Synoeca Cyanea
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''Synoeca cyanea'', commonly known as the marimbondo-tatu in Brazil, is a swarm-founding
eusocial Eusociality ( Greek 'good' and social) is the highest level of organization of sociality. It is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generations wit ...
wasp. Native to
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
and
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
, ''S. cyanea'' is one of the largest and most aggressive species of social wasps and is feared in many rural areas. It begins its colony cycle in the early spring and continues until nest abandonment. Throughout its life, ''S. cyanea'' forage sugary substances and animal carcasses for food and wood pulp for its nest. ''S. cyanea'' is also known for its strong venom, which is enough to cause
haemolytic Hemolysis or haemolysis (), also known by several other names, is the rupturing (lysis) of red blood cells (erythrocytes) and the release of their contents (cytoplasm) into surrounding fluid (e.g. blood plasma). Hemolysis may occur in vivo ...
activity.


Taxonomy and phylogenetics

Johan Christian Fabricius Johann Christian Fabricius (7 January 1745 – 3 March 1808) was a Danish zoologist, specialising in "Insecta", which at that time included all arthropods: insects, arachnids, crustaceans and others. He was a student of Carl Linnaeus, and is co ...
, a Danish zoologist, classified ''S. cyanea'' in 1775. The genus '' Synoeca'' is a part of the paper wasp tribe
Epiponini The Epiponini (formerly known as Polybiini) are a large and diverse tribe of social wasps inhabiting the Neotropical region, with some species' ranges extending into the Nearctic The Nearctic realm is one of the eight biogeographic realms ...
and consists of five species (''S. chalibea'', ''S. virginea'', '' S. septentrionalis'', '' S. surinama'', and ''S. cyanea''). ''Synoeca'' is a basal genus in the tribe. Cyanea, the adjective describing the species, translates to “dark blue,” referring to the metallic blue patches on the exterior of ''S. cyanea''.


Description and identification

''S. cyanea'' is black with metallic blue patches on its body. In addition, ''S. cyanea'' has a red clypeus. While other species in the genus ''Synoeca'' do not demonstrate
caste A caste is a Essentialism, fixed social group into which an individual is born within a particular system of social stratification: a caste system. Within such a system, individuals are expected to marry exclusively within the same caste (en ...
differences, ''S. cyanea'' sometimes shows differentiation between the size of workers and the queen. A queen of a nest that is early in development will often be the same size as the workers and intermediates as it has not fully developed its reproductive organs. As the colony and nest grow, the queen also grows in size, due to ovarian development, until it is larger than the workers and intermediates. When males are present, they are also smaller than the queen.


Nesting

''S. cyanea'' composes its nest out of a single comb with the cells directly attached to a tree. These types of nests are known as astelocyttarous nests. A corrugated, protective envelope made of wood pulp covers the nest, with the entrance on the superior portion of the nest. It is suggested that the architecture of the nest serves as protection against the predation of ants.


Distribution and habitat

While the genus ''Synoeca'' is found from Mexico to Argentina, ''S. cyanea'' is native to almost the whole country of Brazil. Typically, nests are found in the
Restinga Restingas () are a distinct type of coastal tropical and subtropical moist broadleaf forest in eastern Brazil. They form on sandy, acidic, and nutrient-poor soils, and are characterized by medium-sized trees and shrubs adapted to the drier and n ...
and the Atlantic Rain Forest. However, ''S. cyanea'' has been known to forage and make nests in
mangrove A mangrove is a shrub or tree that grows mainly in coastal saline water, saline or brackish water. Mangroves grow in an equatorial climate, typically along coastlines and tidal rivers. They have particular adaptations to take in extra oxygen a ...
areas.


Colony cycle

''S. cyanea'' starts a new colony cycle during the spring as it gets warmer. The first stage of a new colony is known as the pre-emergence state, in which workers have not been produced. No adult offspring are present and the larvae are still very young. A pre-emergent colony can have one or more queens. Following the pre-emergent state, the colony grows to the emergent stage. Workers are present along with larvae of different ages and multiple adult generations. In nests with fewer queens, intermediate females can be found. ''S. cyanea'' males are present in the colony during the male-producing stage. Upon nest abandonment during the late summer, the colony shifts to the quiescent stage, i.e. the nest site no longer houses wasps.


