Polistes erythrocephalus

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Polistes erythrocephalus
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. erythrocephalus
Binomial name
Polistes erythrocephalus
(Latreille, 1813) [1]
Synonyms

Polistes canadensis erythrocephalus [2]

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. [3] [4] The cooperation between the two castes to raise young demonstrates the altruistic nature of these wasps. [5] 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.

Contents

Taxonomy and phylogeny

Polistes scholars such as K. Yoshikawa, Joseph Charles Bequaert, and Mary Jane West-Eberhard, originally considered Polistes erythrocephalus to be a subspecies of Polistes canadensis. For this reason, much of the research done on P. erythrocephalus has been classified under P. canadensis. Additionally, it was left out of Yoshikawa's list of Polistes species around the world in 1963. [2] More recently however P. erythrocephalus has been recognized as a separate species by Owain Richards though it is sometimes still referred to as Polistes canadensis erythrocephalus. [6] Its closest relatives are Polistes canadensis, Polistes annularis , and Polistes infuscatus. [7]

Description and identification

Polistes erythrocephalus is about 21 millimetres (0.83 in) in length and is characterized by its black thorax and abdomen. P. erythrocephalus bodies are typically narrower than those of its closest relatives such as Polistes annularis. The thorax is unmarked and the abdomen is slightly shinier than the rest of the body. This black body color contrasts with the color of its head which is a dull ferruginous (rust) color. [8] On top of the head are ferruginous colored antenna which grow paler at the tips. P. erythrocephalus wings are dark bluish in color and have a noticeable glossy sheen. P. erythrocephalus legs are black with yellowish hues at the tarsi and knee joints. [9]

For a long time P. erythrocephalus was considered to be a subspecies of P. canadensis and thus the two were often confused. However, P. canadensis possesses a ferruginous thorax, abdomen, and head. Additionally, the wings of P. canadensis are often a much lighter color than its body. [2]

Sexual dimorphism

Male and female P. erythrocephalus are hard to distinguish unless they are observed at a close distance. Under close examination the abdomen of female P. erythrocephalus has six banded segments while male abdomens contain seven. [10] In addition the females abdomen narrows to a point with a stinger at the end. The male abdomen on the other hand is more rounded and does not contain a stinger. Female antennas also have 12 segments and are hooked at the end while males have 13 segments and have a more subtle curve. [5] [10] It has been suggested that such sexual dimorphic traits within a species occur to make females more recognizable. This recognition makes it easier to kick males out of nests so that they can disperse and encounter more foundresses. Sexual dimorphism also allows males to more quickly recognize females, thus, providing the benefit of not having to waste time accidentally trying to mate with other males. [5]

Nests

Like most paper wasps Polistes erythrocephalus construct their nests by chewing plant and wood fibers with saliva to create a paper-maché-like material. [11] When dried, their nests are somewhat horizontal and due to the divergence of cells have a concave upper surface. [12] P. erythrocephalus nesting habits are very similar to P. annularis in that they prefer to build their nests in areas protected from direct sunlight, rain, wind, and other insects. [2] The locations in which they find this protection vary greatly from the countryside to the city. P. erythrocephalus in urban areas have been observed to build nests under the eaves of roofs, under the wood of basement floors, in barns/stables, under bridges, and in lofts. This species also appears to prefer to build nests in close proximity to a water source. [10] In more rural areas nests are found in clear areas such as the trunk of a solitary tree or the underside of a large Heliconia leaf. In these cases several nests can be built on the same tree at heights ranging from three to twenty feet off the ground. Again these nests are often in close proximity to a water source such as a stream. [2]

Distribution and habitat

Polistes erythrocephalus is found in Central and South America. Although this species is most concentrated in Costa Rica and Panama, it has also been observed in Nicaragua, Venezuela, Colombia, Ecuador, Peru, Bolivia, and Southern Brazil. [1] [2] [13] In Costa Rica, P. erythrocephalus nests are specifically highly concentrated in Finca Taboga and on the Osa Peninsula. In these countries P. erythrocephalus build nests in protected yet open areas both among humans and in the wild. [2]

Life cycle

The average P. erythrocephalus life cycle is 110 days with an observed range of 91–131 days. P. erythrocephalus go through the common Polistes maturation phases of eggs, larva, pupa, and adult. The egg stage is typically 14 days, the larva stage 29, the pupa stage is about 22 and the adult stage continues until death, which is on average 45 days. [10]

