12]

Behavior

Dominance hierarchy

Foundress associations in Polistes species establish clear dominant and subordinate relationships in which the dominant gains the most reproduction success. Unlike in many species, the queen (most dominant foundress) is not necessarily the largest female. Rather, the first foundress to arrive becomes the queen and any subsequent joiners become subordinates.^[14] While the queen may not be the largest foundress, field studies have shown that the dominant foundress usually has the largest ovaries of all foundresses present at the nest.^[14] Precedence rather than size hierarchy indicates that the earliest foundress may simply be the fittest. This is supported by evidence that the first female to emerge from hibernation has the most developed corporar allata (the site of juvenile hormone synthesis) and high juvenile hormone synthesis is correlated with dominance and ovarian development in Polistes. ^[14] This convention could also be a method to prevent fighting amongst near equals, thus decreasing the risk of injury for both dominants and subordinates.^[14]

Adopting a nest

During the Founding Phase of the colony cycle, many foundresses leave their nest and will join another. Foundresses that move most frequently join the nest of a full sister yet do not increase their reproductive rank upon joining a new nest.^[14] The moving foundress usually lays her eggs at her previous nest so joining a new nest is a surprising behavior as the offspring will not be directly related to her.^[16] These moving foundresses help care for the offspring which has been shown to be very important for the survival of the colony. Colonies with subordinate foundresses have a much higher success rate and higher productivity than solitary colonies.^[14]

Reproductive suppression

The queen dominates reproduction, but not exclusively. Queens produce about 60 percent of total offspring and several subordinates produce the remaining 40 percent of offspring. Only about 20 percent of subordinates associated with any given nest do not participate in reproduction.^[14] According to skew theory, as resources become more scarce the queen will concede less of her reproductive rights to subordinates (reproduction will become more skewed toward the queen) since subordinates have more limited nesting options. Additionally, Polistes carolina does not participate in egg eating, which supports the theory that aggression is not the primary method of determining reproductive share.^[14]

Mating behavior

In Polistes carolina several foundresses will mate; however, each foundress only mates once.^[14] There is an absence of early males in Polistes carolina, which is unique when compared to other Polistes species. This lack of early males however, indicates that female workers do not become inseminated queens in this species^[14] as in common in other wasp species.^[17]

Genetic relatedness of colonies

Polistes carolina like many eusocial insects follow a haplodiploidy sex determination system, meaning males are haploid and develop from unfertilized eggs while females are diploid and develop from fertilized eggs.^[18] This means that daughters will share one identical allele from their haploid father and receive the other allele from their diploid mother's two alleles. This knowledge allows us to determine the relatedness among individuals via algorithms and computer programs such as Relatedness 4.2 and Kinship 1.1.2.^[14] Due to this system of reproduction, genetic relatedness among nest mate foundresses is about 0.75.^[14] Mating males, however, are not related to other males, nor are they related to the females they mated with.^[14]

Kin recognition and discrimination

Polistes carolina demonstrate altruistic behavior in their kin feeding. Many Polistes females will feed their own progeny preferentially in order to increase their survival success. Polistes carolina, however, do not recognize or preferentially care for their own progeny.^[16] This behavior could also account for the lack of conflict often observed in this species.^[19]

Feeding

Diet

Polistes carolina feed mostly on caterpillar meat and nectar fluids.^[19] They have also been observed to prey on Chrysomelidae larvae and cicada.^[9]They have even been found to eat sweet food discarded by humans.^[4]

Feeding behavior

Female Polistes carolina feed offspring by putting her head into the cell containing the larvae. Feeding of larvae can involve visible transfers of food, such as caterpillar meat held in their jaws or non visible transfers of nectar fluids.^[19]

Human importance

Since Polistes carolina nest in sheltered areas, it is common for them to construct their nests in close proximity to humans, such as the open space under a roof. They are also often attracted to humans due to loud noises, bright colors, and sweet smells, such as food and perfume.^[4] Typical paper wasps are relatively unaggressive, only attacking humans and animals if they or their nests are being threatened. Red Paper wasps are more aggressive and their stings can be more painful;^[15] however, only females have the ability to sting.^[20] Unlike bees, wasps do not lose their stinger and thus they are able to sting multiple times.^[4]

Stings

Red wasp stingers do not remain in their victim. A red wasp sting is often painful and causes localized swelling and itchiness. In some cases, people report developing a full-body itchy rash.^[4] Applying a cold compress or ice to the area can help relieve swelling, while application of calamine lotion or 1 percent hydrocortisone cream can help relieve itching and pain. If neither of these creams are available, a paste made from baking soda and water is a very effective sting remedy. The paste should be thick enough to stay on the area of the sting until it dries.^[15] Some individuals are known to have severe allergic reactions, including anaphylaxis, to wasp stings and should seek immediate medical attention if severe symptoms begin to present.^[15] In very rare cases, children may develop shock after a red wasp sting.^[4]

References

External links

• Bug Eric Blog
• BugGuide