Polistes canadensis

Red Paper Wasp
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Hymenoptera
Family:	Vespidae
Genus:	Polistes
Species:	P. canadensis

Polistes canadensis, commonly known as the red paper wasp, is a Neotropical, primitively eusocial wasp. A largely predatory species, it hunts for caterpillar meat to supply its colony, often supplementing its developing larvae with nectar.^[1] The most widely distributed American species of the Polistes genus, it colonizes multiple combs, which it rears year-round.^[2] Emerging from hibernation in the spring, the females found nests built out of plant material such as dry grass and dead wood. These nests are not covered with an envelope and feature hexagonal cells in which eggs are laid and larvae develop.^[1] The Polistes canadensis colony divides its colony among several combs and does not reuse these combs as a defense mechanism against parasites such as the tineid moth. A single female queen with, on average, 9.1 foundresses, usually initiates the construction of new combs and cells to form nests. The more foundresses in a colony, the more combs produced. On average, combs grow for 15.4 days and achieve a size of 30.8 cells.^[3] One female queen exercises absolute dominance over all other females, often using lateral abdominal vibrations and stroking to suppress the aggressive behavior of her nestmates.^[4] While the queen handles all the nest reproduction, the subordinates work to care for, defend, and feed the nest instead. The divisions of labor within the nest correlate with the ages of the red paper wasps.^[5] Aside from the female division of labor, male red paper wasps engage in two alternative mating tactics: the role of the territorial male (who chases away intruding patrollers) and the role of the patroller (who flies from tree to tree and does not chase other males).^[1] The study of the dominance relations within the Polistes canadensis has provided insight into the social organization that characterizes many social invertebrates.^[4]

Taxonomy and phylogeny

The Swedish zoologist Carl Linnaeus described the red paper wasp in 1758. Its species name is a New Latin term canadensis used to refer to species associated with Canada. Linnaeus was misinformed about the location of his specimen, because although it has an extensive range throughout the United States, the Polistes canadensis does not reach Canada.^[2]

Part of the social vespidae family, the Polistes genus (the paper wasps) marks the transition between solitary and highly social behavior among wasps. The red paper wasp is a member of the New World Polistes, which are found in the subgenus Aphanilopterus. Because morphological variation among the Aphanilopterus is small, many closely related species such as the Polistes annularis (from the Eastern US to Texas), Polistes erythrocephalus (Nicaragua through Brazil), and Polistes infuscatus (northern South America) had been grouped with the red paper wasps as a species, but now have been determined to be distinct species of their own.^[6]

Description and identification

Both male and female Polistes canadensis have a uniformly light to dark mahogany-brown body, sometimes with the head and thorax of a lighter shade. Its rust colored body gives the red paper wasp its common name. Some feature a yellow apical margin of the first tergum as well. The wings are purplish-black, and the veins and stigma are either black or reddish-brown.^[2] The Polistes canadensis is a large-bodied wasp with a wing length ranging from 17.0 to 24.5 mm.^[7] Nests are constructed from plant fibers such as dry grass and dead wood, which like in other paper wasps, that they mix with saliva to create water-resistant nests made out of papery material. These nests are not covered with an envelope and feature hexagonal cells in which eggs are laid and larvae develop. The Polistes canadensis colony divides its colony among several combs, with an average size of 30.8 cells per comb. A large, mature but still growing colony has about 838 cells distributed among 38 combs.^[1] Combs on vertical (walls and tree trunks) and sloping surfaces (tree limbs) hang with the petiole at the upper end. Colonies on tree trunks tend to add secondary combs above or below the first comb resulting in a linear arrangement. In contrast, nests on horizontal surfaces away from edges (i.e. ceilings) feature eccentric petioles with secondary combs bordering the central primary comb in a semicircular perimeter. No comb has more than one petiole. Petioles average 9.9 mm in length. When combs reach their final size, adult wasps can often step directly from one comb to another; the average distance between petioles of nearby combs is 2.9 cm. Although rare, adjacent combs sometimes grow to touch.^[3]

