Paper wasps umbrella wasps

These wasps are long-legged and usually brown with fewyellow markings; their middle tibia has two apical spurs. Nests for colonies founded by individual queens (Polistes, Mischocyttarus) are made ofa single comb without an envelope, and attached to the substrate by a strong pedicle (giving the appearance of an upside-down umbrella). Queens apply a chemical secretion from a sternal gland on the abdomen to the pedicel and other nest surfaces. This chemical repels potential predators, including ants, parasitic flies, and yellowjackets. Nests for colonies founded by several queens have an envelope that extends over several combs. Adults partially feed on nectar and honeydew.

Females hunt caterpillars, and often masticate the prey at the capture site. They carry a solid prey pellet back to the nest to feed other adults and larvae. In temperate regions, nests are used one season and queens overwinter. A new nest is often built next to the queen's natal nest, and the result may be a large number of new and used nests in one site. Colonies of the honey wasp, Brachygastra mellifica, are perennial. Tropical paper wasp colonies may last only 2-3 months, whereas the nests of others are perennial and may last up to 25 years.

Other wasps often visit the open nests of Polistes spp. In some cases, the visitors are homeless conspecifics whose nests were destroyed by predators, such as birds or people. In others the visitors are there to steal eggs to feed larvae in their own active colony. Before allowing visitors on to their nest, the wasps have to distinguish between orphaned kin, which will be helpers, and unrelated wasps, which are threats. Paper wasps make this distinction using chemical odors. Each wasp assimilates from its nestan odor specific to the insects thatlive there. This smell, which serves as a recognition cue, is locked into the wasp's epicuticle. The source of the compounds that make up the odor is the plant fibers used to construct the nest and secretions from the worker wasps in the colony. Because each colony uses a unique mixture of plants in nest construction, nonthreatening family members are more likely to share this environmentally acquired label.

Polistes is the most widespread of the social wasp genera, and its range encompasses that of nearly all others. There are about 200 species, and they are distributed almost worldwide in temperate and tropical regions. These wasps are often dominant or the only wasp genus present in many regions, and usually very successful. Polistes nests are relatively small and inconspicuous, and they can nest in a variety of above-ground locations. Their abundance and nesting habits often bring them into contact with people, especially in urban environments.

Pest status of paper wasps is based on their aggressive nest-defense behavior and their painful sting, which is life-threatening for some people. These wasps are pests when they nest near or on buildings, and workers respond aggressively toward movement near the nest. The cavity-nesting species, such as Agelaia in tropical regions, often use wall spaces or discarded domestic containers to build their nests. The aggressiveness of species in the South American genus Stelopolybia is variable; at least one species, S. areata, is considered nonaggressive. Species of Synoeca are considered dangerous because of the barbed sting that frequently remains after stinging. In Southeast Asia, colonies of Polybioides have been used in booby traps in warfare. Species of Apoica forage at night, and many are a stinging hazard to persons working in tropical forests at night. Polistes are serious pests in the urban and agricultural ecosystem. In southwestern USA and regions of Mexico, populations ofPolistes exclamans threaten laborers picking citrus fruits. Nests per tree vary from 1 to 26 and the number of wasps per nestis 100-200. P. apachus and P./uscatus have been a problem for agricultural workers.

African paper wasp, Belonogaster junecus Adults are about 35 mmlongand dark brown. Nests consistofabout50 cells, but may sometimes be 100-200 cells. There are no sterile workers in these wasps, and the colony consists of one or more founding females, daughters, and males. Older females are involved in egg-laying and younger females forage. Males remain at the nest but they do not forage and are fed by females. When colonies reach a certain size, small groups offemales leave to form new nests elsewhere. Nests may be built on the outside of buildings. This species occurs inmostparts of southern Africa.

Honey wasps, Brachygastra spp. These are the honey-producing wasps, which are maintained inasemidomesticated state by native people in Mexico. The nests have several horizontal combs with the cells facing downwards. The combs are surrounded by a thin, paper envelope. Colonies may continue for many years with nests composed of several thousand workers.

Petiolate paper wasp, Mischocyttarus flavitarsis Workers are 16-20 mm long, and the body is black, with orange-yellow and reddish brown markings. The petiole is primarily black, and the abdomen (metasoma) is primarily yellow. It occurs in southwestern USA, including California. Species in this genus are distributed from northern Argentina to British Columbia in the west and through the West Indies to Florida in the east. Females overwinter under bark, and they have been found with Polistes fuscatus aurifer and P. dorsalis californica. They have also been found in an abandoned nest ofVespula arenaria.Thenests of M. flavitarsis may be established under the eaves of houses and branches of trees, and are very similar to the nests ofPolistes,but smaller. A related species, M. drewseni,inGuyanaattacks honey bee hives and causes problems with beekeepers. M. phthisicus and M. mexicanus are neotropical, and extend into Texas and Florida; M. flavitarsis has been introduced into Hawaii.

