Polistes pacificus

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Polistes pacificus
Polistes pacificus (26907465).jpg
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. pacificus
Binomial name
Polistes pacificus
(Fabricius, 1804)
Synonyms [1]
  • Polistes flavopictusDucke, 1918
  • Polistes liliaceusculusSaussure, 1855
  • Polistes liliaciosusSaussure, 1855
  • Polistes modestusSmith, 1862
  • Polistes trinitatisBequard, 1937

Polistes pacificus is a Neotropical species of social paper wasp belonging to the subfamily Polistinae and the family Vespidae. [2] P. pacificus can be found distributed throughout most of Central and South America and parts of southern North America. First discovered by Johan Christian Fabricius in 1804, P. pacificus is much darker in color than some other more recognizable Polistes wasps, and is one of the insects commonly eaten by several indigenous groups in Venezuela and Colombia. [3] [4]

Contents

Taxonomy and phylogenetics

Phylogenetic tree

P. pacificus is part of a monophyletic clade that makes up the membership of Richards’ (1973, 1978) subgenera Epicnemius. [5] P. pacificus is most closely related to its sister species, P. cinerascens. Although previous studies found Epicnemius to be paraphyletic, the most recent and complete study done by Picket et al. reveals that it is likely monophyletic. The species included in this clade are P. cinerascens, P. pacificus, P. geminatus geminatus, P. occipitalis, P. testaceicolor. [5]

Synonyms

P. pacificus was originally discovered in 1804 by the famous Danish entomologist, Johan Christian Fabricius (1745-1808). However, because P. pacificus has wide color variation depending on its geographic region, over time it has been mistaken for several new species and given new names. These names included P. flavipictus Ducke, 1918 P. liliaceusculus de Saussure, 1854, P. liliaciosus de Saussure, 1855, P. modestus Smith, 1862, P. trinitatis Bequard, 1937. These names are relatively obsolete, but may appear in erroneous or outdated literature. [6]

Description and identification

Distinguishing P. pacificus from similar species

P. pacificus has varied coloration patterns with markings that can be black, ferruginous and yellow. It is darker and less brightly colored than most other Polistes species, and it is morphologically very similar to P. boharti. In fact, the two species are classified as the same under Richard's key. [4] However, there are some distinguishing characteristics between the two: Firstly, the digitus structure of the genitalia of P. pacificus males has many dark, stout, distally curved or hooked setae. Secondly, female P. pacificus will have a genal ridge that is usually weak below, and rarely reaches below the level of the lower one-fifth to one-fourth of the eye; on the rare occasions that it extends below this level, it will most likely not reach the base of the mandible and will be very weak. Additionally, in P. pacificus, the lobate humeral angle is not defined, and the lower lateral extension of the pronotal ridge will not be depressed. [4]

The defined morphological criteria

Wasp morphology Wasp morphology.png
Wasp morphology

A specimen of P. pacificus may also be identified by the following morphological criteria: [5]

Front of insect head diagram Front of insect head diagram.svg
Front of insect head diagram

Nest identification

P. pacificus builds nests that are relatively small and made of paper. These nests are most likely to be found in areas of low second-growth vegetation, [3] and are often sheltered by large leaves. [7] The nests, which hang from a plant supported by one or more pedicels, are made up of a single comb of hexagonal cells and do not have an envelope. [3] Polistes produce the water-resistant paper used to make nests by gathering wood fiber and other plant matter and combining it with saliva. [8]

Distribution and habitat

Geographic distribution

P. pacificus can be found from the southern United States to Uruguay and Argentina. P. pacificus can be found in Southern parts of Texas, various areas of Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, Panama, Colombia, Ecuador, Peru, Venezuela, Trinidad, Guyana, Suriname, French Guiana, parts of Brazil, Paraguay, Bolivia, and Uruguay. [9]

