Polistes chinensis | |
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Polistes chinensis antennalis | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Hymenoptera |
Family: | Vespidae |
Subfamily: | Polistinae |
Tribe: | Polistini |
Genus: | Polistes |
Species: | P. chinensis |
Binomial name | |
Polistes chinensis (Fabricius, 1793) | |
Synonyms [1] | |
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, [2] having arrived in 1979. [3]
The wasps prey on invertebrates, especially caterpillars. [4] In this species, queens destroy up to 70% of worker-laid eggs and are aided by workers in a process known as worker policing. [5]
P. chinensis is a member of the cosmopolitan genus Polistes , the largest genus in the family Vespidae, with over 300 recognized species and subspecies. Two subspecies are known: [6]
The Asian paper wasp has a slender body about 13 to 25 mm (0.51 to 0.98 in) in length. Their bodies are reddish brown or black, with yellow rings and reddish areas on the abdomen. Their wings are reddish or amber brown, and they have long legs that hang down during flight. [7] The general morphology of the spermatheca is similar in both reproductive and non-reproductive individuals. The spermathecal gland attaches to the central region of the spermatheca. [8]
The Asian paper wasp frequently constructs nests on man-made structures including houses and buildings. It will also build nests on trees or bushes, usually on branches, but sometimes on leaves as well. P. chinensis prefers to colonize urban habitats, but nests have also been recorded near forest clearings. They sometimes build their nests in dense shrubs, making them difficult to locate. [7] Large populations usually develop in warm, lowland areas with open habitat such as shrublands, swamps and meadows.
The paper wasp is native to East Asia. It has been recorded in China, Japan, Korea and Mongolia. [9] The subspecies P. chinensis antennalis is invasive in New Zealand. [7] It is widespread in the North Island and the northern South Island and competes with native species for food, [10] It has since been recorded in Norfolk Island and New South Wales. [9] At least two independent invasion events, linked to Kanto and Kyushu, Japan, have been proposed as sources of the New Zealand wasps. [11] The Asian paper wasp was first recorded in New Zealand in 1979, in Kohimarama and the Whangaparaoa Peninsula. It remained confined to these areas of Auckland until 1984, when it began appearing in inner city areas and western suburbs. [12]
An overwintered female, who has been inseminated during the previous autumn, emerges during the spring and begins nest cell construction and egg laying. The first broods emerge in late spring or early summer and are entirely female. Males are produced from early summer onwards. Following male production, no more females are produced for the rest of the season. [7] This colony cycle can be divided into three stages: a solitary stage, a superindividual stage and a social stage. During the solitary stage the foundress builds the nest and cares for the first generation of brood. This stage ends when the first workers emerge. The superindividual stage ends when the first reproductive forms emerge. The social stage ends when the colony disintegrates. [13]
Food collection is very costly to the foundress of a colony and carries a high risk of mortality. However, it is a necessary activity for colony survival. Most collected proteinaceous material is consumed by larvae. A significant amount, fourteen percent, is allocated to oral secretions. Polistes chinensis, like many social wasps, uses a proteinaceous oral secretion to glue nest material to maintain and build the nest. The oral secretion of P. chinensis is 73% proteinaceous materials. [14] The queen will augment this investment in response to the environment. Oral secretions increase during rainy seasons, because the foundress must smear oral secretion frequently over the nest surfaces to avoid risk of destruction. Nests in exposed sites vs. sheltered sites show no significant difference in dry weight of secretions produced by the foundress. However, foundresses that nest in sheltered sites bring in more prey then those at exposed sites. Therefore, the relative amount of proteinaceous resources allocated to oral secretions is greater, at 22%. [15]
Foraging by a lone foundress has a high cost of predation risk for both herself and her brood. Furthermore, foraging success of a lone foundress has been estimated to be low. Rather than increase foraging time, P. chinensis will cannibalize their own larvae when there is not enough honey or prey available during the solitary stage of colony development. [16] The foundress targets larvae of different developmental stages depending on whether prey or honey is the limiting resource. Under scarce prey condition, foundresses cannibalize young larvae and intensively feed them to older larvae to ensure successful production of the first generation of workers. It is important that the queen successfully rear the first set of brood because she has a high risk of mortality while maintaining the nest on her own. In a honey limited environment, older larvae are more likely to be targeted. When larvae do not consume honey, they no longer produce saliva, which is an important nutrient resource for the adult wasp. When large larvae cannot produce enough saliva they are more likely to be cannibalized as a nutrient resource for the foundress herself. [16]
