Polistes canadensis

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Polistes canadensis
Polistes canadensis.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Vespidae
Subfamily: Polistinae
Tribe: Polistini
Genus: Polistes
Species:
P. canadensis
Binomial name
Polistes canadensis
Synonyms [1]

Polistes canadensis is a species of red paper wasp found in the Neotropical realm. It is a primitively eusocial wasp as a member of the subfamily Polistinae. A largely predatory species, it hunts for caterpillar meat to supply its colony, often supplementing its developing larvae with nectar. [2] The most widely distributed American species of the genus Polistes , it colonizes multiple combs, which it rears year-round. [3]

Contents

Emerging from hibernation in the spring, the females found nests built from 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. [2] 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. On average, a single female queen with 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. [4] 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. [5]

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 Polistes canadensis. [6] Aside from the female division of labor, male Polistes canadensis 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). [2] The study of the dominance relations within the Polistes canadensis has provided insight into the social organization that characterizes many social invertebrates. [5]

Taxonomy and phylogeny

The Swedish zoologist Carl Linnaeus described this species in 1758. The species epithet is a New Latin term canadensis used to refer to organisms associated with Canada. Linnaeus was misinformed about the origin of his specimen, because although it has an extensive range throughout North America, Polistes canadensis only occurs as far north as southern Arizona. [3] [7]

Part of the social Vespidae, Polistes (the true paper wasps) marks the transition between solitary and highly social behavior among wasps. Polistes canadensis is a member of the New World Polistes subgenus Aphanilopterus, and closely related to Polistes annularis (from the Eastern US to Texas), Polistes erythrocephalus (Nicaragua through Brazil), and Polistes infuscatus (northern South America). [8]

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. [9] Its rust-colored body gives this species of red paper wasp its common name.[ citation needed ] 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. [3] Polistes canadensis is a large-bodied wasp with a wing length ranging from 17.0 to 24.5 mm. [9]

Nests are constructed from plant fibers such as dry grass and dead wood, which, as with other paper wasps, are mixed with saliva to create water-resistant nests made out of papery material. These nests are not covered with an outer envelope and feature hexagonal cells in which eggs are laid and larvae develop. A growing Polistes canadensis colony often engages in fission into several combs, with an average size of 30.8 cells per comb. A large, mature but still growing colony may have over 800 cells distributed among over 30 combs. [2] 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 fuse if they make contact. [4]

Distribution and habitat

Polistes canadensis is widely distributed over most of the Neotropical region, ranging from Arizona to Argentina. Some locations include, but are not limited to, Mexico, Guatemala, Honduras, Colombia, Ecuador, Venezuela, British Guiana, Trinidad, Brazil, Peru, Bolivia and Paraguay. [3] [10] [7] 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. [4]

Colony cycle

On average, a single female queen with 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. [4] Polistes canadensis 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. [11]

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 non-queens' interactions 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 become 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. [5]

Reproductive suppression

While queen control in other Polistes species involves more non-confrontational behavior, Polistes canadensis is characterized by confrontational dominance—the queen will face aggressors and subordinates with a 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. [5] This is similar to the behavior of queens in Polistes instabilis , who suppress ovary development in workers by performing aggressive dominance interactions. [12]

Mating behavior

Male Polistes canadensis 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. [2]

Kin selection

As they work to feed, rear, and defend the nest, helper Polistes canadensis 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. [13]

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 Polistes canadensis, 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. [14]

Defense

Appropriate visual stimuli, such as the movements of nearby large or dark-colored objects, can elicit attacks by Polistes canadensis, 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 Polistes canadensis 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. [15] It is not known whether Polistes canadensis 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. [16]

Parasites

Present in a majority of colonies, the tineid moth (belongs to an undescribed genus) is the most common pest of Polistes canadensis. 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, many inefficiencies are associated with constructing several smaller combs rather than single larger combs like much of the rest of the Polistes. For example, the need to build a separate petiole for each comb comes with the construction of multiple combs. 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. [4] Other common parasites include three species of the obligate hymenopterous parasitoid: Seminota, Toechorychus albimaculatus and the Pachysomides iheringi. 4.4% of Polistes canadensis pupae are lost to these Hymenoptera. [4]

Related Research Articles

<i>Polistes</i> Genus of insects

Wasps of the cosmopolitan genus Polistes are the most familiar of the polistine wasps, and are the most common type of paper wasp in North America. Walter Ebeling coined the vernacular name "umbrella wasps" for this genus in 1975 to distinguish it from other types of paper wasp, in reference to the form of their nests. It is also the single largest genus within the family Vespidae, with over 300 recognized species and subspecies. 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. The European paper wasp, Polistes dominula, was introduced into the US about 1981 and has quickly spread throughout most of the country, in most cases replacing native species within a few years. This species is very commonly mistaken for a yellow jacket, as it is black, strongly marked with yellow, and quite different from the native North American species of Polistes. The cuckoo wasp, Polistes semenowi, is an obligate social parasite, whose only host is P. dominula. Polistes metricus adults malaxate their insect prey by chewing them into a pulp, sucking out and ingesting the body fluids, then feeding the rest of the morsel to their larvae. The most widely distributed South American wasp species, Polistes versicolor, 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. Polistes wasps can be identified by their characteristic flight; their long legs dangle below their bodies, which are also more slender than a yellow jacket.

