Ropalidia Revolutionalis
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''Ropalidia revolutionalis'', the stick-nest brown paper wasp, is a diurnal social wasp of the family
Vespidae The Vespidae are a large (nearly 5000 species), diverse, cosmopolitan family of wasps, including nearly all the known eusocial wasps (such as ''Polistes fuscatus'', ''Vespa orientalis'', and ''Vespula germanica'') and many solitary wasps. Each ...
. They are known for the distinctive combs they make for their nests, and they are mostly found in
Queensland, Australia ) , nickname = Sunshine State , image_map = Queensland in Australia.svg , map_caption = Location of Queensland in Australia , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , established_ ...
in the areas of
Brisbane Brisbane ( ) is the capital and most populous city of the states and territories of Australia, Australian state of Queensland, and the list of cities in Australia by population, third-most populous city in Australia and Oceania, with a populati ...
and
Townsville Townsville is a city on the north-eastern coast of Queensland, Australia. With a population of 180,820 as of June 2018, it is the largest settlement in North Queensland; it is unofficially considered its capital. Estimated resident population, 3 ...
. 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.


Taxonomy and phylogenetics

''Ropalidia revolutionalis'' were first classified by
Henri Louis Frédéric de Saussure Henri Louis Frédéric de Saussure (; ; 27 November 1829 – 20 February 1905) was a Swiss mineralogist and entomologist specialising in studies of Hymenoptera and Orthopteroid insects. He also was a prolific taxonomist. Biography Sauss ...
in 1853. They are part of the ''Polistine'' tribe ''
Ropalidiini Ropalidiini is a tribe of social wasps inhabiting the Afrotropical, Indomalayan and Australasian biogeographical regions. Genera and selected species *'' Belonogaster'' **'' Belonogaster juncea'' ( Fabricius, 1781) **''Belonogaster petiolata' ...
'', which includes paper wasps in the
Australasia Australasia is a region that comprises Australia, New Zealand and some neighbouring islands in the Pacific Ocean. The term is used in a number of different contexts, including geopolitically, physiogeographically, philologically, and ecologica ...
n region, specifically in the
Lesser Sunda Islands The Lesser Sunda Islands or nowadays known as Nusa Tenggara Islands ( id, Kepulauan Nusa Tenggara, formerly ) are an archipelago in Maritime Southeast Asia, north of Australia. Together with the Greater Sunda Islands to the west they make up t ...
and the northern regions of
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
. They are also part of the genus ''Ropalidia,'' which is the only genus that contains both swarm-founding and independent founding wasp species.


Description and identification

''R. revolutionalis'' are small, dark, reddish brown wasps with body lengths of 8 to 10 mm. When viewed closely, they have remarkable patterns on their bodies, and they have eyes with multiple
lenses A lens is a transmissive optical device which focuses or disperses a light beam by means of refraction. A simple lens consists of a single piece of transparent material, while a compound lens consists of several simple lenses (''elements''), ...
. They are slightly smaller than species from the family of ''
Polistes Wasps of the cosmopolitan genus ''Polistes'' (the only genus in the tribe Polistini) 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 ...
''. This is because their
abdomens The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the torso. ...
have a segment after the waist that is slimmer than the following segment. Although these wasps are not considered especially aggressive, they will sting if their nests are disturbed.


Distribution and habitat

The nests of ''R. revolutionalis'' consist of several combs each about 100 mm long with two columns of cells that resemble sticks hanging from a branch. Their nests are usually found on shrubs, hanging baskets, under
eaves The eaves are the edges of the roof which overhang the face of a wall and, normally, project beyond the side of a building. The eaves form an overhang to throw water clear of the walls and may be highly decorated as part of an architectural styl ...
, or near other sheltered areas. These wasps are found mainly in south-eastern to northern Queensland, Australia.


