Polistes carolina

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Polistes carolina
Red Paper Wasp (Vespidae, Polistes carolina (Linnaeus)) (35549913493) (cropped).jpg
Female P. carolina from Kendall County, Texas
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Vespidae
Subfamily: Polistinae
Tribe: Polistini
Genus: Polistes
Species:
P. carolina
Binomial name
Polistes carolina
(Linnaeus, 1767) [1]
Synonyms [2]
  • Polistes carolinus(Linnaeus, 1767)
  • Vespa carolinaLinnaeus, 1767

Polistes carolina is one of two species of red paper wasp found in the eastern United States (the other being Polistes rubiginosus ) and is noted for the finer ridges on its propodeum. It is a social wasp in the family Vespidae and subfamily Polistinae. The species is native to the United States from Texas to Florida, north to New York, and west to Nebraska. [3] The wasp's common name is due to the reddish-brown color of its head and body. P. carolina prefer to build their nests in protected spaces.

Contents

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. [4] In this volume, he referred to the species as Vespa carolina. [4] Henri Louis Frédéric de Saussure later moved it to the genus Polistes in 1855 after Pierre Andre Latreille coined the new genus in 1802. [5] 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 Polistinae (paper wasps), which is the second-largest of the subfamilies within the Vespidae. The Polistinae contain two main behavioral groups: swarm founding, involving a large numbers of workers and several foundresses, and independent founding, which involve a few workers and foundresses. (P. carolina uses the latter.) [6]

P. carolina has been found to be most closely related to P. metricus. Recent phylogenetic analysis has shown that both P. carolina and P. metricus share a common ancestor with P. aurifer and P. fuscatus . [7]

Description and identification

Male P. carolina photographed in Austin, Texas Red paper wasp, male (Vespidae, Polistes carolina) (30901479432) (cropped).jpg
Male P. carolina photographed in Austin, Texas

Both sexes of P. carolina are about 25–32 mm (0.98–1.26 in) long with black wings of lengths ranging from 15–25 mm (0.59–0.98 in). [1] [3] P. carolina is often confused with P. rubiginosus due to its strikingly similar reddish-brown coloring. [3] [2] Females can be separated by the bare genae of P. carolina in contrast to the silvery pubescence on the genae P. rubiginosus. Both sexes can also be differentiated by the coarser transverse ridging of the propodeum of P. rubiginosus when compared with P. carolina. [3] [2] Additionally, female P. carolina specimens have mostly bare malar spaces (the distance between the lower eye orbit and the mouth). [8]

Sexual dimorphism

Females of 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 scuta, or an incomplete median stripe on their propodea. Bands on sternum 2 or terga 3 and 4 can also be present. [3] Additional very restricted yellow markings can be observed on mandibles, clypei, inner orbits, terga 1, the outer surfaces of the tibiae, and tarsi. [3] Females also have more triangular faces with shorter antennae. [9]

Males often have more developed black or brown markings such as spots on the midfemur and sterna. Yellow markings vary, but have been reported on the face and sterna 1 through 4. Additionally, males have more squarish faces with longer hooked antenna. [9]

Nests

Like most paper wasps, P. carolina constructs nests by chewing plant and wood fibers with saliva to create a paper-like material. When dried, their nests form an upside-down umbrella or dome shape with exposed honeycomb-like cells, opening at the bottom. [1] P. carolina prefers to nest in protected spaces, [10] such as naturally occurring locations in vegetation or the cavities of trees. [9] They also frequently nest in man-made structures, such as the underside of bridges, roofs, chimneys, and eaves. [9] [11]

Distribution and habitat

P. carolina is most commonly found in the eastern United States (along with another "red paper wasp", Polistes rubiginosus) from Nebraska to Texas and along the Atlantic coast from New York to Florida. [3] It has also been recorded as an adventitious species in Ontario, Canada, and was introduced to Bermuda. [3]

It prefers to nest in protected areas such as hollow trees and is often observed in woodlands. [3] However, given the opportunity, it will also construct nests near humans, [10] such as the undersides of roofs. [9]

Colony cycle

The Polistes colony cycle involves four separate phases which often overlap: the founding phase, the worker phase, the reproductive phase, and the intermediate phase. [9]

