Polistes nimpha

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Polistes nimpha
Polistes nympha, female.jpg
Female on nest
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Vespidae
Subfamily: Polistinae
Tribe: Polistini
Genus: Polistes
Species:
P. nimpha
Binomial name
Polistes nimpha
(Christ, 1791)
Synonyms
  • Vespa nimpha(Christ, 1791)
  • Vespa diadema(Latreille, 1802)
  • Polistes opinabilis(Kohl, 1898)
  • Polistes moltonii(Guiglia, 1944)
  • Polistes nimpha irakensis(Gusenleitner, 1962)

Polistes nimpha is a eusocial paper wasp found all over Europe, with particular sightings in Turkey, Finland, Estonia, and Latvia. [1] [2] It is also found in northern Africa, Pakistan, Iran, India (especially in the northern states of Jammu and Kashmir and Himachal Pradesh), Kazakhstan, Mongolia, and China. [3] The climate in these areas is relatively cold and snowy in the winter, while summers are usually hot and dry, with steppe vegetation. [4] Polistes nimpha colonies are relatively small and easily manipulated. [5]

Contents

Taxonomy and phylogeny

The genus Polistes is known for its morphological and behavioral uniformity. Richards (1973) was the first to propose a global classification of Polistes. [5] The Polistinae have a large tropical distribution and are the most diverse subfamily of the Vespidae. The genus Polistes is extensive and inhabits North America, all the way to Eurasia. Polistes members exhibit a wide range of varying black and yellow color patterns. Polistes nimpha is generally more black-patterned than its relative, Polistes dominula . One may distinguish between closely related Polistes species by comparing the color traits. [2] Wasps of the genus Polistes (Latreille, 1802) are good examples for studying alternative phenotypes in social insects. Their small colony size, accessible nests, and moderate aggressiveness enable them to be studied comprehensively. [6]

Polistes nimpha Vespidae - Polistes nimpha.jpg
Polistes nimpha

Description and identification

The Polistes nimpha is usually more black-patterned compared to Polistes dominula , but it is very difficult to morphologically differentiate between the females of Polistes nimpha and dominula. In females, the color of the malar area (between the mandible and compound eye) is yellow and the 6th gastral sternum is black. In males, the clypeus (the broad plate at the front of the head) has lateral ridges and the antennal segments are dark at the tips. [1] The venom gland of Polistes nimpha has a muscular poison sac in the shape of an oval. The tip of the stinger is curled at the peak towards the middle, and the palps are shorter than the stinger. The terminal palps are covered with substantial feathers. [7]

Nests appearance and materials

The nests are beige and grey with dark grey lines. Their sizes vary, with measurements ranging from 8.5 cm × 9.6 cm (3.3 in × 3.8 in) to 3.8 cm × 5 cm (1.5 in × 2.0 in). The central cells are oriented in relation to the ground morphology. [8] Polistes nimpha usually nest in trees and sometimes in cavities. A nest consists of a single resinous pedicel and a comb not covered by envelope. [4] Since there are no envelopes on Polistes nests, the temperature is not internally maintained inside the nest. Thus, outer temperatures must coincide with the species needs for offspring development. [1]

The species uses a mixture of oral secretions and plant fibers, called paper pulp, to build their nests. These chewed plant fibers from weathered wood and other sources constitute the nest's make-up. The fibers are gathered from areas proximate to the nesting site. The oral secretions ensure the durability of the nest during rain and weathering. The duration of chewing ultimately determines the absorbency of the nest paper. The chewing period of pulps may differ amongst individual colonies.

The nests also consist of organic and inorganic materials; nitrogen is used for the production of the oral secretion, while oxygen, carbon, silisium, calcium, aluminum, potassium, and iron are found in fragments of the nest and within its walls. The amount of protein incorporated into the construction of nests may depend on environment conditions. Correspondingly, the amount of oral secretion used for the nest is positively correlated to the nest's exposure to rainfall. [4]

Distribution and habitat

Polistes nimpha prefer low and relatively warm, uncultivated lands. [1] The wasps prefer to nest on plants, under eaves of roofs and buildings, and in closed areas. Colonies with only one female foundress reside on vegetation, while colonies with two or more female foundresses are usually found in covered and sheltered areas. [8]