Colony cycle

Polistes erythrocephalus colony cycle follows that of many other Polistes species in that it involves four separate phases. the founding phase, the worker phase, the reproductive phase, and the intermediate phase. The length of a colony cycle for P. erythrocephalus varies depending on if the particular group is located in a tropical or more temperate region. The colony cycle of tropical P. erythrocephalus is six to seven months long. This is one to three months longer than the typical colony cycle of P. erythrocephalus in more temperate areas. This varying colony dissolution time represents an optimization of the tradeoffs of predation before reproduction and the number of reproductives produced should the colony continue to survive. [5]

The founding phase

The founding phase begins in the spring and involves individual reproductive females (called foundresses) building new nests. [5] While one female founds the nest she is often joined by other foundresses several days later. In field studies, P. erythrocephalus were observed to have an average of 4.9 foundresses with a range of 1 to 10. If a foundress is not joined by other females within two weeks of starting the nest she will abandon her nest. [6] During the founding phase the foundress composition of any given nest changes daily as foundresses move from nest to nest. [5] In this way the foundress continues to reassess her reproductive options. [14]

The worker phase

During the worker phase in many Polistes species adult workers and early males are eclosed (emerge as adults from pupae). As workers emerge, they begin to assume colony tasks such as nest maintenance, foraging, and larva care. A typical P. erythrocephalus nest grows to contain around 95 cells. If a colony loses its queen or if she stops ovipositing, then all construction on the nest will stop. This is because the queens are the "primary initiators." While the workers may do much of the building and maintenance, queens with mature eggs in their ovaries demonstrate cell initiation behavior and are responsible for beginning the process. [5]

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. During this time males wait on perches near nests in order to have the opportunity to mate with a virgin foundress. In some cases males will defend their perch in order to ensure the continuation of their genes. In the reproductive phase workers provide larvae with food. This causes hierarchies among workers to emerge as more dominant individual workers forage significantly less than subordinates. [5]

The intermediate phase

In P. erythrocephalus the termination of nest growth and start of brood decline occurs when the queen ceases laying eggs or disappears. This demonstrates that the colony cycle of P. erythrocephalus is a function of the queen's reproductive cycle. [15] During 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. The foundresses of the colony often disappear as males accumulate in the nest. [5] Adults often remove paper from old nests and recycle it to use for new ones. [7]

Behavior

Caste importance

As is common in many wasp species the queen of P. erythrocephalus nests has great influence over the behavior of the colony. The presence of an active reproductive queen in P. erythrocephalus is a necessity for normal colony expansion and nest growth. Under nutritional stress when an active queen is removed and a former inactive queen is substituted the brood will decline. However, under normal conditions when a queen is removed the nest can continue to grow (at a reduced rate) if there are workers present. These experiments demonstrate that the simultaneous presence of an active queen and an effective caste of workers have a huge effect on the success of the nest and colony. While it is often thought that a colony begins to decline with the cessation of the queen, this removal is often accompanied with an absence of workers. These two factors combined lead to colony decline. [15]

Sex ratios

Polistes erythrocephalus nests of less than 50 cells contain no males. This absence confirms that only females accompany the reproductive queen (foundress) in the forming of a new nest. In nests of greater than 50 cells males begin to appear indicating that they are the progeny of the founding queen. The number of males in a nest increases as the number of cells increases until there is an equal ratio of males and females. As the number of cells in a nest increases so does the number of adults associated with the nest. While the number of females tends to increase proportionally more in smaller nests the sex ratio equals out in nests of 300 cells of more. [10]

Reproductive competition

As in many multifoundress wasp species, P. erythrocephalus foundresses compete viciously in order to become the dominant female. These interactions include chasing, biting, lunging, "sting threats," and aggressive mounting. Fights between individuals can sometimes even be fatal. These competitions lead to only one reproductively dominant female while the rest become subordinates. Once reproductive dominance is achieved then that female asserts great control over the behavior of the nest. [5]

Feeding

Diet

P. erythrocephalus are predatory wasps and feed on a wide variety of larvae. A colony's predatory capacity depends on the number of their own larvae present in their nest rather than the number of adults present. They are known to prey upon hornworm larvae and were observed to eat an average of 0.5 hornworm larvae per day (maximum of 1.3 and a minimum of 0.08). [10]

