Polistes atrimandibularis | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Hymenoptera |
Family: | Vespidae |
Subfamily: | Polistinae |
Tribe: | Polistini |
Genus: | Polistes |
Species: | P. atrimandibularis |
Binomial name | |
Polistes atrimandibularis Zimmermann, 1930 | |
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. [1]
P. atrimandibularis is in the subfamily Polistinae (paper wasps). It is closely related to the other three obligate social parasites: P. austroccidentalis , P. maroccanus , and P. semenowi . [2] These four Polistes inquiline species are more closely related to P. nimpha and P. dominula rather than to P. gallicus and P. biglumis. [2]
P. atrimandibularis is the smallest of the social Polistes parasites. The size of the host's species brood cells determines P. atrimandibularis size, with the smaller P. atrimandibularis emerging from the nests of P. gallicus and P. associus. Likewise, the larger P. atrimandibularis emerge from the nests of P. dominula instead. This paper wasp has enlarged mandibles, used as weapons to injure hosts that resist parasite intrusion. They also have an enlarged first femur and a longer posterior tibia that is useful for when they must maintain a dominant position within the host colony. [3]
P. atrimandibularis is rare in Europe, found mainly around the Mediterranean and Caspian basin, [1] but it ranges from south and south-central Europe to western Asia. [4] They typically position themselves at higher elevations, although it is not unheard of to find colonies at lower elevations. [5] Accordingly, one of the species that P. atrimandibularis parasitizes, P. biglumis, lives mainly in montane climates in Southern Europe. [6] They do not possess the ability to build their own nests, so they must parasitize other wasps’ colonies.
Similar to the other Polistes social parasites, P. atrimandibularis usurps colonies in the late spring, which is roughly one month before the emergence of host workers. [5] This allows the parasite sufficient time to reproduce and exploit the worker force. [3] As soon as the usurper has successfully dominated the host's queen, it begins to lay its own eggs, and will remain in the host colony until the end of its life cycle. [5] It is not until late summer that the newly emerged parasites migrate to the mountains to mate [5] and then overwinter for several months under a thick blanket of snow and ice. [3]
When P. atrimandibularis mate, males and females migrate to the tops of high mountains. At the top of the mountains, the males will occupy and aggressively defend their mating territories from male competitors. [5] Once inseminated, females remain for overwintering for several months, under a thick blanket of snow and ice. To ensure that they arrive and usurp their host colonies at the correct time, P. atrimandibularis postpone the overwintering exit. These severe, high altitude climate conditions extend parasite diapause for about a month compared to the lowland host colonies, which allows the P. atrimandibularis to usurp colonies that are more developed. Since Polistes social parasites are very rare, these altitudinal migrations also give them an advantage. The migrations promote encounters between the two sexes and give more opportunities for matings between non-relatives. [3]
Each colony has its own unique hydrocarbon chemical signature that allows P. atrimandibularis to distinguish nestmates from non-nestmates. [7] When a female Polistes wasp first emerges, she is unaware of the colony's unique odor. However, after a few hours the female Polistes wasp will be able to recognize and identify the odor of the nest and create a template that it can refer to in the future. [8] Being a social parasite, the queen has the ability to modify its own chemical signature along with the host's nest signature. [7]
Studies have shown that the presence of P. atrimandibularis lowers the reproductive capacity of the host queen and also inhibits the fertility of host workers. Host foundresses in colonies that were parasitized by P. atrimandibularis were found to have repressed ovarian development, with the female parasite becoming the sole female laying eggs in the colony. [9] P. atrimandibularis is interesting in that the usurper cohabits with the host foundress for a long period of time. This suggests that the host foundress is needed in order to exert reproductive control on the host workers.