Colony cycle

''Ropalidia revolutionalis'' is an independent-founding wasp species that begins building nests each spring. Each nest is founded by a small association of females that begin by creating a single comb. During this pre-emergence stage, the number of egg-laying females decreases to only one being present in the nest (haplometrosis). However, after the emergence of the first progeny of adults, the females create smaller combs alongside the largest comb, and more potential egg-layers join the colony as the construction progresses (pleometrosis). The progeny join the adults in taking care of the nests. Near the end of summer, the queen produces both male wasps and
fertile Fertility is the capability to produce offspring through reproduction following the onset of sexual maturity. The fertility rate is the average number of children born by a female during her lifetime and is quantified demographically. Fertilit ...
female wasps. These progeny then mate with wasps from other colonies, and then the females find secure locations in which they can reside during the winter. At this point, the colonies begin to deteriorate, and the females start their own colonies the following spring.


Division of labor


Nest building

Each colony begins as a single comb constructed by a few females or by one female, and eventually the colony gains smaller
satellite A satellite or artificial satellite is an object intentionally placed into orbit in outer space. Except for passive satellites, most satellites have an electricity generation system for equipment on board, such as solar panels or radioisotope ...
combs as the year progresses. Workers create these combs by using
pulp Pulp may refer to: * Pulp (fruit), the inner flesh of fruit Engineering * Dissolving pulp, highly purified cellulose used in fibre and film manufacture * Pulp (paper), the fibrous material used to make paper * Molded pulp, a packaging material * ...
and decaying wood which they chew and mix with their saliva. Each comb is made up of two rows of cells attached by a pedicle (stalk) at the upper end. Also, occasionally females reconstruct old nests that partially deteriorated during the winter, which suggests that some colonies last more than one year.


Caring for larvae

Each
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
of the ''R. revolutionalis'' is kept in an individual cell in the nest. Each of these cells has a transparent bottom where the workers remove
feces Feces ( or faeces), known colloquially and in slang as poo and poop, are the solid or semi-solid remains of food that was not digested in the small intestine, and has been broken down by bacteria in the large intestine. Feces contain a relati ...
from the cell. Foragers return to the nest with
caterpillars Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Symph ...
and other soft bodied insects. Before presenting food to the larvae, they drum their heads on the inside of the cell. Females rear the young into August even as the nest deteriorates, and the larvae eventually pupate and become adult wasps.


Worker activity

During the day, foragers search for flowers and small prey. They bring back soft insects for the larvae and
nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
for the adults. When temperatures are high, workers will often raise their
abdomens The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the torso. ...
, draw their legs in, and fan their wings rapidly in an effort to cool down the nest. They also continually construct and take care of the cells in the comb.


Communication


Feeding

Females exchange food either by facing each other and engaging in
trophallaxis Trophallaxis () is the transfer of food or other fluids among members of a community through mouth-to-mouth ( stomodeal) or anus-to-mouth ( proctodeal) feeding. Along with nutrients, trophallaxis can involve the transfer of molecules such as pher ...
(the transfer of liquid food by putting the mandibles in opposition) or in some cases by mounting another female and bending over her thorax to transfer food. The reproductive females solicit the most frequently.


Alarm

When other insects approach the nest, the dominant individuals (functional queens) may vibrate their wings or assume a stance of alarm to warn the other females of approaching danger. Females also demonstrate alarm through agitated movement. This is usually demonstrated by contracting and expanding their abdomens or by holding their antennae straight out from their bodies.


Dominance hierarchy

Each nest usually contains several larger, dominant (queen-like but not reproducing) females. These females occasionally engage in aggressive acts such as darting around for short intervals or climbing onto the backs of
subordinates A hierarchy (from Greek: , from , 'president of sacred rites') is an arrangement of items (objects, names, values, categories, etc.) that are represented as being "above", "below", or "at the same level as" one another. Hierarchy is an important ...
and chewing or biting. They also tend to occupy the largest comb, while subordinates (those that lack developed eggs) are often forced onto smaller combs or reside on the substrate of the ceiling. Acts of dominance also appear to increase as the colony cycle progresses, so it is possible that some acts of dominance may be performed by the parent-generation on the newly emerging adults.