Starvation resistance

The starvation resistance of P. erythrocephalus was tested on both adults and larvae. Adults P. erythrocephalus demonstrated average resistance and died in two to three days when cut off from food and water. The larvae of P. erythrocephalus however proved to be extremely starvation resistant. In one nest six larvae were found alive after 26 days without any food or water. After this time one of the larvae was minced and fed to the remaining larvae which ate it readily. This demonstrates that the larvae will behave cannibalistically when necessary. [2]

Interactions with other species

Predators, parasites and symbionts

One of the most devastating predators of P. erythrocephalus are army ants such as Eciton burchellii. These ants attack wasp nests and consume the larva and pupa, often destroying the nest in the process. [16] P. erythrocephalus has not been observed to have any ability to defend their nests from such predators. However, they will attack and sting larger slow moving threats such as humans if they get within 1–2 meters of a nest. [16]

The most common symbiont found in P. erythrocephalus nests is Pachysomoides stupida, a type of parasitic ichneumon. Papery cocoons of Pachysomoides stupida are found in many P. erythrocephalus nests while adults are seldom found. [2] Other parasites of P. erthrocephalus include some species of Oxysarcodexia or flesh flies and Brachymeria conica ( Brachymeria is a parasitic wasp genus). Both these species parasitize the larva and pupa of P. erythacephalus. [10] Additionally, some Xenon species parasitize P. erythrocephalus adults, acting as permanent entomophagous endoparasites by dwelling in the abdomen. [10]

Microlepidoptera (types of smaller moths) larvae have been observed to feed on the excretory waste of wasp larvae. [2]

Honeydew collection

Polistes eythrocephalus is not commonly observed to tend or collect honeydew from aetalionids (treehoppers) or any other insect species. Honeydew is a good source of carbohydrates, composed of a mixture of oligosaccharides. This mixture provides those that eat it with an energy source higher than that of floral nectar. However, in one study done in Peru researchers observed a single female P. erythrocephalus among an aggregation of Aetalion reticulatum (treehoppers). The female P. erythrocephalus stood among or below the treehoppers and touched them with her antennae. Though they did not directly feed her, the honeydew accumulated below where the A. reticulatum were standing. P. erythrocephalus then collected the honeydew from the branch. This research provides the first bit of evidence that P. erythrocephalus may tend to aetalionids within its range. [17]

Recycling nests

A number of species take advantage of abandoned nests. An unidentified species of Trypoxylon (a mud-daubing sphecid) uses the cells in order to raise its own brood. Moths of family Phycitidae use the cells as a safe place to form a cocoon. Spiders of families such as Salticidae, Linyphiidae, Oonopidae, and Scytodidae have been observed to use the nests for brooding and resting webs. [2]

Human importance

As predatory wasps P. erythrocephalus is a natural pest controller. They have long been considered responsible for feeding on and thus controlling the population of leaf-eating worms in tobacco fields in South America. [10] They were especially important to the large plantations of the Colombian Tobacco Company (la Compañia Colombiana de Tabaco) from 1932-1936. [10]

In experiments P. erythrocephalus were introduced to Bacillus thuringiensis which is a bacterium with insecticidal properties. Bacillus thuringiensis however proved harmless to P. erythrocephalus as nest concentrations and working conditions remained completely unchanged. [10]

Related Research Articles

<i>Polistes</i> Genus of insects

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>Ropalidia marginata</i> Species of insect

Ropalidia marginata is an Old World species of paper wasp. It is primitively eusocial, not showing the same bias in brood care seen in other social insects with greater asymmetry in relatedness. The species employs a variety of colony founding strategies, sometimes with single founders and sometimes in groups of variable number. The queen does not use physical dominance to control workers; there is evidence of pheromones being used to suppress other female workers from overtaking queenship.

<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. Polistes metricus is dark colored, with yellow tarsi and black tibia. Nests of Polistes 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.

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

Polistes carolina is one of two species of red paper wasp found in the eastern United States 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. 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.

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.

<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>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>Mischocyttarus flavitarsis</i> Species of wasp

Mischocyttarus flavitarsis is a social paper wasp found in western North America. Their nests can be found both in forests close to rivers or in close proximity to human life under the eaves of roofs. Despite the fact that M. flavitarsis nests are frequently in close contact with humans, M. flavitarsis typically will not sting, but rather ram into the threatening individual. Their colony cycle typically begins before May and will last until October. The queen will then seek a hibernation site for the winter. Perched near female hibernation sites are males with whom the female will mate. The males have claimed their territory by rubbing sternal brushes along the border of the site, leaving a chemical that deters other individuals from approaching. M. flavitarsis feed on arthropods, nectar, and animal carcasses and are often prey to birds, ants, and praying mantis.

