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Polistes

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Polistes
Polistes gallicus
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Vespidae
Subfamily: Polistinae
Tribe: Polistini
Genus: Polistes
Latreille,1802
Type species
Polistes gallicus
Synonyms[2]
  • EupolistesDalla Torre,1904
  • SulcopolistesBlüthgen, 1938
  • PolistulaWeyrauch, 1939
  • PseudopolistesWeyrauch, 1939
  • LeptopolistesBlüthgen, 1943
P. metricus,female

Polistesis a cosmopolitan genus ofpaper waspsand the only genus in the tribePolistini.Vernacular namesfor the genus includeumbrella wasps,coined by Walter Ebeling in 1975 to distinguish it from other types of paper wasp, in reference to the form of their nests,[3]andumbrella paper wasps.[4]Polistesis the single largest genus within the familyVespidae,with over 200 recognized species.[5]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.[citation needed]

Description[edit]

As part of subfamily Polistinae,Polisteswasps are covered in short and inconspicuous hair, have aclypeuswith a pointed apex, have a gena that is wide throughout,tergum1 of themetasomais almost straight to gently arched in profile, the tibia of the mid leg has two spurs, and the legs end in simple tarsal claws. The genus can be distinguished from other Polistinae by a sessile metasoma (the first segment at most slightly longer than wide) and the fourth tarsomeres of the mid and hind legs being symmetrical.[6]

Polistesshowsexual dimorphism,with males having seven externally visible metasomal segments whereas females have six. This trait is shared with other vespid wasps.[6]

Polistesspecies have single-layered nests which are shaped like an umbrella, with the cells exposed to the air from the bottom, and no layer wrapping around the nest. The nests are suspended from a surface by apetioleand are constructed from a paper-like substance made of a mix of saliva and wood fibres chewed off old and soft wood or dead twigs. ManyPolistesspecies in general often have nests supported by a longer petiole than those ofVespula.[3]

Biochemistry[edit]

Similar to many insects, Carlson et al 1998 findsPolistescuticular hydrocarbonsto be predominantly many branched, methylbranched alkanes.The reviews of Nelson 1978, Lockey 1988 and Nelson 1993 concur.[7]

Life cycle[edit]

The general life cycle ofPolistescan be divided into four phases:[8]

  1. Founding (or pre-emergence) phase
  2. Worker phase
  3. Reproductive phase
  4. Intermediate phase

Founding (or pre-emergence) phase[edit]

The founding stage begins in the spring when a solitary female (the "foundress" ) (or a small group of related females) initiates the construction of a nest. The wasps begin by fashioning apetiole,a short stalk which will connect the new nest to a substrate (often the eave of a house or outbuilding), and building a single brood cell at the end of it. Further cells are added laterally in a hexagonal pattern, each cell surrounded by six others. Although nests can achieve impressive sizes, they almost always maintain a basic shape: petiolated (stellocyttarous), single-combed, unprotected, and open (gymnodomous).

Eggs are laid by the foundress directly into the brood cells and are guarded by the foundress and the assisting females (if present). After the first larvae hatch, the foundress feeds them via progressive provisioning, bringing softened caterpillar flesh to the larvae multiple times throughout their development (as opposed to the one-time provisioning seen in some other hymenopteran groups). Each of this first seasonal brood of new paper wasps is exclusively female and destined to a subordinate worker position inside the nest; they do not found their own nests and instead assist their mother in the care and maintenance of future sisters.

Some foundress wasps do not build their own nests, but rather attempt to usurp that of another female. These usurpation attempts may or may not be successful, but almost always result in impressive displays of aggression and violence. Females may also adopt a more peaceful alternative reproduction strategy by joining the nest of a close relative (usually a sister) and working as assisting females. In the latter case, such cofounding females are generally, but not exclusively, close relatives.[8]

Worker phase[edit]

The worker phase usually begins in the early summer, roughly two months after colony initiation, with the emergence of the first workers. These new females take up most of the colony's work duties, foraging, caring for brood, and maintaining the structure of the nest. Around this time, those females which assisted in nest foundation (if present) are driven from the nest by aggressive behavior on the part of the foundress, and leave either to start their own late-season nests or usurp another's.[citation needed]

Reproductive phase[edit]

The reproductive phase of the colony begins when the first female reproductives (thegynes) emerge from their brood cells. These reproductives differ from their worker sisters by having increased levels of fat stores andcryoprotectantcarbohydrate compounds (allowing them to survive the overwintering period). These reproductives contribute genes directly to the next generation, while their worker sisters normally pass along their genes indirectly.[citation needed]

Intermediate phase[edit]

Once male reproductives emerge and both males and females disperse from the natal nest for mating flights, the so-called intermediate phase begins. Brood care and foraging behavior decline and worker numbers drop as dying individuals are no longer replaced by new ones. Intracolonial aggression increases and the social cohesion of the nest declines. In temperatePolistesspecies, individuals (almost exclusively inseminated females) gather in groups of up to 50 individuals and seek a sheltered location (called ahibernaculum) in which to overwinter.[citation needed]

Behavior[edit]

Kin selection[edit]

