Australian paper wasp | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Hymenoptera |
Family: | Vespidae |
Subfamily: | Polistinae |
Genus: | Polistes |
Species: | P. humilis |
Binomial name | |
Polistes humilis | |
Subspecies | |
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Distribution of Polistis humilis |
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. [1] They have been known to re-utilize old nests. [2] While the species does not exhibit morphological class differences, [2] 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. [3]
Polistes humilis is in the genus Polistes , which consists of 150 species that can be found in all regions except the world's coldest climates. Polistes humilis was the only species of the tribe Polistini found in New Zealand after it was accidentally introduced from Australia, [4] prior to the introduction of the Asian paper wasp, Polistes chinensis in 1979. [5] There are 11 different Polistes species found in Australia, and the Polistes humilis is the most prevalent one. Historically, Polistes variabilis has often been misidentified as a sub-species of Polistes humilis. But, recent photogenic analysis has indicated that these are two separate species. [6] The species can be divided into two subspecies, P. humilis humilis which is found in northern Australia and P.humilis synoeus which can be found in the south. [2]
This species of wasp can be identified by its slender body structure and banded coloring. Adults have yellow faces and are 10–15mm long with a tannish-red coloration. Males can be distinguished from females by a yellow marking on the abdomen area. [7] In addition, Polistes humilis are generally bigger than most other paper wasps. [8] Nests of this species can be identified by their cone shaped appearance with multiple hexagonal cells. The species builds its nest out of grey wood fiber material, which is a mixture of its own saliva and wood. Unlike most species of wasps, Polistes humilis colonies have been known to re-utilize old nests from year to year. Some colonies overwinter and shelter above the comb in colder months. [2]
Polistes humilis is mostly found in Australia, particularly in southern Queensland, New South Wales, Victoria, and South Australia. [8] While this species is native to Australia, the species was accidentally introduced to New Zealand in the 1880s and has established a stable population in that country, largely confined to the North Island north of Tauranga and west of Te Kuiti. Environmentally, the highest abundance of P. humilis can be found in shrublands. However, the species can also be found in flax swamps and forest sites. [9]
The colony cycle of Polistes humilis starts in the spring and can begin in three ways. Firstly, the colony cycle can begin with the founding of a new nest, typically by several foundresses. [2] Secondly, the cycle can start with the re-utilization of an old nest, which can be several years old. Thirdly, the cycle can also begin with the continued use of an old nest as this species can overwinter. [2] In situations where old nests are used, new eggs are typically laid around the old cells.
Throughout the year, production of Polistes humilis nests typically follow a distinctive cycle. In the spring, nests are founded by females that have overwintered and have been previously inseminated. After founding, worker females are produced in the late spring and early summer. Males and queen females are produced in the late summer and early autumn. [2]
In Polistes humilis nests, there is an observable hierarchy between reproducing queens and sterile workers. There are no outward physical differences between females; even small females have been known to produce eggs. [10] Since there are no morphological differences between females, behavioral characteristics are the primary determinate of whether a female reproduces or not. These females can be determined by observation as they often receive solid food, honey, and water from worker females returning from foraging. Dominance is often displayed by a "tail wagging" behavior of the lower segment of the species. However, unlike most species that use behavioral characteristics to determine reproductive success, Polistes humilis does not exhibit high competition between females within the nest. [2] This is likely due to the high degree of relatedness among nestmates.
Polistes humilis queens are singly mated, meaning that the eggs of the queen are fertilized by a single male and only one fertilization event occurs per female. However, within each colony, there are an average of two queens. This multiple queen behavior likely developed as a defense mechanism for the nest. Since different males can fertilize each female in the nest, genetic diversity is increased within the nest, which can help defend against disease. In addition, a distinctive characteristic of this species is that there is no inbreeding within P. humilis colonies. This suggests that dispersal is primarily male based and that males do not reproduce in their native colony. Dispersal of males likely happens before mating. [7]
Polistes humilis are a haplodiploid species, with females developing from a fertilized egg and males developing from an unfertilized egg. In this system, queens are singly mated with full sisters sharing the paternal genetic information and either one of the maternal alleles. [7] Therefore, sisters that share the same mother are 75% related in accordance with the haplodiploidy hypothesis. However, this ratio is skewed, making siblings less related, by having multiple queens in a nest. [11] While this might benefit disease defense, skewing the relatedness ratio could disrupt the fragile eusocial behavior within the nest as some females within the nest are more related than others.
In Polistes humilis there are distinct benefits to having larger nest sizes. As nest size increases, the number of reproducing females directly increases. On one hand, this decreases the genetic relatedness of individuals and could cause a breakdown of the dominance hierarchy of the nest as it increases the gene pool of the nest. This correlation likely evolved as a defense against disease transmission, as increased genetic variability has been shown to reduce disease transmission within nests and could prevent a nest from dying out. [7] On the other hand, as nests get larger, there are increased numbers of individuals and therefore increased nest defense. For this reason, larger nests have better defense against predators and human intervention.
Since queen females have few morphological differences from worker females, they must use behavioral means to prevent them from laying eggs. As each queen selfishly wants to have as many offspring as possible, it is likely that oophagy occurs in Polistes humilis. [2] This practice limits the eggs that weaker females can produce and reduces the genetic variability of the nest. Dominance of females can also be established through behavioral displays. Dominant females display their dominance by horizontal vibrations of the gaster and by mounting and wrestling with other females. [2] These behaviors help the queens maximize their offspring's ability to survive and increases the genetic relatedness within the nest.