<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.

References

  1. 1 2 Carpenter, James M. (1996). "Distributional checklist of species of the genus Polistes (Hymenoptera, Vespidae, Polistinae, Polistini)". American Museum Novitates (3188): 1–39. hdl:2246/3649.
  2. 1 2 3 4 5 6 7 8 9 10 11 Nelson, John M. (1971). "Nesting habits and nest symbionts of Polistes erythrocephalus Latreille (Hymenoptera Vespidae) in Costa Rica" (PDF). Revista de Biología Tropical. 18: 89–98.
  3. ↑ Wilson, Edward O. (1971). The Insect Societies . Cambridge. Massachusetts: Belknap Press of Harvard University Press. ISBN   9780674454903.
  4. ↑ Wilson, Edward O.; Bert Hölldobler (20 September 2005). "Eusociality: origin and consequences". Proceedings of the National Academy of Sciences of the United States of America. 102 (38): 13367–13371. Bibcode:2005PNAS..10213367W. doi: 10.1073/pnas.0505858102 . PMC   1224642 . PMID   16157878.
  5. 1 2 3 4 5 6 7 8 9 10 Ross, Kenneth G. (1991). The Social Biology of Wasps. Cornell University Press. p. 104. ISBN   0-8014-9906-2.
  6. 1 2 Jeanne, Robert (1978). "Construction and utilization of multiple combs in Polistes canadensis in relation to the biology of a predaceous moth". Behavioral Ecology and Sociobiology. 4 (3): 293–310. doi:10.1007/bf00297649. S2CID   36132488.
  7. 1 2 Young, Allen M. (October 1986). "Natural history notes on the social paper wasp Polistes erythrocephalus Latreille (Hymenoptera: Vespidae: Polistinae) in Costa Rica". Journal of the Kansas Entomological Society. 59 (4): 712–722. JSTOR   25084849.
  8. ↑ Erythrocephalus means "red-headed".
  9. ↑ Bequaert, Joseph C. (1943). "Color variation in the American social wasp Polistes canadensis (Linnaeus), with descriptions of two new forms (Hymenoptera, Vespidae)". Boletín de Entomología Venezolana. 2: 107–124.
  10. 1 2 3 4 5 6 7 8 9 10 11 Martin P., Carlos A.; Bellotti, Anthony C. (1986). "Biología y comportamiento de Polistes erythrocephalus Ltr. (Hyrnenoptera: Vespidae), predador del "Gusano cachón" de la yuca Erinnyis ello L. (Lepidoptera: Sphingidae)". Acta Agronómica (in Spanish). 36 (1): 63–76.
  11. ↑ "Paper Wasp". Everything About. Retrieved 2014-09-23.
  12. ↑ West-Eberhard, Mary Jane (December 3, 1969). "The social biology of polistine wasps". Miscellaneous Publication, Museum of Natural History, University of Kansas. 140: 1–101. Retrieved 16 November 2014.
  13. ↑ "Fauna entomologica de Nicaragua: Polistes (Aphanilopterus) erythrocephalus LATREILLE, 1813". BIO-NICA. Retrieved 2014-10-14.
  14. ↑ Seppa, Perttu; David C. Queller; Joan E. Strassman (September 25, 2012). "Why wasp foundresses change nests: relatedness, dominance, and nest quality". PLOS ONE. 7 (9): e45386. Bibcode:2012PLoSO...745386S. doi: 10.1371/journal.pone.0045386 . PMC   3458021 . PMID   23049791.
  15. 1 2 Malcolm, Keeping G. (11 December 1996). "Social behavior and brood decline in reproductive-phase colonies of Belonogaster petiolata (Degeer) (Hymenoptera: Vespidae)". Journal of Insect Behavior. 10 (2): 265–278. doi:10.1007/BF02765559. S2CID   41829420.
  16. 1 2 Young, Allen M. (1979). "Attacks by the army ant Eciton burchellii on nests of the social paper wasp Polistes erythrocephalus in northeastern Costa Rica". Journal of the Kansas Entomological Society. 52 (4): 759–768. JSTOR   25083991.
  17. ↑ MacCarroll, M.; Reeves, W. K. (2004). "Attendance of Aetalion reticulatum (Hemiptera: Aetalionidae) by Polistes erythrocephalus (Hymenoptera: Vespidae) in Peru". Entomological News. 115: 52–53. Retrieved 16 October 2014.
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