Polistes biglumis | |
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Female | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Hymenoptera |
Family: | Vespidae |
Subfamily: | Polistinae |
Tribe: | Polistini |
Genus: | Polistes |
Species: | P. biglumis |
Binomial name | |
Polistes biglumis (Linnaeus, 1758) | |
Synonyms | |
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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. [1]
Swedish zoologist Carl Linnaeus described Polistes biglumis in 1758. [2] Its species name biglumis is a Latin phrase meaning "two husks." While no common name for the wasp exists in English, it is referred to as berg Feldwespe in German (meaning "mountain field wasp"). P. biglumis has been studied alongside Polistes snelleni and Polistes chinensis for comparison. P. biglumis was originally classified as a hornet in the genus Vespa, but was reassigned to the genus Polistes, which is the largest genus of paper wasps in the family Vespidae. [3]
This species resides mainly in mountainous zones in Southern Europe, and it is the only paper wasp that inhabits a mountainous clime. Due to its divergence from the more common temperate climes of its genus, it has developed several distinctions from other paper wasps that arise mainly from selective pressure due to the severe climate it commonly experiences. The climate shortens the nesting season of P. biglumis to around four months. In addition, the nests exhibit darker coloration, which allows them to absorb more heat from the sun. [1] Workers are also often nonexistent in some populations of P. biglumis, which demonstrates a large dichotomy from other social wasps, whose most commonly seen specimens are the workers. Finally, because of the reduced nesting season and parasitism by other wasps, P. biglumis has developed nesting strategies that are both distinct from other paper wasps and variable among conspecific populations. [4]
Polistes biglumis can reach a length of up to 16 mm (0.63 in) (queen), 14 mm (0.55 in) (workers), 15 mm (0.59 in) (males). It is a larger species of wasp in comparison to its relatives in Polistes. It also exhibits darker coloration compared to other paper wasps; it has a black petiole for both sexes. The females exhibit black abdomens, as well, with rare yellow spots. Males, however, have largely yellow abdomens. [3] Adult wasps can be distinguished from the young because they are darkly colored and can fly, whereas young wasps have paler stripes and are flightless. [7]
Workers and queens do not exhibit morphological differences, but they can be distinguished physically by the abundance of their fat layers and behaviorally through their relative foraging efforts. Queens have more abundant fat layers and are also significantly less likely to participate in foraging for the nest. [4] In addition, the color of the fat layers is distinct for workers versus queens; the workers exhibited yellow fat layers while the queens exhibited milky fat layers. [1]
The nests of P. biglumis are circular or elliptical in shape and are hung vertically by one pedicel. They are largely produced by the foundress during the phases of egg and larval production. The peripheral cells are generally uninhabited. Nests are more likely to be closer to the ground versus other Polistes wasps because the ground offers thermal inertia and shelter from strong winds. [1] The nest material is also key to nestmate recognition because nest surfaces are impregnated with the epicuticular hydrocarbons that establish the framework by which the individuals discriminate between other wasps. [2]
This species resides mainly in mountainous zones in Southern Europe, [8] especially in Italy and France, including the relatively cooler climate of the Alps and the relatively warmer climate of the Apennines (although neither climate is temperate, making P. biglumis anomalous among Polistes species). [4] Montgenèvre, a commune located in the French Cottian Alps, has been used extensively as a region to study the behavior of the wasp in its natural environment. [7] [9] In addition, P. biglumis can be found extending into Uzbekistan, Sweden, Germany, and Austria. [3] While a few ecologists attribute the name P. biglumis to a wasp species inhabiting Okkaido, Japan, as well, the Japanese species is an undescribed species more closely related to P. nimpha than to the predominantly European P. biglumis described here. [1] The species found in Japan has been closely studied alongside Polistes snelleni , another common Japanese paper wasp. The wasps of this species generally nest on the sides of rocks in meadows in the mountains or in alpine areas consisting of Pinus sylvestris and Larix decidua . [1] [10] The colonies inhabiting these nests are both small and rare, as the colony cycle for the wasp is truncated to only four months. On average, the colonies consist of about 30 individuals. [4]