While the other Polistes social parasites employ more aggressive tactics of usurping host colonies, P. atrimandibularis has a different approach. P. atrimandibularis uses a passive strategy to successfully invade and control a host nest. [3] They do not have specific strategies for each of the host species they invade, instead, they utilize a generalized strategy for all of their invasions. This passive strategy consists of initial submission to frequent attacks of the host foundress, followed by the parasite gradually executing more and more dominant acts against the host foundress. [3] Once the host foundress is subdued and the parasites take possession of the nest, they destroy the host's immature brood, eggs, and small larvae. [3]
Unlike the other Polistes social parasites, P. atrimandibularis simultaneously usurps surrounding host nests, in addition to invading the primary nest which are used solely for reproductive purposes. From these nests, the parasites take larvae and pupae to use as food for feeding their own larvae. [3] By attacking and raiding secondary host nests, the parasites reduce host reproductive success and in return, provide great advantages to the primary nest. Despite P. atrimandibularis passive strategy for usurping colonies, they still maintain morphological fighting characteristics to aid them in their aggressive behavior while pillaging secondary nests. [3]
The host species that P. atrimandibularis decides to invade, affects the size of the newly emerging parasites. [5] Female P. atrimandibularis that were raised by the larger P. dominula, were bigger than others raised by smaller species such as P. associus and P. gallicus. The bigger parasites were the ones that became the most successful usurpers, with only a few of the smaller parasites succeeding. [5] The newly emerging P. atrimandibularis coming from P. dominula host colonies match the size of successful usurpers, suggesting that selection is acting on a population of parasites to increase average sizes and to favor the host P. dominula. [5]
In order to fully take control and integrate themselves into the host colony, the P. atrimandibularis queen needs to alter her distinct chemical order, as to not get rejected by the hosts. Before invading host colonies, the parasites have a species-specific epicuticular signature: unsaturated hydrocarbons (alkenes), which are absent in host species. [8] After the invasion, these alkenes start decreasing in the parasite epicuticular profiles, and compounds abundant in the host chemical signature start increasing. [7] Within one month, when the host workers emerge, the female P. atrimandibularis’s chemical signature perfectly mimics the host signature. At this point the parasites and host foundress are chemically indistinguishable. [8] There have been cases where P. atrimandibularis was so successful in fooling the host species, that the hosts tended the parasite's larvae to maturity weeks after the parasites had left. [5]
P. atrimandibularis queens need the one month before host workers emerge to reproduce and to modify the host's chemical signature. They do so by supplementing the paper nest surface with parasite specific compounds. [7] By the time host workers emerge, they have already learned the modified colony chemical signature, containing both original and parasitic labels. Therefore, even when the parasites’ offspring emerge with their parasite-specific labels, and no colony-specific chemical labels, host workers will tolerate them. [7]
Paper wasps are vespid wasps and typically refers to members of the vespid subfamily Polistinae, though it often colloquially includes members of the subfamilies Vespinae and Stenogastrinae, discussed elsewhere, which also make nests out of paper. Paper wasp nests are characterized by open combs with down pointing cells. Some types of paper wasps are also sometimes called umbrella wasps, due to the distinctive design of their nests.
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.
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. 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.
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 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 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 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.
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.
The name cuckoo paper wasp refers to a monophyletic species group of brood-parasitic paper wasps in the genus Polistes. This species group contains only four species; Polistes atrimandibularis, P. austroccidentalis, P. maroccanus, and P. semenowi, all of them obligate social parasites of other Polistes species.
Nest usurpation is when the queen of one species of eusocial insects takes over the colony of another species.
Van der Vecht's gland or Van der Vecht's organ is a gland which is located in an area of modified cuticle on the rearmost gastral sternite of female wasps. This gland secretes chemicals which are important in the determination and maintenance of the hierarchy of groups of eusocial wasps and are used in the defence of the nests in others. In the Asian giant hornet the Van der Vecht's gland is used to scent mark hives of honey bees to attract other members of their colony to cooperatively attack the hive; the only known case of the gland's use to scent mark a food source. In the cleptoparasitic paper wasp Polistes semenowi the female usurps the host foundress, usually Polistes dominula and uses an enlarged Van der Vecht's gland to produce large quantities of hydrocarbons and to control the host workers, and even sometimes the host foundress. The gland was discovered by, and named in honour of, the Dutch entomologist Jacobus van der Vecht.