Behavior


Dominance hierarchy

The hierarchy of ''S. cyanea'' is based on physiology, specifically ovarian development. Wasps containing undeveloped filamentous
ovariole An ovariole is a tubular component of the insect ovary, and the basic unit of egg production. Each ovariole is composed of a germarium (the germline Stem-cell niche, stem cell niche) at the anterior tip, a set of developing Oocyte, oocytes containe ...
s, slightly developed oocytes, or some mature oocytes are considered workers and intermediates. Wasps with well-developed and longer ovarioles with mature oocytes become queens. The number of queens in the colony determines the development of a queen’s ovaries. If there are more queens present, the ovaries are at a lower degree of development. One benefit to this is that as a swarming insect, queens with smaller abdomens are able to fly more easily than queens with larger abdomens. This makes the queen less susceptible to predators.


Foraging

Foraging activity for the swarm-founding wasp ''S. cyanea'' begins early in the morning and stops in the evening. During the hottest hours of the day, activity of the wasps is highest. In addition to temperature, ''S. cyanea'' increase their activity with decreased levels of humidity and heightened intensities of light. While foraging, the wasps collect water for temperature control, wood pulp for building and repairing the nest, sugary substances for both the larvae and adults and finally animal protein for larvae.


Nest repair

Following damage by rain to the nest, ''S. cyanea'' will seal a leak of water with their mouth-pieces. Individuals begin by suctioning the water from the wall of the nest and then finish by expelling the water to the exterior of the colony. The hole is then repaired using wood pulp.


Kin selection


Worker-queen conflict

In ''S. cyanea'', discrimination between the queen and workers is often not determined by morphology, but by physiology. Young females in the colony can develop ovaries at different points in the cycle, so castes must be determined by the adult wasps rather than at the larval stage. Often, adult wasps will manipulate young females and suppress their ovaries because a wasp becomes a queen once she has mated. The number of queens allowed in a nest often fluctuates. When queen repopulation is not needed, worker policing helps to prevent female egg-layers through egg-eating and aggressive behaviors.


Interaction with other species


Diet

The wasps forage for sugary substances to feed to both larvae and adult wasps. Additionally, ''S. cyanea'' is among the species of Neotropical wasps that practices
necrophagy Scavengers are animals that consume dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a herbivorous feeding beh ...
. ''S. cyanea'' feed larvae the animal proteins collected. The wasps are predators of the coffee-leaf-miner, ''
Leucoptera coffeella ''Leucoptera coffeella'' (coffee leaf miner) is a moth in the family Lyonetiidae. It is found in every coffee-growing country in South America, Central America and the West Indies. It is considered one of the worst pest species of coffee. The w ...
''.


Defense

Characteristic to the genus, ''S. cyanea'' will sound an alarm when disturbed. This sound is made when the wasps scrape their mandibles on the carton of their nest. In addition, as one of the most aggressive social wasps, ''S. cyanea'' often will bite and sting when threatened. Their barbed sting often remains in the skin as the wasp pulls away.


Human importance


Venom

''S. cyanea'' venom is strong enough to cause
haemolytic Hemolysis or haemolysis (), also known by several other names, is the rupturing (lysis) of red blood cells (erythrocytes) and the release of their contents (cytoplasm) into surrounding fluid (e.g. blood plasma). Hemolysis may occur in vivo ...
activity.
Rhabdomyolysis Rhabdomyolysis (shortened as rhabdo) is a condition in which damaged skeletal muscle breaks down rapidly. Symptoms may include muscle pains, weakness, vomiting, and confusion. There may be tea-colored urine or an irregular heartbeat. Some o ...
and hemorrhage may also occur. In mice, abdominal spasms, ataxia, defecation, dyspnoea, hyperactivity, hypoactivity, sweating, and throes were observed following venom injection. ''S. cyanea'' venom also contains some antibacterial activity. Human accidents with wasps can occur one of two ways. Either a human may receive no more than one or two stings, or a swarm may attack a human. Symptoms following the attack can range from inflammatory reactions to severe allergic reactions resulting in anaphylactic shock. In some cases, death may occur following several stings and a large amount of venom injection; however, a wasp sting typically will not produce a reaction more severe than local symptoms that affect only the skin.


Biocontrol agent

In Brazil, the fruit fly ''
Zaprionus indianus ''Zaprionus indianus'', the African fig fly, is a species of vinegar fly in the family Drosophilidae. The species has become widely distributed in tropical regions of the world, as well as southern Europe and the U.S. Although the holotype was ...
'' is a common pest species that inflicts damage to multiple types of fruit trees. Since ''S. cyanea ''may obtain many nutrients from these fruit trees, researchers have observed the predatory nature of these wasps on ''Z. indianus''. When ''S. cyanea'' detect the fruit fly larvae, they soften, remove, and transport them back to the colony. For this reason, many fruit farmers consider ''S. cyanea'' a prospective candidate for use in pest management.


References

{{Taxonbar, from=Q10375500 Vespidae Insects of South America Insects described in 1775 Taxa named by Johan Christian Fabricius