Polistes humilis are observed to subsist on food brought back to the nest by worker wasps. Once food is brought back to the nest, workers distribute the food to the queen and larvae. The species consumes primary water, pulp, carbohydrate and protein prey. In southern Australia, Polistes humilis appears to specialize in consuming lepidopteran larvae for protein as well as small spiders. Polistes humilis has to compete with other species for food, particularly the newly introduced, invasive species Vespula germanica . [12] While Polistes humilis is very common now, competition for food could reduce its population in the future.
As a stinging wasp, Polistes humilis has a very powerful defense mechanism. The venom in the sting of the species helps with prey capture and nest defense from predators. The venom comes from the two tubular glands and is secreted by powerful muscles that coat the reservoir and squeeze out the venom. The venom is known to be used in fights between species and is often used in limited amounts. The venom is largely made of serotonin, histamine, tyramine, and dopamine which are all considered major pain producing components of the venom. [13]
Polistes humilis also has developed two defenses against disease: genetic diversity and the production of cuticular antimicrobial compounds. There is a trade-off between these mechanisms, with larger colony sizes having more genetic diversity and smaller colonies having increased antimicrobials. This trade-off has likely developed by evolution as the cuticular compounds are metabolically expensive to produce so larger colonies may not be able to support this mechanism. In successful colonies, these mechanisms work together, as shown by the Pc80 locus. Genetic analysis shows that increased variation at this locus is positively correlated with antimicrobial strength. However, as the success of these mechanisms is very important for colony survival, these mechanisms are important limiting factors for maximum colony threshold size. Since antimicrobial compounds in Polistes humilis are found in the species' venom and only females produce the sting venom, larger colonies with more males might have lower effectiveness of this mechanism. [14] Group size and microbial defense are important characteristics to consider when determining disease susceptibility.
Polistes humilis is probably most known to humans by its painful sting. These stings often result from agitation of nest sites. Polistes humilis nests are often found in "modified habitats" where there is a mix of human structures and vegetation. Since nests are typically found in walls, eaves of buildings, and fences, humans are particularly at risk of accidentally disturbing a nest site. [10] However, while the sting is very painful, humans are not at risk of serious injury from the sting unless they are allergic. The best way to avoid a sting is to simply stay away from known nest sites if possible, as these wasps typically only sting as a defense mechanism.
Polistes humilis are important to the overall ecosystem as pollinators. Since this species is one of the most populous species in Australia, it is responsible for the survival of many native wildlife. Polistes humilis appear to be widespread pollinators, not only in abundance but also in the species of plants pollinated. While this behavior is good for native plants, it has important implications for non-native species. The success of an introduced invasive species largely depends on the availability of pollinator relationships. Since Polistes humilis are general pollinators, they have been observed to contribute to the success of invasive species, such as the milkweeds Asclepias curassavica and Gomphocarpus fruticosus. [3]
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.
Oophagy or ovophagy, literally "egg eating", is the practice of embryos feeding on eggs produced by the ovary while still inside the mother's uterus. The word oophagy is formed from the classical Greek ᾠόν and classical Greek φᾱγεῖν. In contrast, adelphophagy is the cannibalism of a multi-celled embryo.
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 instabilis, or unstable paper wasp is 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.
Dolichovespula arenaria, also known as the common aerial yellowjacket, sandhills hornet, and common yellow hornet, is a species of wasp within the genus Dolichovespula widely distributed in the North American continent.
Polybia occidentalis, commonly known as camoati, is a swarm-founding advanced eusocial wasp. Swarm-founding means that a swarm of these wasps find a nesting site and build the nest together. This species can be found in Central and South America. P. occidentalis preys on nectar, insects, and carbohydrate sources, while birds and ants prey on and parasitize them. P. occidentalis workers bite each other to communicate the time to start working.
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 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 revolutionalis, the stick-nest brown paper wasp, is a diurnal social wasp of the family Vespidae. They are known for the distinctive combs they make for their nests, and they have been found in Queensland, Australia in the areas of Brisbane and Townsville. They are an independent founding wasp species, and they build new nests each spring. They can be helpful because they control insect pests in gardens.
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.
Ropalidia romandi, also known as the yellow brown paper wasp or the yellow paper wasp. is a species of paper wasp found in Northern and Eastern Australia. R. romandi is a swarm-founding wasp, and manages perennial nests. Its nests are known as 'paper bag nests' and have different architectural structures, depending on the substrates from which they are built. The specific name honors Gustave, baron de Romand, a prominent French political figure and amateur entomologist.
Synoeca surinama is a Neotropical swarm-founding wasp of the tribe Epiponini. It is known for its metallic blue and black appearance and painful sting. S. surinama builds nests on tree trunks and can be found in tropical climates of South America. When preparing to swarm, there are a number of pre-swarming behaviors that members of S. surinama colonies partake in, such as buzzing runs and occasional brood cannibalism. In S. surinama, social environmental conditions determine the caste ranks of individuals in the developing brood. Unlike less primitive Hymenoptera species, S. surinama display little morphological variation between egg laying queens and workers. S. surinama wasps visit flowering plants and are considered pollinators. When these wasps sting, the stinger is left in the victim and the wasp ultimately dies.
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.