Agelaia pallipes

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Agelaia pallipes
Marimbondo carniceiro (Agelaia pallipes).jpg
Adult Agelaia pallipes
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Vespidae
Subfamily: Polistinae
Tribe: Epiponini
Genus: Agelaia
Species:
A. pallipes
Binomial name
Agelaia pallipes
(Olivier, 1792)
Synonyms= [1] [2]
  • Polybia lutea Ducke, 1904
  • Stelolybia pallidipes(Olivier, 1792)
  • Gymnopolybia pallidipes(Olivier, 1792)
  • Vespa pallipesOlivier, 1792

Agelaia pallipes is a species of social paper wasp found from Costa Rica to Brazil, Argentina and Paraguay. A. pallipes is ground-nesting and is one of the most aggressive wasps in South America. This species is a predator of other insects, including flies, moths, and ground crickets, as well as baby birds.

Contents

Taxonomy and phylogeny

A. pallipes is part of the family Vespidae, subfamily Polistinae, tribe Epiponini, and subsequently the genus Agelaia . Vespidae is characterized by the eusociality of its members. Eusociality refers to the complex organization of animal behavior and is defined by these four attributes: adults live in groups, cooperative care of juveniles, reproductive division of labor where only specific members have the capacity to reproduce, and an overlap of generations. [3] In Polistinae, prey and other food resources are masticated and fed to larvae, who in turn give back a clear but nutritious liquid which the adults then consume. The Epopini is a tribe of Polistinae characterized by being polygynic, with many queens and reproduction in swarms. [4]

Relatives of A. pallipes in the Agelaia genus include: A. vicina, A. flavipennis, A. areata, A. angulicollis, A. cajennensis, A. fulvofasciata, A. myrmecophila, A. yepocapa, and A. panamaensis.

Description

Depending on their location and altitude, A. pallipes can vary from 9.05 to 9.21 mm (0.356 to 0.363 in) in length and are yellow with black maculation. Their head displays a range of colors of black, yellow and brown. Their bodily terga vary from yellow to brown and black. Their legs and wings are also yellow. [5] There is clear dimorphism between queens (egg-layers) and sterile female workers with the former displaying physical superiority and size in 17 of 22 characteristics relating to the head, mesosoma, metasoma and wings. [6] There is also clear dimorphism between ovary development of the two different castes of females. Longer and developed ovaries are seen in queens whereas short, thread-like ovaries are seen in workers. There is a stark absence of no transitional females (workers with developed ovaries) thus showing unambiguous division between castes. Furthermore, what separates workers from queens is the presence of a brown or black stripe on the humeri.

Their nests are composed of vertical combs made of horizontal combs forming a single form row of concentric hemispheres. [7] Found in ground cavities, their nests are found to have a maximum diameter of 50 cm (20 in) with a maximum of 16,500 individuals. [8]

Distribution and habitat

Being a species of tropical distribution, A. pallipes is found in the central and northern regions of South America and is commonly restricted to the high-altitude mountain forests and open fields at over 3,000 m (9,800 ft) in altitude. [9] [10] The wasp favors regions of more temperate climate as their preferred regions showed a range of temperatures from 4 °C-12 °C. A. pallipes builds nests in these forests and open fields; however, it is observed to be a ground nesting species with frequent colony displacement. [11] These wasps will not construct envelopes for their nests and, instead, will prefer to obscure themselves in the cavities of tree trunks. [12] In addition to tree trunks, A. pallipes will construct their nests in armadillo tunnels, hollow logs, and man-made constructions. In temperate climates, A. pallipes is found to be among the most abundant species along with its close relative, A. vicina . [13]

Colony cycle

An important characteristic of A. pallipes is its swarm-founding behavior. In swarm-founding species, a large group of workers and one or more queens initiate a new colony and subsequently display polygynic behavior. In contrast to independent-founding species that engage in continuous construction of nests, swarm-founding A. pallipes will engage in periodic construction where the nest is rapidly built to sustain a specific population size. After which, the nest size stays static until capacity is reached and subsequent expansion is necessary where thousands of cells can be built within days. [14]