Life cycle

Colonies may either be haplometrotic or pleiometrotic. Haplometrotic colonies have a single foundress female, while pleiometrotic colonies have two or more foundress females. [8] The fertile foundresses live through the winter and build a nest in the beginning of May. They raise the first generation of workers, which emerge in the first half of June. Over the summer the colony develops and switches from rearing workers to raising sexual individuals; these sexual individuals include males and future foundresses. In August, there is a mass emergence of the male species, and only after this, do the future foundresses appear. The colony begins to disintegrate in late summer and declines throughout autumn. Once the reproductive individuals mate, the males and workers die. By the end of autumn, only the future foundresses are left to survive the winter and begin the cycle again in June. [9] Polistes colonies found in Turkey are found to be univoltine (have one brood of offspring per year). Polistes nimpha colonies are small, averaging less than 100 workers per colony. [4]

Behavior

Division of labor and resources

The specialization of workers occurs due to age-related polyethism; workers take on specific functions depending on their age within the colony. The Polistes nimpha also have specialized foragers: those who collect and those who utilize. Foragers bring food (prey and nectar) and building materials (wood pulp and water) into the nest and give them to the workers. Individual foragers may deliver one type or both load options throughout its active period. However, if a forager switches to a different kind of load, it usually belongs to the same functional group (food or net materials). There are three functional groups that divide the workers. This division is based on their foraging and handling specializations. The first group is builders: workers who prefer hunting to building and who tend to pass proteinaceous food to other individuals. The second group is prey foragers: workers who deliver building materials but do not pass prey to other workers. The third group includes the non-foraging workers, who engage solely in activities within the nest. Active builders are involved in establishing and maintaining the worker's dominance structure. While food was shared amongst individual foragers, building material and water were used only by the individual who had initially brought them. [10]

Nesting and hierarchy

The behavior and process of nest building consists of foragers delivering the paper material to the construction place; the foundress moving over the area and sometimes exhibiting wagging and/or trapping movements with its abdomen; the foundress grasping the entire amount of materials or some part of it; and the construction of the petiole, cell base, or cell wall. The foundresses wag their abdomens while keeping the batch of pulp in their mandibles. They continue this wagging movement until they find a suitable place to apply the pulp. The foundress may sometimes take the nest material (partly or completely) from the foragers who brought it to the nest. This particular behavior is more common amongst foundresses who occupy larger nests. Foundresses may exhibit dominant behavior towards workers whenever workers attempt to lay eggs. If this happens, the worker's wing tips can become damaged from the foundress's aggressive attacks. [10]

Foundresses and kin recognition

Females often retreat under eaves after hibernation and before the foundation of new nests. Nests are likely to be destroyed during the winter; therefore foundresses sometimes find refuge in parental nests. This retreat is more likely to nests that are sheltered, rather than nests built on vegetation or perches. Since foundresses are siblings, their return to maternal nests enables them to come together before founding new colonies. This philopatric occurrence guides how foundresses establish their colonies in the spring and also influences with whom they build their nests near. The relative reproductive potential of a female may play a role in whether their colony is pleiometrotic or haplometrotic. [8] If a foundress's first nest is destroyed, she will either rebuild it or join an alien colony. More than 15% of foundresses mate twice, instead of just mating once. Foundresses differ from other female generations in phenotype; and this is a result of population adaptation to their environment. Since efficient foundresses are favored for emergence, they contribute to the species overall adaptation. A female's surrounding environment ultimately influences their phenotypic variation. Some examples of these effects include: long absences of rain, cold winters, predators and parasitoids. Phenotypic variation of a new generation of foundresses can also be caused by genetic shifts. For example, population density and larval diet can impact the wasps’ individual success. The more irregular and unpredictable a given habitat is, the more phenotypic changes occur in future foundresses. Thus, it can be inferred that Polistes nimpha are quite sensitive to climate changes. [6]

Living in groups

Two types of foundresses can be identified for their difference in founding behavior: generalists and specialists. Generalists are females that find nesting places first. They can attract other foundresses, and they are vulnerable to attracting predators. These pioneering colonies are the first to be attacked by entomophages, such as the parasitic wasp L. argiolus, the hornet Vespa crabro , and ants. Females who nest later and establish colonies separately or on the edge of large aggregations are not attacked by entomophages as often and are usually invaded at later dates. Specialists on the other hand, are females that prefer to nest proximate to already founded nests. If specialists join the colonies of foundresses from the first group, the specialists take on subordinate positions. [11]