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. [9] In field studies, P. carolina was observed to have a range of one to eight foundresses in surviving colonies. [10] During the founding period, many foundresses move between nests, sometimes settling at another nest (movers) and sometimes returning to their own nest (visitors). [10] In this way, the foundress continues to reassess her reproductive options. [12] During these visits, foundresses were also observed to lay eggs in other nests. [10] While most nests are initiated by one foundress, they are usually joined by full sisters which become subordinates during this period. [12]

The worker phase

During the worker phase in many Polistes species, adult workers and early males are enclosed. [9] P. carolina, however, lacks early males during this time and instead only produces worker females. [10] As workers emerge, they begin to assume colony tasks, such as nest maintenance, foraging, and larval care. [9]

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. [9] During this time, each foundress mates with a different male and lays her eggs, with the dominant foundress laying the majority of the eggs. [10]

The intermediate phase

The time between colony decline and the founding of new colonies, the initial colony begins to disperse as new reproductives search for locations to initiate their own nests. [9] The foundresses of the colony often disappear during this time, as males accumulate in the nest. [9]

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. [10] 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. [10] 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. [10] This convention could also be a method to prevent fighting amongst near equals, thus decreasing the risk of injury for both dominants and subordinates. [10]

Adopting a nest

During the founding phase of the colony cycle, many foundresses leave their nests and 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. [10] 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. [12] 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. [10]

Reproductive suppression

The foundress dominates reproduction, but not exclusively. Foundresses produce about 60% of total offspring and several subordinates produce the remaining 40%. Only about 20% of subordinates associated with any given nest do not participate in reproduction. [10] According to skew theory, as resources become more scarce, the queen concedes less of her reproductive rights to subordinates (reproduction becomes more skewed toward the queen) since subordinates have more limited nesting options. Additionally, P. carolina does not participate in egg eating, which supports the theory that aggression is not the primary method of determining reproductive share. [10]

Mating behavior

In P. carolina, several foundresses mate; however, each foundress only mates once. [10] The absence of early males in P. carolina 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 [10] as is common in other wasp species. [13]

Genetic relatedness of colonies

P. carolina, like many eusocial insects, follows a haplodiploidy sex determination system, meaning males are haploid and develop from unfertilized eggs, while females are diploid and develop from fertilized eggs. [14] Daughters share one identical allele from their haploid father and receive the other allele from their diploid mother's two alleles. This knowledge allows determination of the relatedness among individuals via algorithms and computer programs such as Relatedness 4.2 and Kinship 1.1.2. [10] Due to this system of reproduction, genetic relatedness among nestmate foundresses is about 0.75. [10] Mating males, however, are not related to other males, nor are they related to the females with which they mated. [10]

Kin recognition and discrimination

P. carolina demonstrates altruistic behavior in kin feeding. Many Polistes females feed their own progeny preferentially to increase their survival success. P. carolina, however, does not recognize or preferentially care for its own progeny. [12] This behavior could also account for the lack of conflict often observed in this species. [15]

Feeding

Diet

P. carolina wasps feed mostly on caterpillars as larvae and nectar fluids as adults. [15] They have also been observed to prey on Chrysomelidae larvae and cicadas. [3] Females feed offspring by putting their heads into the cells containing the larvae. Feeding of larvae can involve visible transfers of food, such as caterpillar flesh held in their jaws or invisible transfers of nectar fluids. [15]

Human importance

Since P. carolina nests in sheltered areas, it commonly constructs nests in close proximity to humans, such as the open space under a roof. Typically, paper wasps are relatively unaggressive, only attacking humans and animals if they or their nests are being threatened. As in other aculeate wasps, only females have the ability to sting. [16] Unlike bees, wasps do not have barbed stingers that can be lost, so they are able to sting multiple times to defend a nest. [17]

Related Research Articles

<i>Polistes</i> Genus of wasps

Polistes is a cosmopolitan genus of paper wasps and the only genus in the tribe Polistini. Vernacular names for the genus include umbrella wasps, coined by Walter Ebeling in 1975 to distinguish it from other types of paper wasp, in reference to the form of their nests, and umbrella paper wasps. Polistes is the single largest genus within the family Vespidae, with over 200 recognized species. Their innate preferences for nest-building sites leads them to commonly build nests on human habitation, where they can be very unwelcome; although generally not aggressive, they can be provoked into defending their nests. All species are predatory, and they may consume large numbers of caterpillars, in which respect they are generally considered beneficial.