Distribution and habitat

The red paper wasp is widely distributed over most of the Neotropical region, ranging from the United States to Argentina. Some locations include, but are not limited to, Mexico, Guatemala, Honduras, Colombia, Venezuela, British Guiana, Trinidad, Brazil, Peru, Bolivia and Paraguay.^[2] Nests can often be found on human constructions such as buildings, in open habitats on trunks and large limbs of trees, and in sheltered sites such as in caves, sheds, or under peeling bark.^[3]

Colony cycle

A single female queen with, on average, 9.1 foundresses, usually initiates the construction of new combs and cells to form nests. The more foundresses in a colony, the more combs produced. The preemergence period of comb development lasts from initiation of the first comb until the first adult offspring emerges, when the postemergence period begins. On average, combs grow for 15.4 days and achieve a size of 30.8 cells. Although comb enlargement tends to stop when the oldest brood in that comb had pupated, in many cases the brood is still in the egg stage, so the stage of brood development in the comb does not seem to provide the cue that causes the queen to cease comb development. The egg to adult development period takes around 40 days, during which time the first helpers emerge in the nest. Unlike other species of Polistes, no more than one generation of broods is housed in a comb. Therefore, as older combs are abandoned, brood rearing moves into new combs, with most colonies having about 38 combs. This multiple comb-building tactic is thought to be a defense mechanism against brood loss to moth infestations. Broods are reared year round and two or three generations of offspring are reared during the colony cycle of several months.^[3] The red paper wasp is an asynchronous species, meaning that there appears to be no relationship between the time of the year and the colony’s development. Colonies are observed to be founded or abandoned at any time of the year, but the mean duration of the brood development stages differ between cold-dry and warm-rainy seasons.^[8]

Behavior

Dominance hierarchy

One female queen exercises absolute dominance over all other females. The queen can only effectively control her subordinates if they are within her physical reach. She often suppresses the aggressive behavior of her nestmates through lateral abdominal vibrations and stroking. In contrast to the aggressive actions of the queen, the unmated females are unaggressive and often workers, even if they possess developed ovaries. Even the non-queen female with the most developed ovaries—the queen’s greatest potential challenger—will await the despotic queen’s demise without being aggressive. Because the interactions between non-queens are rarely aggressive, if there is a queen, no other hierarchy amongst the subordinates will be seen. Only when no queen reigns does the frequency of aggressive interactions amongst the foundresses becomes significant; the level of aggressiveness and position within the nest often mirrors the reproductive state of the wasp. As the despotic queen ages and her dominance wanes, the colony cycle will gradually end as the nest is abandoned and the younger females of the nest, with the most developed ovaries, form new colonies.^[4]

Reproductive suppression

While queen control in other Polistes species involves more non-confrontational behavioral, the red paper wasp is characterized by confrontational dominance—the queen will face aggressors and subordinates with physical action that can sometimes escalate into fatal battles. This aggressiveness can especially be seen in the queen’s territorial defense of the egg-laying region of the nest. In the absence of the dominant queen, most of the subordinates within the nest would be capable of laying eggs if given access to empty cells. The queen is able to inhibit the ovary development and reproduction of her subordinates through physical attacks on any subordinates who approach the empty cells and newly laid eggs that she guards.^[4] This is similar to the behavior of queens in Polistes instabilis, who suppress ovary development in workers by performing aggressive dominance interactions.^[9]