Research on Female identification
In their research article on phenotypic variability of foundresses, Rusina and Orlova suggest the two groups of Polistes nimpha females are phenotypically distinguishable. Generalists are identified with non-infested individuals reared under favorable conditions, while specialists are identified with infested females, reared under conditions of insufficient nutrition. Infested individuals are inferior in size to non-infested females, and they exhibit a lighter pigment on the mesoscutum. There also seems to be a correlation between the worker population and the foundresses’ pigmentation: the more workers there are in a colony, the darker the foundresses mesoscutum is. [11]

Territorial behavior

In the late summer, males patrol and defend small areas of territory, usually around hedges and bushes. A male generally lands with his head directed towards the upper edge of the leaf. Then he rotates in a circle while dragging the ventral part of his abdomen, ending with his head facing in the opposite direction. Rotational direction is the same for a given wasp, and he usually cleans his posterior legs and abdomen afterwards. This movement may enable sexual pheromones to be deposited over territorial perches. The exocrine glands, found in the sternal gastral hypodermis of males, are the site for biosynthesis and storage of various excretions. These secretory glands are involved in the production of a wide range of substances, such as repellent, venom in females, and potentially sexual pheromones in males. [12] Resident males attack anything that flies near their territories, including other males, flies, other insects, and even paper models attached to the tips of grass stalks. Territorial males assault and bite the head and legs of intruders, sometimes leading to permanent impairment of the antennae and legs. Males can take on aggressive postures before attacking; they exhibit open mandibles and raised wings and antennae. Butting wasps typically hover in the air, and fights between neighboring territories are quite common. Territories of male Polistes nimpha seem to be purely symbolic and are comparable to leks of vertebrates, as well as the pheromone-marked sites of many bees. [13]

Mating system

Mate-searching is concentrated to a limited portion of the Polistes nimpha habitat. Males gather in great numbers; either flying high above the ground where matings can take place or perched near hibernation places waiting for uninseminated females. The criteria for these locations depend on accessibility to the nests, appropriate hibernacula, foraging areas, hilltops, landmarks and other environmental necessities. Males land under specific conditions of temperature and insulation; if clouds cover the sun, the wasps may leave the hedges and return when the sun is back out. [13] It is common for male individuals to rub their gastral sterna, mandible, and/or legs on various perches. This behavior seems to be associated with a pheromonal release and has been interpreted to function as scent-marking. Throughout September and October, males congregate in low enclosure walls, fences, or shrub and tree hedges in order to defend and scent-mark selected perches. They leave their mark by rubbing their gastral sterna in a circular motion across the surface of branches. Individual territorial boundaries inevitably overlap and differ depending on the density and aggressiveness of males. Contrary to Polistes dominula behavior, Polistes nimpha exhibit no evidence for alternative strategies or conspicuous size differences among individuals. Polistes nimpha's mate-finding strategy incorporates a complex of the two sites: resting and scent-marking. In order to navigate within this network, the species requires well-developed spatial memory. Landmarks and perches can be considered “hotspots,” since individuals are more likely to encounter a receptive mate at those locations. [14]

Parasitoidism

Latibulus argiolus (Rossi) (Hymenoptera, Ichneumonidae) and Elasmus schmitti (Ruschka) (Hymenoptera, Eulophidae) are parasitoids that are harmful to Polistes nimpha. An infestation of Latibulus argiolus is determined by the presence of oval slanted, light yellow or light orange remains in the larval cuticle on the cell margins. An infestation with Elasmus schmitti consists of dark gray covers in the cells, with the parasitoid meconium prior to pupation. Both parasitoids develop in two generations; the females of the first generation infest the host nest towards the end of May to the middle of June. The second generation infests the host nest from the middle of July to the beginning of August. Both parasitoids are more present in colony clusters and usually infest larger nests. If there is a large amount of parasitoids, the host colony will become infested earlier. If infestations occur before worker emergence, the host population density can be greatly impacted. Therefore, spatial distributions of the nests, as well as seasonal aspects, affect the colony's development and vulnerability to the parasitoids. Polistes nimpha may coexist with the parasitoids, but this stability depends on annual changes in their life cycles, as well as on the various features of the host habitat, since this can directly affect the intensity of predation. The influence of parasitoids on the overall population of the species depends not only on Polistes nimpha’s defensive behavior, but also on the intra- and interspecific competition of the parasitoids. [15]