<span class="mw-page-title-main">European paper wasp</span> Species of wasp

The European paper wasp is one of the most common and well-known species of social wasps in the genus Polistes. Its diet is more diverse than those of most Polistes species—many genera of insects versus mainly caterpillars in other Polistes—giving it superior survivability compared to other wasp species during a shortage of resources.

<i>Polistes gallicus</i> Species of wasp

Polistes gallicus is a species of paper wasp found in various parts of Europe, excluding England, Denmark, and Scandinavia, from warmer climates to cooler regions north of the Alps. Nests of these social insects are created in these various conditions. The Polistes species use an oral secretion to construct their nests, which consist of a combination of saliva and chewed plant fibers. This structural mixture physically protects the nest from various harsh elements and from weathering over time.

<i>Polistes chinensis</i> Species of wasp

Polistes chinensis is a polistine vespid wasp in the cosmopolitan genus Polistes, and is commonly known as the Asian, Chinese or Japanese paper wasp. It is found in East Asia, in particular China and Japan. The subspecies P. chinensis antennalis is an invasive species in New Zealand, having arrived in 1979.

<i>Polistes annularis</i> Species of wasp

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.

<i>Polistes metricus</i> Species of wasp

Polistes metricus is a wasp native to North America. In the United States, it ranges throughout the southern Midwest, the South, and as far northeast as New York, but has recently been spotted in southwest Ontario. A single female specimen has also been reported from Dryden, Maine. P. metricus is dark colored, with yellow tarsi and black tibia. Nests of P. metricus can be found attached to the sides of buildings, trees, and shrubbery.

<i>Polistes fuscatus</i> Species of insect

Polistes fuscatus, whose common name is the dark or northern paper wasp, is widely found in eastern North America, from southern Canada through the southern United States. It often nests around human development. However, it greatly prefers areas in which wood is readily available for use as nest material, therefore they are also found near and in woodlands and savannas. P. fuscatus is a social wasp that is part of a complex society based around a single dominant foundress along with other cofoundresses and a dominance hierarchy.

Belonogaster juncea juncea is a subspecies of Belonogaster juncea and is classified as a primitively eusocial wasp, meaning that the species is social while exhibiting a morphology that is indistinguishable from that of other castes. It is also classified as a type of African Paper Wasp. Many of the studies relating specifically to B. j. juncea take place at the University of Yaoundé in Cameroon.

Ropalidia fasciata, a common paper wasp, is a wide-ranging species that is distributed from India to the Lesser Sunda Islands, Palawan, and Ryukyu Islands, occupying the northern edge of Ropalidia's larger distribution. These primitively eusocial wasps are unique in that they do not exhibit the strict matrifilial, single-queen social structure found in many species of social insects. Instead, colonies are founded based on associations between several females, or 'foundresses'.

<i>Polistes nimpha</i> Species of wasp

Polistes nimpha is a eusocial paper wasp found all over Europe, with particular sightings in Turkey, Finland, Estonia, and Latvia. It is also found in northern Africa, Pakistan, Iran, India, Kazakhstan, Mongolia, and China. The climate in these areas is relatively cold and snowy in the winter, while summers are usually hot and dry, with steppe vegetation. Polistes nimpha colonies are relatively small and easily manipulated.

<i>Polistes bellicosus</i> Species of insect

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.

<i>Polistes canadensis</i> Species of wasp

Polistes canadensis is a species of red paper wasp found in the Neotropical realm. It is a primitively eusocial wasp as a member of the subfamily Polistinae. A largely predatory species, it hunts for caterpillar meat to supply its colony, often supplementing its developing larvae with nectar. The most widely distributed American species of the genus Polistes, it colonizes multiple combs, which it rears year-round.