Nest-building habitats

P. pacificus builds its nest on plants, using large leaves of the host plants as shelter. While some reports have been made that suggest that social paper wasps build nests where arboreal ants are also present, P. pacificus is only rarely associated with arboreal ants and frequently chooses nesting sites on plants that are unattractive to ants. [7] Some of the most common plants for P. pacificus to build nests on are Clusia grandiflora and Vismia sessilifolia, but they have also been known to build nests that connect to long, thin thorns under the leaves on juvenile Astrocaryum sciophylum palm trees. [7]

Colony cycle

Lek diagram for mating Lek-diagram.jpg
Lek diagram for mating

P. pacificus is one of several social paper wasp species that migrate seasonally depending on the times of the wet and dry seasons. In areas with less pronounced dry seasons, P. pacificus is more likely to brood continuously. However, when P. pacificus live in regions with a defined dry season, the dry season is the non-nesting phase of the colony cycle. [10] During this period, wasps leave the low elevation habitats in which they normally nest and reproduce during the rainy season, and move to cooler areas at higher elevation. The aggregating wasps are sometimes found to be in small groups of individuals from different nests. [10] Due to the fact that dry season aggregates of P. pacificus have been found in evergreen old growth forests, a place distinct from where they nest, it has been suggested that this behavior is evidence of altitudinal migration. [10] The benefits to this type of seasonal migration could be that wasps in these cooler high elevation conditions have lower metabolic needs and costs. [10] This behavior is similar to migrating behavior observed in temperate zone Polistes wasps during winter. Mating can occur either at leks away from nests or at newly founded nests, often late in the nesting season, but before the seasonal dormancy. Generally the inseminated females will pass the dormant season hibernating in a group of other females in a type of shelter, and at this time males most likely die off. [10]

Interactions with other species

Diet

The Polistes have high nutritional demands because they nurse their broods. [11] P. pacificus is an animal of prey, but it is also a flower-visiting wasp that gets carbohydrates from flower nectar. [12] When P. pacificus is in the larval state, the majority of its diet comes from animal protein. In addition to the caterpillars which Polistes mostly preys on, the P. pacificus also visits flowers, thus picking up carbohydrates. [12] Carbohydrates collected through flower nectar are a complement of its diet and thought to help maintain its own energy metabolism. [13]

Predators

P. pacificus has at least several known predators: arboreal ants and vertebrates, including humans. The larvae of the P. pacificus is what predators eat, as it is localized within the nest and immobile. The larvae makes a very valuable food source because it is rich in amino acids and fats. [3] [14]

Defense

Clusia grandiflora, the tree on which P. pacificus often constructs nests because it is not appealing to predators. Clusia grandiflora 137-8387.jpg
Clusia grandiflora, the tree on which P. pacificus often constructs nests because it is not appealing to predators.

P. pacificus nest site selection and structure have evolved under selective pressures of weather and predation. [14] In order to keep predators out, P. pacificus nests are built in specific places in which they are less susceptible to attack by arboreal ants or vertebrates. [14] The placement of the nest not only aims to reduce predation, but also to provide protection from extreme weather. [14] Nests are most often built on the underside of large leaves, which protect against weather and limit predator’s views of the nest from above. The most preferable large leaves have spines along the central vein, which act as a pedicel and protect the nests from ant attacks. [14] In the neotropics, P. pacificus nests are most closely associated with a specific plant, the Clusia grandiflora, which is most likely an adaptation due to the selective pressure of ant predation as Clusia is not an attractive host for arboreal ant nests. [7]

Communication

Lack of visual signaling

Communication is a critical for any species and can benefit both the sender and receiver of information. [15] Some wasps have high visual acuity and use recognition of facial patterns to establish dominance hierarchies; [16] however, P. pacificus differs from these species in that it does not use visual signals, and must rely on other modes of communication. [17] It is likely that P. polistes detects pheromones and other non-visual recognition cues to distinguish nestmates from nonnestmates and determine dominance. [18]