Predators

The Polistes nimpha is prey to not only parasitoids, but also to entomophages, such as the parasitic wasp L. argiolus, the hornet Vespa crabro, and ants. [11] Pleiometrotic colonies are better protected from invertebrate predators, and they can usually rebuild their nests if it is destroyed. The most vulnerable time in the wasp's life cycle is the interval between nest foundation and worker emergence. [6]

Related Research Articles

<i>Polistes</i>

Wasps of the cosmopolitan genus Polistes are the most familiar of the polistine wasps, and are the most common type of paper wasp in North America. Walter Ebeling coined the vernacular name "umbrella wasps" for this genus in 1975 to distinguish it from other types of paper wasp, in reference to the form of their nests. It is also the single largest genus within the family Vespidae, with over 300 recognized species and subspecies. 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. The European paper wasp, Polistes dominula, was introduced into the US about 1981 and has quickly spread throughout most of the country, in most cases replacing native species within a few years. This species is very commonly mistaken for a yellow jacket, as it is black, strongly marked with yellow, and quite different from the native North American species of Polistes. The cuckoo wasp, Polistes sulcifer, is an obligate social parasite, whose only host is P. dominula. Polistes annularis, whose species name is Latin for "ringed", is also known for its distinctive red body color. Polistes metricus adults malaxate their insect prey by chewing them into a pulp, sucking out and ingesting the body fluids, then feeding the rest of the morsel to their larvae. The most widely distributed South American wasp species, Polistes versicolor, 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. Polistes wasps can be identified by their characteristic flight; their long legs dangle below their bodies, which are also more slender than a yellow jacket.

European paper wasp 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 that of most Polistes species—many genera of insects versus mainly caterpillars in other Polistes—giving it superior survival ability compared to other wasp species during a shortage of resources.

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

Polistes gallicus is a fairly common 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>

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 employees 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 throughout southern Canada, the United States, Mexico, and Central America. 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 queen along with other cofoundresses and a dominance hierarchy.

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

Belonogaster juncea is a typical quasisocial paper wasp from sub-saharan Africa and south-western Asia. It is the type species for the genus Belonogaster.

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

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.

<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. They are native to the United States from Texas to Florida, north to New York, and west to Nebraska. They have also been found in Bermuda and Canada, where they are considered non-native. The wasp's common name is due to the reddish-brown color of its head and body. P. carolina are known to construct some of the largest nests of any wasp species and prefer to build their nests in protected spaces.

<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.

Polistes atrimandibularis is one of three obligate social parasites among the Polistes wasps found in Europe. Of the three social paper wasp parasites, 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>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 life cycle 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 utilize an odor based recognition system that is the basis for all wasp to wasp interaction of the species. The wasp's life cycle is highly intertwined with that of Polistes atrimandibularis, an obligate social parasite wasp that frequently invades the combs of P. biglumis wasps.

Polistes sulcifer is a species of paper wasp in the genus Polistes that is found in Italy and Croatia. It is one of only three 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. sulcifer 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 is a subtropical social wasp within Polistes, the most common genus of paper wasp. The most widely distributed South American wasp species, P. versicolor 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 substratum 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. The P. versicolor colony cycle broadly ranges from 3 to 10 months, although there appears to be no relationship between the colony's development and the season of the year. While yellow paper wasps do have clear annual colony cycles, many young queens have the opportunity to hibernate during the winter, forming optional winter aggregations. Dominance hierarchies within these aggregations are characterized by physical aggression of the dominant female(s) towards the associated females, who tend to be sisters. Wagging movements are also often used as a form of communication within the colony. The yellow paper wasp is generally predatory, capturing a wide range of insects, although it often feeds on pollen and nectar as well. Therefore, P. versicolor can be useful as a pollinator or as effective pest control.

<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.