The colony cycle is the developmental period between one reproductive episode and the next. For swarm-founding species like A. pallipes, new colony reproduction is not necessarily coupled with queen production. [15] Due to their region of preference, the neotropics, the onset of nest foundation varies over a number of months and even seasons exclusive of only the deepest of winter periods. Therefore, unlike its relatives in temperate regions, nest foundation is not limited to the beginning of the favorable season. [16]

Behavior

As seen in the dimorphism between queens and workers, castes can be determined based on both behavior and morphology. As part of the Epiponini tribe, A. pallipes is characterized by being neotropical, polygynous and in relative abundance. The queens’ primary job is to reproduce whereas the workers are charged with foraging for food, caring for the larvae and defending the nest. [17] As necessary with this type of hierarchy, there must be a mechanism for distinguishing between females’ ability to reproduce in a social setting. This reproductive control or suppression has been recently credited to pre-imaginal caste determination via the mechanism of insemination. [18] The lack of intermediate females (workers with developed ovaries) implies heavy control of reproduction. [19] It is hypothesized that only with the advent of insemination, the female will then be able to develop their ovaries adequately to become a layer. [20] Observation has denoted that interactions among queen A. pallipes are unaggressive therefore implying that workers are most important in queen selection. [21]

As a swarm-founding species, it has been observed that in order to communicate at a macro level, individuals will leave a leave a pheromone trail for other members of the nest to follow. Rubbing their gastral sternites on objects when moving in the swarm leaves this trail. [22]

Polyethism

There is clear polyethism among A. pallipes where the young workers care for the larvae and older workers forage and defend the nest. [23] It is hypothesized that the young workers are delegated to stay in the nest and care for larvae because it is significantly safer within the confines of the nest while the older workers are outside being exposed to predation in order to increase life expectancy of the workers. Other activities performed by the older workers include removing larvae from the nest, rubbing gaster on the nest and destroying cells. Intra-cell activities like checking cells, moving gaster and self-grooming were all-indicative of younger workers. [24] Therefore, there is a clear relation between age and function thus implying that A. pallipes displays temporal polyethism behavior.

Morphology

There is clear dimorphism between queen and worker A. pallipes with the former caste observed to be physically superior in 22 characters. [25] Pre-imaginal determination has been referenced to be the reason for differentiation between them. A stark difference between the two castes of females can be seen through examining their ovaries – type A ovaries display filament-appearance with no developed oocytes whereas type B ovaries were longer and well developed and coupled with several mature oocytes. [26] Type B ovaries also contained sperm and could very clearly be categorized as those of queens where as type A ovaries were observed as uninseminated thus identifying as those of workers. This dimorphism displays strong control of sexual organs upon pre-imaginal determination.

A. pallipes morphology is also affected by the altitude with an inverse relationship between higher altitude and reduction in body size, especially hind femur length and head width. [27] The cloud forest environment, where some of these A. pallipes are found, contribute negatively to foraging activity and force individuals to complete life cycles with a smaller body size. [28] This study along the altitudinal gradient provides insight on A. pallipes capacity to adapt morphologically to more severe conditions.

Kin and mate selection

Pre-imaginal determination is the predominant theory as to why certain females become reproductively capable over others. There is a lack of queen-queen aggression that suggests that only workers are involved in the queen selection process. [29] The mechanism for workers choosing their queens is still unclear – however, the pervading theory is related to body size and that the largest females are chosen due to higher survivability. [30] Research has also suggested that the workers have a mechanism for identifying those females who are more fecund and allowing them to identify, and discard, those with less favorable ovaries. Naturally, in a polygynic society of swarm-founding individuals with so many queens, kin selection becomes a challenge. With so many queens, it is expected to have a lower relatedness between workers and the larvae for which they care. [31] Though it is expected that workers will strive to reproduce those who are most like themselves in order to pass on their genes, eusociality still persists. [32]