P. biglumis colonies are always founded by a single foundress wasp. [11] New colonies become active in late May or early June each year, and the colony cycle terminates in September. [4] Half of all P. biglumis nests fail during the pre-emergence stage, and due to the limited colony cycle, foundresses cannot start a new nest for the season. [11] The egg stage of the P. biglumis wasp is around 2 weeks, and the male offspring are produced before female offspring in a sequential fashion. [11] [12] The rate of appearance of future queens, however, is affected by environmental factors in the region; early female offspring in cold areas with high parasitism have fatter, gyne-like bodies and less foraging effort than do female offspring in warm areas with low parasitism. Both the climate and the parasite prevalence affect the first female offspring's abundance of fat bodies independently of one another, but only climate affects the first female offspring's foraging effort. Parasitism has no effect on late female offspring, but climate still affects the fat layers of their bodies. P. biglumis may have been selected for suppressed worker production in the first brood, the only brood not destroyed by parasites, so that new queens would survive to produce new colonies. [4] The colony cycle is characterized by a pre-emergence period that lasts from foundation by the single gyne of the colony to the emergence of the first new worker, and a postemergence period, from the emergence of the new worker to the end of the cycle (as an annual species, this marks the end of the colony). [1] At the end of a season, the future queen females of the colony overwinter to reproduce in the spring. [4]
Polistes biglumis wasps exhibit a homogeneous odor that is both species- and colony-specific, and it is used by individuals to recognize nestmates. [9] This system of recognition uses epicuticular-saturated hydrocarbons. The queen produces the odor specific epicuticular hydrocarbons and transfers them to the nest paper. Wasps learn to recognize nestmates by the nest paper odor, which serves as a template for recognizing any wasp and determining whether or not to act aggressively toward it. [12] Although epicuticular hydrocarbons cover the adults and larvae of Polistes biglumis, as well as their nest surfaces, newly emerged P. biglumis wasps are accepted in foreign colonies of the same species, whereas adult non-nestmates are met with violent aggression, usually until the invading wasp is killed. [2] The distinction between aggression toward foreign newly emerged versus adult wasps could arise from a low level of epicuticular hydrocarbons developed in newly emerged wasps with respect to adult wasps. This low level is tantamount to a weakening of the signal. Similarly, a dead adult wasp that has been stripped of its epicuticular hydrocarbon layer elicits no reaction in a foreign nest. [9]
P. biglumis females have also developed a method for discriminating between their own and foreign conspecific eggs that will become queens; they tend to eat foreign wasp eggs. Cues for distinguishing the eggs are located on the eggs themselves. This discrimination is a useful tactic both for foundresses that are in conflict with usurpers or joiners, and for usurpers and joiners themselves. No such system of recognition exists for eggs that develop into workers. [12]
P. biglumis males typically return to sunlit landmarks in their patrol flights repetitiously. Males adopt certain tiny territories, usually small stones or scrubs within a larger mating region that they patrol, defend against intruders, and mark with scent. Both the suitability of the microclimate of the territory adopted by the male and the conspicuousness of the territory against the terrain affect the mate-locating efficacy of the species. Males attempt to copulate with females resting on their habitual perches in their discrete territories within a larger mating aggregation, characteristic of a lek mating system. Landmark mating in this system is advantageous for the promotion of outbreeding. [10] Each lek is composed of related males that capture contiguous territories. One-third of all P. biglumis females mate with multiple male partners, all of whom aree related. P. biglumis females' polyandrous activity make them unique among all other paper wasps. [11]