Common paper wasps, umbrella wasps, Polistes spp. Workers are about 18 mm long, brown to blackish brown, and with yellow marking on the abdomen. These wasps occur in all regions of the world except the coldest part of continents. In Europe and North America, colonies of Polistes probably outnumber colonies of all other social wasps combined. In the UK, P. dominulus and P. gallicus occur in the southern parts of the country, and establish nests only during very warm summers. P. gallicus occurs in Europe and was introduced into eastern USA in the 1980s. P. humilis occurs in New Zealand; it was accidentally introduced from Australia. The Pacific Island hornet, P. habraeus, occurs on islands in the southern Pacific region, and sometimes in New Zealand. The hymenopterous parasite, Elasmuspolisti (Elasmidae) is an ectoparasite of Polistes spp. (reported from P. exclamans). Species of Sulcopolistes are social parasites in the nests of Polistes in the UK and continental Europe.

Temperate-region Polistes species overwinter as fertilized queens overwinter. Single queens initiate nests in spring. However, she may be joined by other queens that have not constructed a nest of their own. These individuals establish a dominance hierarchy, with the auxiliary queens subordinate to the founding queen; they all cooperate in the care of the young. Nests with multiple founding queens are more likely to survive than nests with single queens. Multiple-queen colony foundation is common in P. exclamans and is the rule in P. annularis, P. apachus, and P. carolinus. Aggressive interaction frequently occurs between the nest-sharing queens until one attains a dominant status.

Tropical Polistes do not hibernate. Females may leave the nest at any time during the year and found new colonies. The nest is usually constructed by multiple queens. Nest architecture is variable, with some species making a single horizontal comb, while others make a vertical comb, or a series ofcells, one cell below the other. The nest may have a single or multiple attaching pedicels. No Polistes species construct envelopes around the comb. Caterpillars are an important prey of these wasps, but they will capture a variety of other slow-moving insects. Adults feed on honeydew, and they are attracted to bruised fruits.

Large paper wasp, Polistes annularis (Fig. 9.12b) Workers are 20-25 mm long, the body is light brown, and there are coarse transverse ridges on the propodeum. The abdomen is dark brown, while the basal segment is distinctly light brown. This species builds combs in bushes and trees.

Polistes apachus Workers are 16-20 mm long; the body is light brown to reddish brown with yellow markings. It has the habit of building a new nest close to that of the previous

Figure 9.12 Hymenoptera: Vespidae. (a) Polistes fuscatus pallipes; (b) P. annularis; (c) P. exclamans; (d) P. rubiginosus.

year. In one case, this habit was observed for 10 years. New colonies are typically founded by 2-6 queens. Nests are located in agricultural and urban environments. This species occurs in Mexico, Texas, New Mexico, and California.

European paper wasp, Polistes dominulus This species was accidentally introduced into the eastern coast of the USA in the 1970s and has been rapidly spreading westward, displacing P. Juscatus. The native species, P. dominulus, appears to have a number of advantages over P. Juscatus, including earlier production of workers, higher per capita foraging rates by queens and workers, higher queen survivorship, and lack of conspecific pressures. Nest site and prey availability may be limiting factors in the competition between the two species. This species typically has an Old World distribution.

Polistes exclamans (Fig. 9.12c) Workers are about 18 mm long; the body is brown to dark brown, with yellow markings. This species is common in southeastern USA, especially along the Atlantic seaboard. Two or three cooperating queens form nests.

Dark paper wasp, Polistes fuscatus complex Workers are 9-15 mm long, the head and thorax are black with yellow markings; the abdomen is yellow with black bands on segments

1 and 2. Nests are found under the eaves of buildings and sometimes in attics. P. Juscatus aurifer is the most widespread subspecies of P. fuscatus. It ranges from British Columbia to southern California, where it merges with P. Juscatus centralis, and at the eastern part of its range, P. Juscatus aurifer merges with P. Juscatus variatus and P.fuscatus utahensis.

Polistes fuscatus pallipes ( = P. metricus) (Fig. 9.12a) Workers are 17-21 mm long and black and yellow. The abdomen is black; the first segment has a yellow band posteriorly. This species is common in northeastern USA and ranges west to British Columbia.

Orange paperwasp, Polistesrubiginosus(Fig. 9.12d) Workers are 18-24 mm long; the head, thorax, and legs are uniformly orange brown. This species occurs in western North America, and nests in hollow trees or the walls of houses or other buildings.

Asian umbrella wasps, Polybiodes spp. These wasps occur in Southeast Asia and central Africa. They construct nests varying in structure from spirally produced combs to vertical combs surrounded by an envelope. Colonies may have 3000 workers, and nests of species in the genus have been used in booby traps during hostilities in Vietnam.

Asian paper wasps, Ropalidia spp. These wasps occur in Southeast Asia, India, and Australia, and in Africa and Madagascar. There are more than 100 species in Ropalidia. Nests are made of chewed wood fiber, and the type of nest constructed by the various species is diverse. There are simple nests comprised of only a few cells, and large nests with thousands of cells. The nests may be attached to the underside of tree branches, electric wires, and under the eaves of houses. The large nests and peridomestic nesting sites of some species may provide a stinging hazard for humans.

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