Feeding and foraging

Agelaia pallipes will normally feed on other insects (flies, moths, butterflies), however, it has been recently discovered that A. pallipes will also feed on carrion. Though the collection of bodily fluids is preferred, the wasp will also settle to take carrion morsels to their larvae; furthermore, it has been documented in recent studies that they will chew tissues of cavities in order to find liquids. [33] A. pallipes was observed to feed on carrion primarily during the fresh stage (98% of occurrences) which can lead to significant forensic importance when determining the post-mortem interval. [34]

A. pallipes has been observed to display social facilitation for recruitment to food resources. This type of behavior is seen in other carrion eaters like crows and ravens where the pooling of independent search efforts is the most efficient way to locate food sources and subsequently defend them from other organisms. This behavior can very quickly develop into an Evolutionary Stable Strategy for the species. [35]

Competition

A. pallipes have been observed to directly compete with several species of ants when contending for food resources. The A. pallipes workers, when approaching an occupied food source, will fly low and flap their wings in order to free areas. The workers will recruit other members of their species from the nest as a form of social facilitation and thus display increased dominance at resources by displaying their competitive behavior. [36] Therefore, like social facilitation for feeding and foraging, the A. pallipes will attempt to recruit its nest mates in order to gain advantages over competitors.

A. pallipes will compete with other species of wasp and will, due to its aggressiveness, generally be successful as seen in their superior abundance on carcasses in the primary stages of decay. [37] However, even though A. pallipes is usually victorious in its interactions, it has been observed that A. pallipes will often be out-competed by certain species of stingless bees.

Communication

By virtue of such large numbers, swarm-founding members must necessarily have a method of communication that will allow the entire nest to coordinate movements efficiently. In a swarm-founding species like A. pallipes, the main conduit of communication is through the use of pheromones. [38] Most individuals of the tribe Epiponini have a Richard's gland to release the pheromones, which is an exocrine gland located on the underside the abdomen. However, it is absent in Agelaia and instead, it is hypothesized that the van der Vecht's gland is the alternative source for the communicative pheromone. By rubbing their gastral on objects, they leave a pheromone trail for their worker counterparts to follow suit. As noticed in A. pallipes foraging habits, the eusocial wasps also receive visual stimulus that communicates to the individual to join the larger group. [39] It was observed through bait stations that those stations with wasp models were better dominated by wasps when ant competition was also present. Furthermore, scientists have proved that even without their olfactory senses, the mere presence of wasp models was enough for recruitment. [40] Thus it is clear that though A. pallipes communicate through the release of trail pheromones, they can also be recruited through simple visual stimuli.

Human importance and current research

Social wasps use venom to defend the nest from intruders and predators and have, in many instances, caused severe deleterious effects in major organs in the human body and other organisms. Due to the aggressiveness of A. pallipes and the number of instances between humans and wasps, the characterization of its venom has become an important issue for scientists.

Recent studies have characterized two novel peptides in the venom of the social wasp: Protonectin and Agelaia-MP. The first venom, Protonectin, is found to be the same venom from relative neotropical social wasp, Protonectarina sylveirae. [41] In cohesion, the two peptide toxins were tested for and displayed potent ability to negatively effect hemolytic activity, mass cell granulation, and chemotaxis in large vertebrates. [42] The inhibition of these activities can result in anaphylactic shock, renal failure and myonecrosis in humans. Certainly, the characterization and identification of these novel A. pallipes toxins will allow scientists to elucidate the mechanism of envenoming which will lead to future pharmacological breakthroughs. [43]

Related Research Articles

<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 more familiar 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>Agelaia</i> Genus of wasps

Agelaia is a genus of Neotropical social wasps, with species from Mexico to northern Argentina. Fifteen of the 31 described species are found in Brazil. These species are swarm founders that nest in cavities. The nest generally is without an envelope.