References

  1. 1 2 3 4 Pekkarinen, A. (1999). "The Polistes Species in Northern Europe (Hymenoptera : Vespidae)". Entomologica Fennica. 10 (4): 191–97. doi: 10.33338/ef.84021 .
  2. 1 2 "Polistes Nimpha (Christ, 1791)". The Global Biodiversity Information Facility: GBIF Backbone Taxonomy, GBIF.ORG.
  3. ↑ Carpenter, James M. (1996). "Distributional Checklist of the Species of the Genus Polistes (Hymenoptera: Vespidae; Polistinae, Polistini)" (PDF). American Museum Novitates.
  4. 1 2 3 4 Bağriaçik, N. "Some Structural Features of Nest Materials of Polistes Nimpha (Christ, 1791) in Several Ecological Conditions (Hymenoptera: Vespidae)". Journal of the Entomological Research Society. 15 (3): 1–7.
  5. 1 2 Carpenter, James M. (1996). "Phylogeny and Biogeography of Polistes". In Turillazzi, S.; West-Eberhard, M. J. (eds.). Natural History and Evolution of Paper-Wasps. Oxford University Press. pp. 18–57. ISBN   0-19-854947-4.
  6. 1 2 3 Rusina, L. Y.; et al. (2011). "Dynamic Stability of Phenotypic Variation in Polistes Wasps (Hymenoptera: Vespidae)" (PDF). Russian Entomological Journal. 20 (3): 321–26. ISSN   0132-8069.
  7. ↑ İşcanoğlu, Ş; Bağriaçik, N. (2011). "Polistes gallicüs (L.), Polistes nimpha (Christ) ve Vespula germanica (Fab.) (Hymenoptera: Vespidae) Türlerinde Zehir Aygıtının Ultramorfolojik Karşılaştırılması". Kafkas Universitesi Veteriner Fakultesi Dergisi (in Turkish). 17 (4): 621–624. doi: 10.9775/kvfd.2010.4380 .
  8. 1 2 3 4 Cervo, R.; Turillazzi, S. (1985). "Associative Foundation and Nesting Sites in Polistes Nimpha". Naturwissenschaften. 72 (1): 48–49. Bibcode:1985NW.....72...48C. doi:10.1007/bf00405334.
  9. ↑ Rusina, L. Yu (2011). "Host Discrimination by Elasmus Schmitti (Hymenoptera, Eulophidae) and Latibulus Argiolus (Hymenoptera, Ichneumonidae), Parasitoids of Colonies of Polistes Wasps (Hymenoptera, Vespidae)". Entomological Review. 91 (9): 1081–087. doi:10.1134/S0013873811090016.
  10. 1 2 Rusina, L. Yu.; Firman, L. A.; Ch; Starr, K. (2011). "Pulp Partitioning and Worker Specialization in Polistine Wasps (Hymenoptera, Vespidae, Polistinae)". Entomological Review. 91 (7): 820–27. doi:10.1134/s0013873811070025.
  11. 1 2 3 Rusina, L. Yu.; Orlova, E. S. (2011). "The Relationship between Phenotypic Variability in Future Foundresses of Polistes Nimpha (Christ) (Hymenoptera, Vespidae, Polistinae) and Infestation of Their Larvae by the Mite Sphexicozela Connivens Mahunka (Acari, Astigmata, Winterschmidtiidae)". Entomological Review. 91 (6): 685–91. doi:10.1134/S0013873811060017.
  12. ↑ Delfino, G.; et al. (1991). "Preliminary Ultrastructural Findings on the Sternal Glands of Male Polistes Nimpha (Christ)". Ethology Ecology & Evolution. 3 (Sup1): 55–58. doi:10.1080/03949370.1991.10721911.
  13. 1 2 Turillazzi, S.; et al. (1982). "Territorial Behaviour in Males of Polistes nimpha (Christ) (Hymenoptera, Vespidae)". Zeitschrift für Tierpsychologie. 58 (2): 174–80. doi:10.1111/j.1439-0310.1982.tb00315.x.
  14. ↑ Beani, L.; et al. (1992). "Landmark-Based Mating Systems in Four Polistes Species (Hymenoptera: Vespidae)". Journal of the Kansas Entomological Society. 65 (3): 211–17. JSTOR   25085358.
  15. ↑ Rusina, L. Yu. (2013). "The Role of Parasitoids in Regulation of Polistes Wasp Population (Hymenoptera, Vespidae: Polistinae)". Entomological Review. 93 (3): 271–280. doi:10.1134/S0013873813030019.
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