Reproduction


Mating

''Ropalidia revolutionalis'' females almost always mate singly, increasing the amount of
relatedness The coefficient of relationship is a measure of the degree of consanguinity (or biological relationship) between two individuals. The term coefficient of relationship was defined by Sewall Wright in 1922, and was derived from his definition of th ...
within the colony. Also, there is little to no
inbreeding Inbreeding is the production of offspring from the mating or breeding of individuals or organisms that are closely related genetically. By analogy, the term is used in human reproduction, but more commonly refers to the genetic disorders and o ...
within the colonies. This is because males leave the nest and mate with females from other colonies, and newly fertile females also mate with non-related males before hiding during the winter.


Reproductive dominance

Sometimes eggs laid by subordinate females are eaten by dominant females, but multiple females can still lay eggs after the colony has reached the post-emergence stage. Also, there is usually one primary dominant reproducer, and that producer stays in the nest and sits on capped cells containing
pupae A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
.


Queen cycling

The dominant reproductive female remains dominant until she dies. After this occurs, the colony goes through a short period of having multiple
queens Queens is a borough of New York City, coextensive with Queens County, in the U.S. state of New York. Located on Long Island, it is the largest New York City borough by area. It is bordered by the borough of Brooklyn at the western tip of Long ...
(often daughters of the previous queen). Eventually, one queen becomes dominant again, but the nest still contains a few less dominant queens.


Spatial arrangement

The individuals in each
colony In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the ''metropole, metropolit ...
can usually be separated into three groups: on the nest, next to the nest, or absent from the nest. Those who are on the nest usually consist of the dominant reproductive female, queens, and those caring for the
brood Brood may refer to: Nature * Brood, a collective term for offspring * Brooding, the incubation of bird eggs by their parents * Bee brood, the young of a beehive * Individual broods of North American Periodical Cicadas: ** Brood X, the largest bro ...
or taking care of the nest. If more than one reproducing female is present in a nest, the less dominant one either stays on a smaller comb or remains next to the nest out of the way of the dominant individual. Non-reproductive individuals can reside in all three groups, but they usually remain either next to the nest or absent from the nest foraging for food. If they are on the nest, the dominants often force these subordinates onto smaller combs.


Defense

If a member of either sex of ''R. revolutionalis'' feels threatened, the wasp assumes a defensive position on the nest. It will warn the intruder to retreat by first lifting and spreading its wings, and then waving its front legs and sometimes shaking its body. If the threat does not go away, the wasp will then fly at and often
sting Sting may refer to: * Stinger or sting, a structure of an animal to inject venom, or the injury produced by a stinger * Irritating hairs or prickles of a stinging plant, or the plant itself Fictional characters and entities * Sting (Middle-eart ...
the intruder.


Kin selection

Since ''R. revolutionalis'' queens mate singly, there is a high level of relatedness within the colony, and the workers are more likely to favor the sons produced by workers rather than sons produced by the queen because the workers are more closely related to each other's sons. This can create potential conflict. However, when there are multiple queens in a colony, the level of relatedness decreases, and the workers are more likely to favor the queen's sons. So, the workers switch between wanting to invest in new queens and wanting to invest in males depending on how many queens already exist in the colony.


Human-wasp interaction

''Ropalidia revolutionalis'' help control
pests PESTS was an anonymous American activist group formed in 1986 to critique racism, tokenism, and exclusion in the art world. PESTS produced newsletters, posters, and other print material highlighting examples of discrimination in gallery represent ...
in gardens by catching caterpillars and other insects for their larvae. If possible, their nests are best left alone. However, if disturbed, they will sting. Their venom contains 5-hydroxytryptamine, which is thought to be a pain-producing compound, though the pharmacological mode of action remains unknown.


References

{{Taxonbar, from=Q14425369 Vespidae Fauna of Queensland Insects described in 1853