Foundresses need to leave the nest to gather prey, nest materials and water. When she does so there is a risk that conspecific females will depredate her brood. These females fly to a nest, pull out a single larva and return to their own nests to feed their own larva with it. Foundresses chase off attacking females by spiral flight, whereby two females fly in the air in a double helix pattern. Attendance on nests successfully deters attacking females. Foundresses use the loss of a larva as a cue to assess predation risk, and increase their defense efforts in response. When a larva is removed, lone foundresses increase the total time spent on the nest by decreasing time spent on each off nest activities, and increasing the time interval between consecutive activities. There is not a difference in the number of off nest activities. After 30 minutes following the removal of a larva, the time spent on the nest begins returning to the pre-removal level. After 60 minutes, time spent on the nest returns to pre- removal levels. [17]
Asian paper wasps are monandrous. Like other species of eusocial wasp, males are haploid and emerge from unfertilized eggs. Workers are .5 times genetically related to their sons, .375 times to their nephews, .25 times to their brothers, and .75 times to their sisters. According to kin selection theory, this relatedness predicts worker queen conflict. Workers should police the foundresses male eggs. However, this model does not seem to apply to P. chinensis. In fact, the opposite occurs, workers police each other. [18]
Workers police each others' eggs. 88.5% of queens' eggs survive to hatching vs 1.4% of workers'. Worker reproduction is frequent in queen-right colonies in which the queen is alive. Both queen and workers replaced workers' eggs. Polistes chinensis was studied alongside Polistes snelleni for comparison in this conflict. In P. snelleni, queens monopolize egg production. [19] Queens contributed 2.4 times more to replacing than workers. Workers sequentially perform oophagy and oviposition in the same cells. The ratio of worker-produced eggs to eggs laid by the queen is 3 or 4 times to one in a colony having between 100 and 500 wasps. [18]
Worker-worker policing is predicted to occur in polyandrous nests. A few hypotheses have been proposed to explain worker policing in the monandrous P. chinensis. The colony efficiency hypothesis proposes that worker policing is selected for because worker production imposes a cost on worker productivity. However worker oophagy is only observed in conjunction with oviposition. The colony efficiency theory predicts that even non reproductive workers would participate in policing. Another explanation is that worker policing occurs because of the size constraints of the colony. Workers collectively produce more eggs than the queen. The difference in egg production becomes more pronounced as the colony grows larger. Therefore, as nest size increases, competition for limited oviposition space becomes more intense, with workers destroying their nestmates eggs so they have room for their own. This theory has yet to be confirmed. [18]
Foundresses in P. chinensis control colony investment but allow worker oviposition. On average, around 40% of males in queen-right colonies are the sons of workers. The observed sex allocation ratio in field observations performed by the Natural History Museum and Institute in Japan was .61, while the expected ratio in queen-right colonies is .48. Variation in sex allocation ratio is likely explained by limited queen control. [20]
P. chinensis collects nectar and honeydew from flowers. The wasp preys on invertebrates, preferring larvae and caterpillars of lepidopteran insects (moths and butterflies) for protein resources, [7] but may also feed on marine invertebrates or fish carrion when the opportunity arises. [21] Asian paper wasps also feed on the larvae of other conspecific females. [17] Larval saliva is an important nutrient resource for adult Asian paper wasps. This contains a high concentration of free amino acids, 50 times that of floral nectars, and is nutritionally analogous to nectar. [16]
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 humilis, known as the Australian paper wasp, is a species of wasp in the family Vespidae that is found throughout Australia and which has been introduced to northern New Zealand. These paper wasps can be identified by their long thin legs and banded yellow and black coloring. They have been known to re-utilize old nests. While the species does not exhibit morphological class differences, there are distinct behavioral differences between queens and workers. In addition, the species is eusocial and benefits from relatedness between individuals. They are known for delivering a painful sting, especially when their nest is disturbed, a behavior that has been developed as a nest defense mechanism. While wasps are often viewed negatively, they play an important pollination role for many plants.
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 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.
Belonogaster juncea is a typical quasisocial paper wasp from sub-saharan Africa and south-western Asia. It is the type species for the genus Belonogaster.
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.
Belonogaster juncea juncea is a subspecies of Belonogaster juncea and is classified as a primitively eusocial wasp, meaning that the species is social while exhibiting a morphology that is indistinguishable from that of other castes. It is also classified as a type of African Paper Wasp. Many of the studies relating specifically to B. j. juncea take place at the University of Yaoundé in Cameroon.
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 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.
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.
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 japonicus is a eusocial paper wasp found in Japan. It was first described by Henri Louis Frédéric de Saussure in 1858. It is closely related to Polistes formosanus. This species lives in small colonies with few workers and a foundress queen. Nests of these wasps are sometimes used as a traditional medicine in Korea, China, and Japan.
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.
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.