Mating behavior

Male red paper wasps engage in two alternative mating tactics: the role of the territorial male (who chases away intruding patrollers) and the role of the patroller (who flies from tree to tree and does not chase other males). Patrollers have a smaller body size than territorial males. Because of their inability to successfully compete against the larger territorial males for territories, these smaller males resort to patrolling. While physical altercations between territorial males and intruding patrollers are rare (<5% of the time) as the intruders typically quickly depart, there is still significant competition over the possession of territories. Owning a territory tends to lead to more copulations although the territory itself does not hold any resources or nests. The defense of these territories represents a lek polygyny mating system that is not based on resources. But rather, females perhaps incentivize males to compete for territories by restricting mating to these territories; this way, females know that the territorial males they choose to mate with are strong, healthy males. To attract females to the territories, males can rub their abdomens across the territories to apply pheromones. The patrollers do not just wait around for territories to be vacated; they will sneak matings with females in territories when the territorial males are temporarily away or distracted.^[1]

Kin selection

As they work to feed, rear, and defend the nest, helper red paper wasps forgo personal reproduction in order to care for the nest. These workers are closely related to their female nestmates (r = 0.47 ± 0.049; n = 28 nests, 145 wasps) and may not only obtain high indirect fitness by helping their nestmates, but also may gain direct fitness (in terms of personal reproduction) by taking over the reproductive role of the queen if the queen dies. Many female workers choose to care for their nestmates while waiting to become breeders in their original nests. But for younger female workers, the benefits of direct fitness may be greater than the benefits of indirect fitness. These younger females can choose to leave their original nests for a more uncertain future (in terms of indirect and direct payoffs) to co-found a new nest. For older females who are more limited reproductively, however, the best and perhaps only option would be engage in risky foraging tasks to take care of others within their nests, thereby maximizing their own indirect fitness.^[10]

Costs and benefits of sociality

Because new nests are often founded near the parent nest by a group of sisters, wasps in neighboring nests tend to be closely related (Mantel test, r=-0.138, p<0.05). Therefore, workers are able to best maximize their indirect fitness by not just taking care of their home nest, but by helping out several related nests nearby. Within the red paper wasps, 56% of females have drifted to nearby nests. While workers share the most genetic similarity with their home nestmates, these genetic benefits are outweighed by the high predation risk associated with only investing in one nest. An egg in a nest has a 40% chance of being eaten by a predator before reaching adulthood. Therefore, drifting to invest in nearby nests increases the chances that at least some of the workers’ investments survive in the face of whole-nest predation. In addition, the presence of nestmates in high nest density can also reduce predation risk—with large number of nestmates, while some workers forage, significant numbers of nestmates can remain to defend the nest.^[11]

Defense

Appropriate visual stimuli, such as the movements of nearby large or dark colored objects, can elicit attacks by the red paper wasps, which will fly at the object in alarm, attempting to sting it. An alarmed Polistes canadensis can release venom onto its nest; the odor of venom will prompt alarm responses in its nestmates, lower their thresholds for attack, and even attract more nestmates to the alarm. Through this chemical means of communicating alarm, the colony is able to rise quickly with its sting chambers open to defend its nest against predators. Because red paper wasps uniquely occupy multiple combs unattached to each other, the chemical alarm substance may have arisen as a necessary adaptation for a more efficient alarm for a sparsely dispersed nest.^[12] It is not known whether these red paper wasps can release venom at the nest independent of stinging behavior in order to communicate alarm to nestmates. While these wasps have been seen to open the sting chamber independently of venom release on the nest, the solo release of venom has not been observed. The Polistes canadensis may open its sting chambers as part of a defensive display (involves bending of the gaster toward the source of alarm) or in preparation for stinging.^[13]