<i>Apoica pallens</i> Species of wasp

The Central American paper wasp is a nocturnal eusocial wasp. It is famous for its swarm based emigration behavior, and is native to the lowlands of Central and northern South America. This species has developed special night vision adaptations to facilitate their night-time swarming and foraging behavior and has important medicinal properties for the Pankararú people of Brazil.

Apoica flavissima is a paper wasp found primarily in South America. The species is distinguishable by its light coloring, unique single comb nests, and nocturnal nature. A notable feature of this species is the size dimorphism between queens and workers. Unlike most Vespidae wasps, Apocia flavissima queens are smaller than their worker counterparts which results in unique intraspecies relationships.

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>Polybia occidentalis</i> Species of wasp

Polybia occidentalis, commonly known as camoati, is a swarm-founding advanced eusocial wasp. Swarm-founding means that a swarm of these wasps find a nesting site and build the nest together. This species can be found in Central and South America. P. occidentalis preys on nectar, insects, and carbohydrate sources, while birds and ants prey on and parasitize them. P. occidentalis workers bite each other to communicate the time to start working.

<i>Agelaia vicina</i> Species of wasp

Agelaia vicina is a species of wasp in the genus Agelaia. They are neotropical social wasps known to have the largest colony sizes and nest sizes among social wasps, with some colonies exceeding over one million individuals. They are predators of land arthropods, consuming both insects and spiders alike. Recent sperm morphology studies have shown that although Vespidae belong to the superfamily Vespoidea, A. vicina may be more phylogenetically related to Apoidea.

<i>Synoeca cyanea</i> Species of wasp

Synoeca cyanea, commonly known as the marimbondo-tatu in Brazil, is a swarm-founding eusocial wasp. Native to Brazil and Argentina, S. cyanea is one of the largest and most aggressive species of social wasps and is feared in many rural areas. It begins its colony cycle in the early spring and continues until nest abandonment. Throughout its life, S. cyanea forage sugary substances and animal carcasses for food and wood pulp for its nest. S. cyanea is also known for its strong venom, which is enough to cause haemolytic activity.

<i>Protonectarina</i> Genus of wasps

Protonectarina sylveirae, commonly referred to as the Brazilian wasp, is a neotropical swarm-founding wasp species that ranges widely across South America. This species relies heavily on the consumption of animal protein rather than nectar. P. sylveirae preys heavily on agricultural pests to coffee crops, keeping pest populations low.

<i>Leipomeles dorsata</i> Species of wasp

Leipomeles dorsata is a neotropical paper wasp that is found across Central America and northern South America. It is a eusocial wasp with little differentiation between reproducing and non-reproducing females. In fact, workers can become temporary reproductives if the main reproductives are killed, allowing reproduction to continue until the main reproductive population recovers. The colony cycles through different ratios of main reproductive females and subordinate reproductive females, starting with few or no primary reproducing females, and increasing until there are only main reproductives.

Protopolybia exigua is a species of vespid wasp found in South America and Southern Brazil. These neotropical wasps, of the tribe Epiponini, form large colonies with multiple queens per colony. P. exigua are small wasps that find nourishment from nectar and prey on arthropods. Their nests are disc-shaped and hang from the undersides of leaves and tree branches. This particular species of wasp can be hard to study because they frequently abandon their nests. P. exigua continuously seek refuge from phorid fly attacks and thus often flee infested nests to build new ones. The wasps' most common predators are ants and the parasitoid phorid flies from the Phoridae family.

<i>Polybia sericea</i> Species of wasp

Polybia sericea is a social, tropical wasp of the family Vespidae that can be found in South America. It founds its colonies by swarming migrations, and feeds on nectar and arthropods.