Worker mouthing and rubbing

Adult female workers of P. pacificus have been observed to mouth and rub newly emerged female wasps, during which time the newly emerged wasps remain stationary. In several cases, the older worker used her gaster to rub the new wasp with discrete anterior to posterior strokes. In other cases, the older female has also been seen to mouth the young wasps with her mandibles, by opening and closing them over the new wasp’s head, thorax and gaster. [18] While in contact with the younger wasp, the older worker kept her sting chamber open. The workers that were observed executing this behavior had few, small, only partially developed eggs in their ovaries, and had shriveled Dufour’s glands. [18] Although it is unclear why exactly the wasps partook in this behavior, it is possible that doing so may give each of the newly emerged wasps the colony specific odor, which could ensure them safety when returning to the nest after foraging. [18] It has also been hypothesized that this behavior may express some order of dominance. [19]

Human importance

Agriculture

P. pacificus, which gets some of its nutrition from flower nectar, is also a pollinator, and thus an important species for agriculture. [12] In Mexico, P. pacificus has been observed to pollinate avocados, and it has been suggested that as a native species to the area P. pacificus started a coevolution with avocado species. [12] Although P. pacificus is not as abundant as some of the other insects known to pollinate avocados, it nevertheless contributes to avocado pollination and production. [12]

Human food source

Not only does P. pacificus help pollinate avocados, thus helping to provide a calorie-dense plant-based food source for humans, but it also is commonly hunted by humans who eat the wasp larvae. [3] Although insect eating is often portrayed as an archaic trait, among the Yukpa-Yuko Indians of Venezuela and Colombia, insect food sources are still vitally important, as they supplement the Yukpa’s diet by providing animal proteins and fats. [3] The Yukpa word for P. pacificus is “nonawu,” and the Indians can recognize the nests as being low to the ground in second-growth vegetation. [3] Although both women and men may participate in identifying nests, collecting the nests is exclusively men’s work. [3] When it comes time to harvest, the men wait until the sun is setting, then build a fire on the ground below where the nest is hanging. Using a hooked stick to detach the nest from its connection site, the nest is detached from the plant and immediately placed in the fire. Once in the fire, the adult insects evacuate as quickly as possible while the larvae die as they are toasted. The combs with the toasted larvae inside are then removed from the fire and brought home. [3] The larvae can be considered celebratory food, and when a boy is newly born it is the father’s duty to collect a nest alone. [3]

Cultural significance of wasp stings

Several tribes from the Amazon basin collected P. pacificus, along with other Polistes wasps, to use their stings as instruments of pain during cultural rights of passage. [20] For example, the Oyana people in Surinam constructed breastplates that were made of woven plant stems, in which live wasps were trapped. During initiation ceremonies, young boys would have to wear the breast plate all night, enduring the pain of continual wasp stings. This would have been especially painful because wasps are able to retract their stings and repeatedly insert them with new venom multiple times. [20]

Related Research Articles

<span class="mw-page-title-main">Vespidae</span> Family of insects

The Vespidae are a large, diverse, cosmopolitan family of wasps, including nearly all the known eusocial wasps and many solitary wasps. Each social wasp colony includes a queen and a number of female workers with varying degrees of sterility relative to the queen. In temperate social species, colonies usually last only one year, dying at the onset of winter. New queens and males (drones) are produced towards the end of the summer, and after mating, the queens hibernate over winter in cracks or other sheltered locations. The nests of most species are constructed out of mud, but polistines and vespines use plant fibers, chewed to form a sort of paper. Many species are pollen vectors contributing to the pollination of several plants, being potential or even effective pollinators, while others are notable predators of pest insect species, and a few species are invasive pests.

<span class="mw-page-title-main">Paper wasp</span> Vespid wasps that gather fibers from dead wood and plant stems

Paper wasps are a type of vespid wasps. The term is typically used to refer to members of the vespid subfamily Polistinae, though it often colloquially includes members of the subfamilies Vespinae and Stenogastrinae, which also make nests out of paper.

<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">Polistinae</span> Subfamily of insects

The Polistinae is a subfamily of eusocial wasps belonging to the family Vespidae. They are closely related to the wasps and true hornets of the subfamily Vespinae, containing four tribes. With about 1,100 species total, it is the second-most diverse subfamily within the Vespidae, and while most species are tropical or subtropical, they include some of the most frequently encountered large wasps in temperate regions.