Since a large set of nests will fail during the nesting season, foundresses constrained by the limited nesting season are forced to attempt female usurpation, where females take over conspecific nests. Thus, nests often have multiple unrelated females. The nests are either fully usurped, where the original female leaves; joined, where the original female remains; or both usurped and joined by other female wasps. Usurpers and joiners are not related to the original nest owner, but instead were random wasps from the overall population, which differs from other Polistes wasp joiners that always join with relatives. [11] Usurping and joining are both inferior strategies to founding a new nest and are only fractionally successful, so these strategies are only chosen after nest failure when the breeding female is not a resident of any nest and is searching for alternative reproductive opportunities. In each case of serial female nesting, the original nest owner no longer has control of the nest. In addition, the newcomer destroys part of the original brood. All new breeding is by the intruder, and kin-selected benefit via raising the new brood is eliminated for the original female's emerging brood. [11]
The average sex ratio of P. biglumis is female biased; however, the specific sex ratio varies from brood to brood from virtually all male to all female. Furthermore, the sex ratios vary temporally, as well, and the sex ratios are biased toward females more toward the end of the pre-emergence period. The presence of multiple females also affects the sex ratios, and single matrilineal nests were more male biased than were nests that underwent serial breeding by multiple females. Because foundresses produce a large number of male offspring early in the season, and nest founders produce similar sex ratios to begin with, a section of the male offspring of the original foundress is culled by usurpers and joiners.
P. biglumis has a haplodiploid sex determination system. All males among a male brood exhibit the relatedness of full brothers, indicating that they are haploid and generally produced by one female. Females, though, are diploid. Sex ratio analysis indicates that the female producing all of the male workers was the original nest founder. [11]
Nest predation on the wasp is minimal throughout its nesting season; however, when it does occur, it is mainly perpetrated by birds. [1] Nevertheless, nest predation by vertebrates reaches its peak during the pre-emergence period when the nest is empty except for the foundress and is, as a result, undefended. Furthermore, such predation is a major contribution to nest failure. [13] In these cases, often the nest is preyed upon, but the foundress survives, which leads to the creation of usurpers and joiners. [11] [13] Unlike with other Polistes species, ants do not often attack the nests of P. biglumis. Accordingly, P. biglumis releases a very reduced amount of ant repellant substance on to the pedicels of the nests. In most Polistes species, this substance is secreted via abdominal glands on pedicels to prevent ant invasion of the comb. [1]
Polistes atrimandibularis is a rare, obligate parasite that permanently invades the nest of the P. biglumis colonies. Its prevalence is highest in relatively cooler P. biglumis habitats in the Alps and is scarcer in their warmer habitats in the Apennines. The parasite wasp queen invades the nest of the foundress host wasp during the pre-emergence phase when the nest is empty except for the foundress. The parasite wasp destroys all of the host eggs, and then represses the foundress's egg-laying capacity, cutting her productivity in half. As a result, often only the first of the P. biglumis wasp broods survive, namely the brood that emerged early and escaped destruction by the parasite queen. [4]
The parasite enters the host nest peacefully and submits to the attacks of the foundress, but over time, the parasite queen begins to dominate the host queen. The parasite queen co-opts the host workers and the host queen to care for her brood by altering their processes of nestmate recognition. Parasite queens also help care for the larvae toward the end of the colony cycle. She enters other local colonies of the host species to steal their larvae and pupae. She uses the pupae and larvae of the foreign colony P. biglumis to feed her own larvae in the parasitized nest. Intrusion and survival of the parasite wasp in the host foundress's nest is based largely on its ability to manipulate various components of the recognition system used by the host wasp. [7]
Parasite queens have levels of epicuticular hydrocarbons key to recognition that are one-third to one-fourth the levels in host wasps, which facilitate a weaker signal that allows them to insinuate themselves into host colonies. [7] As time progresses, the parasite queen can camouflage her odor as a host wasp; at the point of emergence for the workers, the parasite queen is indistinguishable from the host queen. [2] [9] The adult parasitic offspring have no such mechanism for deception and they have unsaturated epicuticular hydrocarbons specific to their species that do not occur in the host species. However, the different odors of parasite and host species do not affect the acceptance by the host species workers of the parasite brood in parasitized nests. As a result, parasitized colonies do not have a homogeneous odor because parasite offspring have a different odor from hosts.