<i>Synoeca surinama</i> Species of wasp

Synoeca surinama is a Neotropical swarm-founding wasp of the tribe Epiponini. It is known for its metallic blue and black appearance and painful sting. S. surinama builds nests on tree trunks and can be found in tropical climates of South America. When preparing to swarm, there are a number of pre-swarming behaviors that members of S. surinama colonies partake in, such as buzzing runs and occasional brood cannibalism. In S. surinama, social environmental conditions determine the caste ranks of individuals in the developing brood. Unlike less primitive Hymenoptera species, S. surinama display little morphological variation between egg laying queens and workers. S. surinama wasps visit flowering plants and are considered pollinators. When these wasps sting, the stinger is left in the victim and the wasp ultimately dies.

Parachartergus fraternus is a neotropical, swarm founding, polistine wasp species that is distributed throughout Central and South America. They live in nests in second growth tropical dry forests, near pasture fields, roadside areas, and urban areas as well. These wasps eat insects, such as caterpillars of Lepidoptera. They also drink nectar, honeydew, and water. The workers capture their prey during foraging. They also use venom to paralyze their prey in order to consume it later. P. fraternus wasps are not very aggressive and they do not attack when the nest is approached.

<i>Synoeca septentrionalis</i> Species of stinging wasp

Synoeca septentrionalis is one of five species of wasps in the genus Synoeca. It is a swarm-founding wasp that is also eusocial, exhibiting complicated nest structure and defense mechanisms and a colony cycle including a pre-emergence phase and a post-emergence phase. It is typically found in areas from Central to South America. This wasp is one of the larger species of paper wasps and exhibits multiple morphological adaptations as a result of this. Synoeca septentrionalis is known for possessing a very painful sting.

<i>Agelaia multipicta</i> Species of wasp

Agelaia multipicta is a swarm-founding, highly eusocial wasp that lives in Mexico, Argentina, Trinidad and southern Brazil. It nests in natural cavities such as hollow trees and aggressively defends the nest from ants, who are brood predators. The workers and queens are morphologically distinguished by ovarian development as well as external features such as a larger petiole and gaster in the queen. Like other carrion-eating (necrophagous) wasp species, A. multipicta plays a scavenging role in the ecosystem. Agelaia multipicta was described by the Irish entomologist Alexander Henry Haliday in 1836.

Brachygastra scutellaris, a honey wasp, is a Neotropical, swarm-founding species that is found in South America and has a medium-sized population of 100-1000 individuals per colony. It stores large amounts of nectar in its nest for the production of honey, and it was even found that at certain times of the year, the nectar is toxic to humans, as they will extract nectar from hallucinogenic plants, depending on the season.

Protopolybia chartergoides, also known as Pseudochartergus chartergoides, is a species of wasp within the genus Protopolybia. It is a social wasp found in southern Central America and northern South America.

<i>Polybia paulista</i> Species of wasp

Polybia paulista is a species of eusocial wasp occurring in Brazil, Paraguay, and Argentina.

<i>Parachartergus</i> Genus of wasps

Parachartergus is a genus of epiponine social wasps belonging to the subfamily Polistinae. Species include:

References

  1. "Agelaia multipicta (Haliday, 1836)". GBIF.org. Retrieved 2 May 2013.
  2. James M. Carpenter. "Tentative Checklist of the Polistine Tribe Epiponini". IUNH. Archived from the original on 29 December 2017. Retrieved 2 May 2017.
  3. Plowes, N. (2010) An Introduction to Eusociality. Nature Education Knowledge 3(10):7
  4. Noll, F, et al. 2004. Evolution of Caste in Neotropical Swarm-Founding Wasps (Hymenoptera: Vespidae; Epiponini). American Museum Novitates.
  5. Hermes, Marcel Gustavo and Kohler, Andreas. The genus Agelaia Lepeletier (Hymenoptera, Vespidae, Polistinae) in Rio Grande do Sul, Brazil. Rev. Bras. entomol.[online]. 2004, vol.48, n.1 [cited 2014-09-25], pp. 135-138
  6. Noll, F.B., D. Simões & R. Zucchi. 1997. Morphological caste differences in Neotropical swarm-founding Polistinae wasps. Agelaia m. multipicta and A. p. pallipes (Hymenoptera: Vespidae). Ethol. Ecol. Evol. 9: 361-372.
  7. Richards O.W. 1978. The social wasps of the Americas excluding the Vespinae. London: British Museum. 580pp.
  8. Rodríguez-Jimenez, Andrea, & Sarmiento, Carlos E.. (2008). Altitudinal distribution and body resource allocation in a High Mountain social wasp (Hymenoptera: Vespidae). Neotropical Entomology, 37(1), 1-7.
  9. Hermes, Marcel Gustavo and Kohler, Andreas. The genus Agelaia Lepeletier (Hymenoptera, Vespidae, Polistinae) in Rio Grande do Sul, Brazil. Rev. Bras. entomol.[online]. 2004, vol.48, n.1 [cited 2014-09-25], pp. 135-138
  10. Rodríguez-Jimenez, Andrea, & Sarmiento, Carlos E.. (2008). Altitudinal distribution and body resource allocation in a High Mountain social wasp (Hymenoptera: Vespidae). Neotropical Entomology, 37(1), 1-7.
  11. Fowler, H.G. 1992. Social Facilitation During Foraging in Agelain (Hymenoptera: Vespidae). Naturwissenschaften. 79, 424.
  12. Hermes, Marcel Gustavo and Kohler, Andreas. The genus Agelaia Lepeletier (Hymenoptera, Vespidae, Polistinae) in Rio Grande do Sul, Brazil. Rev. Bras. entomol.[online]. 2004, vol.48, n.1 [cited 2014-09-25], pp. 135-138
  13. Thiago de Carvalho Moretti et al., 2011. Bait and Habitat Preferences, and Temporal Variability of Social Wasps (Hymenoptera: Vespidae) Attracted to Vertebrate Carrion. Journal of Medical Entomology. 48(5): 1069-1075.
  14. De Oliveira, Otavio, et al. 2010. Foraging Behavior and Colony Cycle of Agelaia vicina (Hymenoptera: Vespidae; Epiponini). Journal of Hymenoptera research.
  15. Hastings, M.D., Strassmann J.E. 1998. Kin selection, relatedness, and worker control of reproduction in a large-colony epiponine wasp, Brachygastra mellifica. Behavioral Ecology. 9(6): 573-581.
  16. F. B. Noll & R. Zucchi (2000) Increasing caste differences related to life cycle progression in some neotropical swarm-founding polygynic polistine wasps (Hymenoptera Vespidae Epiponini), Ethology Ecology & Evolution, 12:1, 43-65
  17. Noll, F.B., Zucchi R. 2002. Castes and the influence of the colony cycle in swarm-founding polistine wasps. Insectes Sociaux. 49: 62-74.
  18. O’Donnell, S. 1998. Reproductive Caste Determination in Eusocial Wasps (Hymenoptera: Vespidae). Annual Review of Entomology. 43: 323-46.
  19. Biao, M., et al. 2003. Shape differences rather than size differences between castes in the Neotropical swarm –founding wasp Metapolybia docilis (Hymenoptera: Veispidae, Epiponini). BMC Behavioral Ecology.
  20. Noll, F.B., Zucchi R. 2002. Castes and the influence of the colony cycle in swarm-founding polistine wasps. Insectes Sociaux. 49: 62-74.
  21. Noll, F.B., Zucchi R. 2002. Castes and the influence of the colony cycle in swarm-founding polistine wasps. Insectes Sociaux. 49: 62-74.