<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>Brachygastra</i> Genus of wasps

Honey wasps are species in the genus Brachygastra of the family Vespidae. Brachygastra comprises 17 species of social paper wasps. The ancestral species are thought to have diverged about 32 million years ago within diverse Amazon rainforest. Subsequent speciation within the genus is thought to have mostly occurred between 23 Ma and 10 Ma, during the time of the Andean uplift when the landscape was significantly altered due to tectonic activity. The current cladistic organisation of the genus has been heavily reliant on morphological characteristics.

<span class="mw-page-title-main">Wasp</span> Group of insects

A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. The wasps do not constitute a clade, a complete natural group with a single ancestor, as bees and ants are deeply nested within the wasps, having evolved from wasp ancestors. Wasps that are members of the clade Aculeata can sting their prey.

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

Polistes humilis, known as the Australian paper wasp, is a species of wasp in the family Vespidae that is found throughout Australia and which has been introduced to northern New Zealand. These paper wasps can be identified by their long thin legs and banded yellow and black coloring. They have been known to re-utilize old nests. While the species does not exhibit morphological class differences, there are distinct behavioral differences between queens and workers. In addition, the species is eusocial and benefits from relatedness between individuals. They are known for delivering a painful sting, especially when their nest is disturbed, a behavior that has been developed as a nest defense mechanism. While wasps are often viewed negatively, they play an important pollination role for many plants.

<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 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 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 has been found in Ontario, Canada and the eastern United States from Illinois down south to Florida and west to Nebraska and California. It is also found in Mexico from Chihuahua to Jalisco, Hidalgo. P. exclamans has shown variability in its range including an absence of the species in eastern Missouri from the 1920’s to 1940’s, a presence in the 1960’s to 1980’s, and an apparent absence again of the species in these same sites in eastern Missouri since 1989. This suggests that their range has either expanded northward and contracted southward or that they have large, long-term cycles of abundance. 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 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.

<i>Brachygastra lecheguana</i> Species of wasp

Brachygastra lecheguana, formerly known as Nectarina lecheguana, is a species of dark paper wasp found across North and South America. It nests in underbrush in grassland-type environments, and produces honey, characteristic of the genus Brachygastra.

<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 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 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.

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.

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

Mischocyttarus mexicanus is a New World species of paper wasp that exhibits facultative eusocial behavior and includes at least two subspecies living in the southern United States and Central America. This social wasp species is a good model for studying the selective advantage of different nesting tactics within a single species. M. mexicanus females can form nests both as individuals and as members of a colony, and are even known to switch between these two nesting strategies throughout their life, which is an unusual phenomenon in the field of social biology. Individuals in a colony have particular social roles that are plastic, as opposed to rigid castes, and brood parasitism and usurpation have been observed between unrelated conspecifics. They nest in a variety of types of plants and human constructions, although they have most frequently been observed in palm trees, and they are known to interact with a number of other species as prey, competitors over resources, or foragers.

<i>Mischocyttarus mexicanus cubicola</i> Subspecies of wasp

Mischocyttarus mexicanus cubicola is a neotropical subspecies of paper wasp found in the New World. It is a social wasp that demonstrates two different types of nesting strategies, depending upon context. This context-dependent trait makes Mischocyttarus mexicanus cubicola a good model to study social biology within social wasps. In detail, this trait allows for the females of this species to form nests both individually and as co-founders with other females within the same colony. This subspecies is also known to exhibit cannibalism, with M. m. cubicola queens feeding on their own larvae for nourishment when unaided by workers.