The survival of the parasite offspring relies on their queen to alter the nest paper from which the new host workers learn nestmate recognition. [9] The parasite wasp impregnates the nest paper with unsaturated, parasite-species hydrocarbons so that the emerging host workers learn to recognize both host and parasite broods as nestmates. [2] As a result, newly emerged parasite wasp females are only accepted in P. biglumis colonies that have already been parasitized. In nonparasitized colonies, they receive highly aggressive responses from host wasps. Thus, the host wasps in parasitized colonies learn a recognition odor template that is more inclusive than the one used by wasps of nonparasitized colonies. [9] This expanded template leads to a far greater error rate in nestmate recognition for the host wasp. The parasitized nest hosts demonstrate an impairment in discrimination, and are much more likely to permit even non-nestmate conspecific individuals. They are also more likely to reject nestmates erroneously. [2]
Polistes is a cosmopolitan genus of paper wasps and the only genus in the tribe Polistini. Vernacular names for the genus include umbrella wasps, coined by Walter Ebeling in 1975 to distinguish it from other types of paper wasp, in reference to the form of their nests, and umbrella paper wasps. Polistes is the single largest genus within the family Vespidae, with over 200 recognized species. 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 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.
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.
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.
Polistes chinensis is a polistine vespid wasp in the cosmopolitan genus Polistes, and is commonly known as the Asian, Chinese or Japanese paper wasp. It is found in East Asia, in particular China and Japan. The subspecies P. chinensis antennalis is an invasive species in New Zealand, having arrived in 1979.
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.
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. P. metricus is dark colored, with yellow tarsi and black tibia. Nests of P. metricus can be found attached to the sides of buildings, trees, and shrubbery.
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.
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.
Polistes carolina is one of two species of red paper wasp found in the eastern United States and is noted for the finer ridges on its propodeum. It is a social wasp in the family Vespidae and subfamily Polistinae. The species is native to the United States from Texas to Florida, north to New York, and west to Nebraska. The wasp's common name is due to the reddish-brown color of its head and body. P. carolina prefer to build their nests in protected spaces.
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.
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.
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.
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. The most widely distributed American species of the genus Polistes, it colonizes multiple combs, which it rears year-round.
Polistes atrimandibularis is one of four 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.
Polistes austroccidentalis is a kleptoparasitic paper wasp that is found in several regions of high altitude in Europe, and until 2017 was universally mistakenly referred to as Polistes semenowi, which is instead the correct name of the species formerly known as "Polistes sulcifer". As one of only four obligate parasites in the subgenus Polistes, it uses the nests of other paper wasps to rear its young. To evade detection by the host nest, P. austroccidentalis employs mimicry by adjusting its cuticular hydrocarbons to match those of the host. Once the host nest has been infiltrated, the parasitic female physically attacks the host queen to subdue her and become the colony's new queen. P. austroccidentalis displays several morphological adaptations for parasitism such as increased mandible size and an enlarged Van der Vecht's organ. This species is unusual because it does not have the ability to produce workers and is only able to produce individuals who have the capacity to reproduce.
Polistes semenowi is a species of paper wasp in the genus Polistes that is found in southeastern and southern central Europe, as well as central Asia, and was until 2017 erroneously known by the name Polistes sulcifer, while a different species was incorrectly believed to represent P. semenowi. It is one of only four known Polistes obligate social parasites, sometimes referred to as "cuckoo paper wasps", and its host is the congeneric species Polistes dominula. As an obligate social parasite, this species has lost the ability to build nests, and relies on the host workers to raise its brood. P. semenowi females use brute force, followed by chemical mimicry in order to successfully usurp a host nest and take over as the queen.
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 substrate 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.
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.
Nest usurpation is when the queen of one species of eusocial insects takes over the colony of another species.