  22. Smith, A., O’Donnell, S. 2002. Evolution of Swarm Communication in Eusocial Wasps (Hymenoptera: Vespidae). Journal of Insect Behavior. 15(6)
  23. Torres, V., et al., Temporal polyethism and life expectancy of workers in the Eusocial wasp Polistes Canadensis Canadensis Linnaeus (Hymenoptera: Vespidae). Sociobiology 60(1):107-113.
  24. Torres, V., et al., Temporal polyethism and life expectancy of workers in the Eusocial wasp Polistes Canadensis Canadensis Linnaeus (Hymenoptera: Vespidae). Sociobiology 60(1):107-113.
  25. Noll, F.B., D. Simões & R. Zucchi. 1997. Morphological caste differences in Neotropical swarm-founding Polistinae wasps. Agelaia m. multipicta and A. p. pallipes (Hymenoptera: Vespidae). Ethol. Ecol. Evol. 9: 361-372.
  26. Biao, M., et al. 2003. Shape differences rather than size differences between castes in the Neotropical swarm –founding wasp Metapolybia docilis (Hymenoptera: Veispidae, Epiponini). BMC Behavioral Ecology.
  27. Rodríguez-Jimenez, Andrea, & Sarmiento, Carlos E.. (2008). Altitudinal distribution and body resource allocation in a High Mountain social wasp (Hymenoptera: Vespidae). Neotropical Entomology, 37(1), 1-7.
  28. Rodríguez-Jimenez, Andrea, & Sarmiento, Carlos E.. (2008). Altitudinal distribution and body resource allocation in a High Mountain social wasp (Hymenoptera: Vespidae). Neotropical Entomology, 37(1), 1-7.
  29. Noll, F.B., Zucchi R. 2002. Castes and the influence of the colony cycle in swarm-founding polistine wasps. Insectes Sociaux. 49: 62-74.
  30. Noll, F.B., Zucchi R. 2002. Castes and the influence of the colony cycle in swarm-founding polistine wasps. Insectes Sociaux. 49: 62-74.
  31. Strassman, J., et al. 1991. Relatedness and queen number in the Neotropical wasp, Parachartergus colobopterus. Animal Behavior. 42: 461-470.
  32. Strassman, J., et al. 1991. Relatedness and queen number in the Neotropical wasp, Parachartergus colobopterus. Animal Behavior. 42: 461-470.
  33. Gomes, L. et al. 2007. Occurrence of Hymenoptera on Sus scrofa carcasses during summer and winter seasons in southeastern Brazil. Revista Brasileira de Entomologia. 51(3)
  34. Gomes, L. et al. 2007. Occurrence of Hymenoptera on Sus scrofa carcasses during summer and winter seasons in southeastern Brazil. Revista Brasileira de Entomologia. 51(3)
  35. Davies, Nicholas., Krebs, John., West, Stuart. 2012. An Introduction to Behavioral Ecology. Oxford: Wiley-Blackwell. p.160.
  36. Fowler, H.G. 1992. Social Facilitation During Foraging in Agelain (Hymenoptera: Vespidae). Naturwissenschaften. 79, 424.
  37. Gomes, L. et al. 2007. Occurrence of Hymenoptera on Sus scrofa carcasses during summer and winter seasons in southeastern Brazil. Revista Brasileira de Entomologia. 51(3)
  38. Smith, A., O’Donnell, S. 2002. Evolution of Swarm Communication in Eusocial Wasps (Hymenoptera: Vespidae). Journal of Insect Behavior. 15(6)
  39. Fowler, H.G. 1992. Social Facilitation During Foraging in Agelain (Hymenoptera: Vespidae). Naturwissenschaften. 79, 424.
  40. Fowler, H.G. 1992. Social Facilitation During Foraging in Agelain (Hymenoptera: Vespidae). Naturwissenschaften. 79, 424.
  41. K Dohtsu, K Okumura, K Hagiwara, M.S Palma, T Nakajima. 1993. Isolation and sequence analysis of peptides from the venom of Protonectarina sylveirae (Hymenoptera-Vespidae). Nat. Toxins, 1 (1993), pp. 272–276
  42. Mendes, Maria. 2004. Structural and biological characterization of two novel peptides from the venom of the neotropical social wasp Agelaia pallipes pallipes. Toxicon. 44(1): 67-74.
  43. Baptista-Saidemberg, Nicoli, et al. 2011. Profiling the peptidome of the venom from the social wasp Agelaia pallipes pallipes. Journal of Proteomics. 74(10): 2123-2137