References

  1. ↑ "Polistes pacificus Fabricius, 1804". GBIF.org. Retrieved 1 May 2017.
  2. ↑ Carpenter, James; Bolivar Garcete-Barrett; Alexander Lopez (2012). "Las Vespidae (Hymenoptera: Vespoidea) de Guatemala". Biodiversidad de Guatemala. 2. Universidad del Valle de Guatemala: 269–279.
  3. 1 2 3 4 5 6 7 8 9 10 Ruddle, Kenneth (1973). "The Human Use of Insects: Examples from the Yukpa". Biotropica. 5 (2): 94–101. doi:10.2307/2989658. JSTOR   2989658.
  4. 1 2 3 Snelling, Roy (1983). "Taxonomic and Nomenclatural Studies on American Polistine Wasps (Hymenoptera: Vespidae)". Pan-Pacific Entomologist. 59 (1–4): 267–280.
  5. 1 2 3 Pickett, Kurt; James Carpenter; Ward Wheeler (2006). "Systematics of Polistes (Hymenoptera: Vespidae), with a phylogenetic consideration of Hamilton's haplodiploidy hypothesis". Ann. Zool. Fennici. 43. Finish Zoological and Botanical Publishing Board: 390–406.
  6. ↑ "The Global Biodiversity Information Facility: GBIF Backbone Taxonomy". 2013.
  7. 1 2 3 4 Corbara, Bruno; J. M. Carpenter; R. Cereghino; M. Leponce; Alain Dejean (2009). "Diversity and nest site selection of social wasps along Guianese forest edges: assessing the influence of arboreal ants". Comptes Rendus Biologies. 332 (5): 470–479. doi:10.1016/j.crvi.2009.01.003. PMID   19393979.
  8. ↑ "Paper Wasp". Microsoft Encarta Online Encyclopedia. 2006.
  9. ↑ Carpenter, James (1996). "Distributional Checklist of Species of the Genus Polistes (Hymenoptera: Vespidae; Polistinae, Polistini)". American Museum of Natural History (3188): 39 pp.
  10. 1 2 3 4 5 Mulkey, Stephen (1999). "Dry Season Migration by Costa Rican Lowland Paper Wasps to High Elevation Cold Dormancy Sites". Biotropica. 31 (1): 192–196.
  11. ↑ Hermes, Marcel; Andreas Kohler (2006). "The flower-visiting social wasps (Hymenoptera, Vespidae, Polistinae) in two areas of Rio Grande do Sul State, southern Brazil". Revista Brasileira de Entomologia. 50 (2): 268–274.
  12. 1 2 3 4 5 Castaneda-Vidozola, A.; A. Equihua-Martinez; J. Valdes-Carrasco; A. F. Barrientos-Priego; G. Ish-Am; S. Gazit (1999). "Insectos polinizadores del aguacatero en los Estados de Mexico y Michoacan". Revista Chapingo Serie Horticultura. 5: 129–136.
  13. ↑ Faegri, K; L. Van der Pijl (1979). The principles of pollination ecology. London: Pergamon Press Ltda. pp. 244 pp.
  14. 1 2 3 4 5 Dejean, Alain; B. Corbara; J. M. Carpenter (1998). "Nesting site selection by wasps in the Guianese rain forest". Insectes Sociaux. 45: 33–41.
  15. ↑ Davies, Nicholas; John Krebs; Stuart West (2012). "Chapter 14: Communication and Signals". An Introduction to Behavioral Ecology. Wiley-Blackwell. pp. 394–423.
  16. ↑ Tibbets, E. A.; J. Dale (2004). "A socially enforced signal of quality in a paper wasp". Nature. 432: 218–222.
  17. ↑ Sheehan, MJ; E. A. Tibbets; J. Jinn (2014). "Coevolution of visual signals and eye morphology in Polistes paper wasps". Biology Letters. 10 (4).
  18. 1 2 3 4 Richter, Monica; Holly Downing; Wayne Richter (1987). "Social Wasp Behavior: Worker Mouthing and Rubbing of Teneral Polistes pacificus (Hymenoptera: Vespidae)". Journal of the Kansas Entomological Society. 60 (2): 347–349.
  19. ↑ West-Eberhard, M. J. (1982). "Communication in social wasps: predicted and observed patterns, with a note on the significance of behavioral and ontogenetic flexibility for theories of worker "altruism."". Communication Ches les Insectes Sociaux: 13–36.
  20. 1 2 Crane, Ethel Eva (2013). The World History of Beekeeping and Honey Hunting. Routledge. 720 pp.
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