<i>Polistes atrimandibularis</i> Species of wasp

Polistes atrimandibularis is one of four obligate social parasites among the Polistes wasps found in Europe. Of the four social paper wasp parasite species known, it is the smallest. It parasitizes multiple species such as P. dominula, P. nimpha, P. associus, P. gallicus, and P. biglumis. Females of P. atrimandibularis are unable to build a nest or produce workers, and therefore rely entirely on the host colony.

<i>Belonogaster petiolata</i> Species of wasp

Belonogaster petiolata is a species of primitively eusocial wasp that dwells in southern Africa, in temperate or subhumid climate zones. This wasp species has a strong presence in South Africa and has also been seen in northern Johannesburg. Many colonies can be found in caves. The Sterkfontein Caves in South Africa, for example, contain large populations of B. petiolata.

<i>Polistes biglumis</i> Species of wasp

Polistes biglumis is a species of social wasp within Polistes, the most common genus of paper wasp. It is distinguished mainly by its tendency to reside in montane climates in meadows or alpine areas. Selection pressure from the wasp's environment has led to several idiosyncrasies of its behavior and lifecycle with respect to its relative species in the genus Polistes. It alone among paper wasps is often polyandrous. In addition, it has a truncated nesting season that gives rise to unique competitive dynamics among females of the species. P. biglumis wasps use an odor-based recognition system that is the basis for all wasp-to-wasp interaction of the species. The wasp's lifecycle is highly intertwined with that of Polistes atrimandibularis, an obligate social parasite wasp that frequently invades the combs of P. biglumis wasps.

<i>Polistes semenowi</i> Species of wasp

Polistes semenowi is a species of paper wasp in the genus Polistes that is found in southeastern and southern central Europe, as well as central Asia, and was until 2017 erroneously known by the name Polistes sulcifer, while a different species was incorrectly believed to represent P. semenowi. It is one of only four known Polistes obligate social parasites, sometimes referred to as "cuckoo paper wasps", and its host is the congeneric species Polistes dominula. As an obligate social parasite, this species has lost the ability to build nests, and relies on the host workers to raise its brood. P. semenowi females use brute force, followed by chemical mimicry in order to successfully usurp a host nest and take over as the queen.

<i>Polistes japonicus</i> Species of wasp

Polistes japonicus is a eusocial paper wasp found in Japan. It was first described by Henri Louis Frédéric de Saussure in 1858. It is closely related to Polistes formosanus. This species lives in small colonies with few workers and a foundress queen. Nests of these wasps are sometimes used as a traditional medicine in Korea, China, and Japan.

<i>Polistes versicolor</i> Species of wasp

Polistes versicolor, also known as the variegated paper wasp or yellow paper wasp, is a subtropical social wasp within Polistes, the most common genus of paper wasp. It is the most widely distributed of South American wasp species and is particularly common in the Southeastern Brazilian states. This social wasp is commonly referred to as the yellow paper wasp due to the distinct yellow bands found on its thorax and abdomen. The P. versicolor nest, made of chewed vegetable fiber, is typically a single, uncovered comb attached to the substrate by a single petiole. The yellow wasp is frequently found in urban areas. New nests and colonies are usually founded by an association of females, sometimes in human buildings.

<i>Polistes dorsalis</i> Species of wasp

Polistes dorsalis is a species of social wasps that can be found throughout various parts of North America. It is classified under the Vespidae within the genus of Polistes. Male Polistes dorsalis wasps can be distinguished from other Polistes species by their distinctly prominent median tubercle of sternum 7. Both sexes can also be recognized due to their v- shaped yellow markings on their head. They are distributed widely across North America and can be found in sheltered nests, typically closer to the ground. These wasps live in a dominance hierarchy in which the queen's role differs from that of ordinary workers. When threatened, these wasps can deliver moderately painful stings. Their venom might also be of human interest for their antimicrobial uses.

Polistes erythrocephalus is a species of paper wasp in the subfamily Polistinae of family Vespidae found in Central and South America. P. erythrocephalus is a eusocial wasp, meaning that it possesses both reproductive and non-reproductive castes. The cooperation between the two castes to raise young demonstrates the altruistic nature of these wasps. P. erythrocephalus exhibits a four-stage colony cycle, as do many other Polistes wasps. This species generally feeds on larvae, occasionally their own, and is preyed upon by species such as army ants.

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