Parasites

Present in a majority of colonies, the tineid moth (belongs to an undescribed genus) is the most common pest of the red paper wasps. The moth lays its eggs and then these eggs hatch into larvae that can burrow from cell to cell and prey on meconia and wasp pupae. While only 0.47% of the brood is lost to predation, the wasps perform many defense mechanisms to protect their nests from these infestations. Adult wasps attempt to remove and kill moth eggs and larvae by chewing down the edges of cells, coating the cells with an oral secretion that gives the nest a dark brownish appearance. The colonies then do not reuse these infested combs to raise another brood. The wasps also clip disused combs, reducing the rate of reinfestation by moths. In some instances, though, the comb clipping is also used to provide room for the construction of new combs. One of the key reasons the Polistes canadensis nests across multiple combs rather than building a single comb like much of the rest of its genus is as a defense mechanism against the infestation of these moths. The average amount of nesting material required to build a new cell in a comb decreases as the number of cells in the comb increases. Therefore, there are many inefficiencies associated with constructing several smaller combs rather than single larger combs like much of the rest of the Polistes genus does. For example, with the construction of multiple combs, comes the need to build a separate petiole for each comb. By building multiple combs and failing to reutilize them, the costs Polistes canadensis pay to defend its nests are significant and quite large in terms of energy expended in nest construction per brood reared.^[3] Other common parasites include three species of the obligate hymenopterous parasitoid: Seminota, Toechorychus albimaculatus and the Pachysomides iheringi. 4.4% of the red paper wasp pupae are lost to these Hymenoptera.^[3]

References

1. Polak, Michal (1993). Competition for Landmark Territories among Male Polistes canadensis (L.) (Hymenoptera: Vespidae): Large-size Advantage and Alternative Mate-acquisition Tactics. Behavioral Ecology 4(4):325–31.
2. Bequaert, Joseph C. (1940). An Introductory Study of Polistes in the United States and Canada with Descriptions of Some New North and South American Forms (Hymenoptera; Vespidæ). Journal of the New York Entomological Society 48(1): 1–31.
3. Jeanne, Robert L. (1979). Construction and Utilization of Multiple Combs in Polistes canadensis in Relation to the Biology of a Predaceous Moth. Behavioral Ecology and Sociobiology 4(3): 293–310.
4. West-Eberhard, Mary Jane (1986). "Dominance Relations in Polistes canadensis (L.), a Tropical Social Wasp." Monitore Zoologico Italiano 20(3): 263–81.
5. Giray, T; Giovanetti, M; West-Eberhard MJ (March 2005). "Juvenile hormone, reproduction, and worker behavior in the neotropical social wasp Polistes canadensis". Proc. Natl. Acad. Sci. U.S.A. 102 (9): 3330–5. doi:10.1073/pnas.0409560102. PMC: 552932. PMID 15728373.
6. Pickett, Kurt M., and John W. Wenzel (2004). Phylogenetic Analysis of the New World Polistes (Hymenoptera: Vespidae: Polistinae) Using Morphology and Molecules. Journal of the Kansas Entomological Society 77(4): 742–60.
7. O'donnell, Sean, and Robert L. Jeanne. (1991). Interspecific Occupation of a Tropical Social Wasp Colony (Hymenoptera: Vespidae:Polistes)." Journal of Insect Behavior 4(3): 397–400.
8. Torres, Viviana De Oliveira. (2008). Bionomics Aspects of the Neotropical Social Wasp Polistes Canadensis Canadensis (Linnaeus) (Hymenoptera, Vespidae)." Revista Brasileira De Entomologia 53(1): 134–8.
9. Molina, Yamile (2006). "Mushroom Body Volume is Related to Social Aggression and Ovary Development in the Paper Wasp Polistes instabilis". Brain Behavior and Evolution.
10. Sumner, S., H. Kelstrup, and D. Fanelli (2010). Reproductive Constraints, Direct Fitness and Indirect Fitness Benefits Explain Helping Behaviour in the Primitively Eusocial Wasp, Polistes canadensis. Proceedings of the Royal Society B: Biological Sciences 277(1688): 1721–8.
11. Sumner, Seirian, Eric Lucas, Jessie Barker, and Nick Isaac (2007). Radio-Tagging Technology Reveals Extreme Nest-Drifting Behavior in a Eusocial Insect. Current Biology 17(2): 140–5.
12. Jeanne, R.L. (1982). Evidence for an Alarm Substance In Polistes canadensis. Experientia 38(3): 329–30.
13. Ross, Kenneth G., and Robert W. Matthews. The Social Biology of Wasps. Ithaca: Comstock Pub. Associates, 1991.