<span class="mw-page-title-main">European paper wasp</span> Species of wasp

The European paper wasp is one of the most common and well-known species of social wasps in the genus Polistes. Its diet is more diverse than those of most Polistes species—many genera of insects versus mainly caterpillars in other Polistes—giving it superior survivability compared to other wasp species during a shortage of resources.

<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>Ropalidia marginata</i> Species of insect

Ropalidia marginata is an Old World species of paper wasp. It is primitively eusocial, not showing the same bias in brood care seen in other social insects with greater asymmetry in relatedness. The species employs a variety of colony founding strategies, sometimes with single founders and sometimes in groups of variable number. The queen does not use physical dominance to control workers; there is evidence of pheromones being used to suppress other female workers from overtaking queenship.

<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 metricus</i> Species of wasp

Polistes metricus is a wasp native to North America. In the United States, it ranges throughout the southern Midwest, the South, and as far northeast as New York, but has recently been spotted in southwest Ontario. A single female specimen has also been reported from Dryden, Maine. Polistes metricus is dark colored, with yellow tarsi and black tibia. Nests of Polistes metricus can be found attached to the sides of buildings, trees, and shrubbery.

<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 is found throughout the United States, Mexico, the Bahamas, Jamaica and parts of Canada. Due to solitary nest founding by queens, P. exclamans has extended its range in the past few decades and now covers the eastern half of the United States, as well as part of the north. This expansion is typically attributed to changing global climate and temperatures. 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 instabilis</i> Species of wasp

Polistes instabilis, a type of paper wasp, is a neotropical, eusocial wasp that can be found in tropical and subtropical areas such as Central America and South America. It can be easily identified with its characteristic yellow, brown, and reddish markings, and it builds nests made from chewing plant fibers and making them into paper.

<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 snelleni</i> Species of wasp

Polistes snelleni, the Japanese paper wasp, is a common social wasp species in central and northern Japan. P. snelleni is also found in northern China, Korea, and the Russian Far East. Due to the different climates in these regions, P. snelleni is able to adapt to different temperatures and climatic conditions. P. snelleni is typically found in hilly or submontane areas, so they are classified in the semi-highland category.

<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 atrimandibularis</i> Species of wasp

Polistes atrimandibularis is one of three obligate social parasites among the Polistes wasps found in Europe. Of the four social paper wasp parasite species known, it is the smallest. It parasitizes multiple species such as P. dominula, P. nimpha, P. associus, P. gallicus, and P. biglumis. Females of P. atrimandibularis are unable to build a nest or produce workers, and therefore rely entirely on the host colony.

<i>Belonogaster petiolata</i> Species of wasp

Belonogaster petiolata is a species of primitively eusocial wasp that dwells in southern Africa, in temperate or subhumid climate zones. This wasp species has a strong presence in South Africa and has also been seen in northern Johannesburg. Many colonies can be found in caves. The Sterkfontein Caves in South Africa, for example, contain large populations of B. petiolata.

<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.

Ropalidia plebeiana is a eusocial temperate paper wasp. It is unique, as it is the only temperate wasp in the typically tropical Ropalidia genus. R. plebeiana is widely distributed in eastern Australia, and recently have been found making huge nest aggregations, with thousands of nests on trunks of trees, in south-eastern New South Wales.

<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 substratum 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. The P. versicolor colony cycle broadly ranges from 3 to 10 months, although there appears to be no relationship between the colony's development and the season of the year. While yellow paper wasps do have clear annual colony cycles, many young queens have the opportunity to hibernate during the winter, forming optional winter aggregations. Dominance hierarchies within these aggregations are characterized by physical aggression of the dominant female(s) towards the associated females, who tend to be sisters. Wagging movements are also often used as a form of communication within the colony. The yellow paper wasp is generally predatory, capturing a wide range of insects, although it often feeds on pollen and nectar as well. Therefore, P. versicolor can be useful as a pollinator or as effective pest control.

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

Polistes dorsalis is a species of social wasps that can be found throughout various parts of North America. It is classified under the Vespidae within the genus of Polistes. Male Polistes dorsalis wasps can be distinguished from other Polistes species by their distinctly prominent median tubercle of sternum 7. Both sexes can also be recognized due to their v- shaped yellow markings on their head. They are distributed widely across North America and can be found in sheltered nests, typically closer to the ground. These wasps live in a dominance hierarchy in which the queen's role differs from that of ordinary workers. When threatened, these wasps can deliver moderately painful stings. Their venom might also be of human interest for their antimicrobial uses.

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.

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