Main article:Kin selection

The reproductive behavior ofPolisteswasps provided some of the first evidence for the mathematical biologistW. D. Hamilton's 1964 theory ofkin selection.Hamilton showed that animals such as workers could be expected to provide assistance to relatives such as their queens according to the costs and benefits involved (K) and their degree of genetic relatedness (r), and gavethe rule that now carries his name,K > 1/r.[9]Early caution existed among researchers as to whether social insects could really assess their relatedness. Hamilton himself suggested an alternative possibility, namely that kin could become associated simply by "population viscosity" —that offspring tend not to disperse far from their birthplaces— and West-Eberhard (1969) found some evidence for this inPolistes.However,Polistesspecies are now known to learn and remember chemical signals (hydrocarbons) picked up from the nest to distinguish nestmates accurately from non-relatives.[10]

Dominance hierarchy system[edit]

Morphologically, the foundress and subordinate reproductive members of the colony differ little. However, behavioral differentiation occurs among females both between and within generations. For example, in the speciesPolistes humilisthe queen displays a "tail-wagging" behavior to assert her dominance over the worker class.[11]Similarly,Polistes canadensisalso possesses behavioral differentiation between the queen and her nestmates, with the queen often suppressing the aggressive behavior of subordinates through lateral abdominal vibrations and stroking. In contrast, unmated females are not aggressive.[12]InPolistes exclamansqueens have different amounts of glucose, fructose, and trehalose which lead to different cryoprotectant levels. This alters their survivability in different temperatures, increasing their odds of reproduction. Females inP. bellicosusare also morphologically similar between caste separations. For example, aP. bellicosusworker could become queen, and egg-layer, if all of the original foundresses die or leave the nest.[13]This is also true forPolistes dorsalis,which also displays dominant behavior. Despite having no distinct morphological caste, roles ofP. dorsalistend to be fixed in a system with division of labor.[14]

Nestmate recognition[edit]

Polistesspp. discriminate colony mates using an acquired (i.e. learned) cue, absorbing hydrocarbons from the natal nest at eclosion.[15]Thiscuticularhydrocarbon "signature" is derived both from the plant material and the foundress-applied substances from which the nest is made. Studies ofPolistes fuscatushave researched the molecular basis of the recognition "pheromone"used by the wasps, and indicate at least some of the recognizable labels have the same chemical constituents as the adult cuticular hydrocarbons. Similar recognition is found inPolistes metricus.[16] Dominant individuals ofP. dominulahave differing cuticular profiles from workers,[17]and the frequent observations of the dominant female stroking itsgasteracross the nest surface, combined with its staying on the nest for longer times than subordinates, suggests the dominant individual may contribute more to the nest odor.[citation needed]

P. carolinafemales do not preferentially feed their own progeny (as larvae).[18]

Further to this recognition of nestmates,Polistes biglumisfoundresses discriminate between 'alien' eggs and their own via differential oophagy.[19]

Gallery[edit]

Species[edit]

203 species were described in the genus by 1996,[5]and new species continue to be described.[20]There are nine species in Europe.[2]

224 species and 126 subspecies are as follows:[21][20][22][23]

Pest status[edit]

Polistes chinensis antennalis,Asian paper wasp has established itself as a pest species in New Zealand.

Along with theGermanandcommon wasps,the Asian and Australian paper wasps (P. chinensisandP. humilis) are considered pests inNew Zealand.Arriving in 1979,[26]the Asian paper wasp has established itself in both the North Island and the northern parts of the South Island. Because it competes with native species (such as thekaka) for insects, nectar, and honeydew,[27][28]it is a hindrance toconservationefforts.[citation needed]

InNorth America,the introduced European speciesPolistes dominulahas rapidly colonized a significant area, and is considered an invasive pest.[29][30]It is a concern for cherry and grape growers inBritish Columbia,as it injures the fruit by biting off the skin. It also spreadsyeastand fungi that harm fruit and can be a nuisance to workers and pickers at harvest.[31]There is evidence it has also displaced native paper wasp species by outcompeting them.[32]

Parasites[edit]

Various other insects areparasitesorparasitoidsofPolistes,including flies (e.g.,Sarcophagidae),mantispids,and wasps in the familiesTorymidae,Mutillidae(rarely),Braconidae,andIchneumonidae(e.g.Latibulus argiolus). Some more specialized groups are more intimately associated withPolistes;this includesstrepsipteransin the familyStylopidae(genusXenos), wasps of the genusElasmus(formerly placed in their own family, "Elasmidae" ), and wasps in the familyTrigonalidae.

The nests of many species of this wasp genus are invaded by theparasitoidcaterpillars of the mothChalcoela iphitaliswhich feed on the wasp larvae andpupasat night, spinning theircocoonsin empty cells.[33][34][35]

Within the subgenusPolistesare four known social obligate parasites:P. atrimandibularis,P. austroccidentalis,P. maroccanus,andP. semenowi,which parasitize otherPolisteswasps.[23]Known host species of these parasites areP. dominulus,P. gallicus,P. nimphus,P. associus,andP. biglumis.[36]Although these parasites differ in their host invasion strategies, their end goal is to successfully infiltrate the host nest and reproduce at the host's expense.[citation needed]

References[edit]

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Further reading[edit]

External links[edit]

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