Belonogaster Juncea Juncea
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''Belonogaster juncea juncea'' is a subspecies of ''
Belonogaster juncea ''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''. Breeding The nest is similar in form to that of the ''Polistes'' paper wasp's ne ...
'' 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 Cameroon (; french: Cameroun, ff, Kamerun), officially the Republic of Cameroon (french: République du Cameroun, links=no), is a country in west-central Africa. It is bordered by Nigeria to the west and north; Chad to the northeast; the C ...
.


Taxonomy and phylogeny

Johan Christian Fabricius Johan 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 cons ...
originally discovered and classified ''Belonogaster juncea juncea'' in 1781. ''Belonogaster juncea juncea'' belongs to the genus ''
Belonogaster ''Belonogaster'' (''belone'' - "needle", ''gaster'' - "belly") is a large genus of mainly Afrotropical quasisocial wasps, although some species occur in Arabia and two extend as far as India. They have characteristics of both the eusocial and th ...
'' which consists of over 80 species of primitively eusocial wasps. In general, the genus is
Afrotropical The Afrotropical realm is one of Earth's eight biogeographic realms. It includes Africa south of the Sahara Desert, the majority of the Arabian Peninsula, the island of Madagascar, southern Iran and extreme southwestern Pakistan, and the island ...
. The genus consists of independent-founding species, meaning that colonies are founded by either one or a few foundresses. Individuals in colonies of various ''Belonogaster'' species, such as '' B. grisea'', '' B. petiolata'', and ''B. j. juncea'' also tend to display aggressive behavior toward other members of the same colony. There are two subspecies of ''Belonogaster juncea'': ''Belonogaster juncea colonials'' and ''Belonogaster juncea juncea''.


Description and identification

''Belonogaster juncea juncea'' has a gaster that is considerably larger and more bulbous than its petiole, which appears very thin and long. Mesoscutum length is 3.8 mm ± 0.2 mm in dominant individuals and 3.6 mm ± 0.5 mm in subordinate individuals, while petiole length is 6.9 mm ± 0.4 mm in dominant individuals and 6.6 mm ± 0.5 mm in subordinate individuals. It also has somewhat lengthy antennae and typically exhibits a black/dark red color. Along with an increased size, a dominant female in a ''B. j. juncea'' colony can be identified by abdominal wiggling behavior, as subordinate females and other males or workers do not exhibit this sort of behavior.


Distribution and habitat

''Belonogaster juncea juncea'' has been sighted primarily in
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
and in various parts of
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
such as
Saudi Arabia Saudi Arabia, officially the Kingdom of Saudi Arabia (KSA), is a country in Western Asia. It covers the bulk of the Arabian Peninsula, and has a land area of about , making it the fifth-largest country in Asia, the second-largest in the A ...
and
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
. They also are found in both
tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
and
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
climates. ''B. j. juncea'' will colonize on buildings with great frequency, as evidenced by the fact that multiple studies have been conducted on colonies found in overhangs and roofs of the buildings of the University of Yaoundé in Cameroon, Africa. Some nests have also been found on large boulders. Their nests are made of paper, as they are a type of African Paper Wasp, and they contain a single comb. Typically, colonies consist of no more than 20 individuals.


Colony cycles

Colony foundation occurs throughout the year, independent of the seasons, and has an average cycle time of roughly seven months. Two phases make up the colony cycle: the pre-emergence phase and the post-emergence phase.


Pre-emergence phase

The pre-emergence phase is the time between the founding of the nest by one or multiple foundresses and the emergence of the first adult. During this phase, labor begins to become divided between dominant and subordinate individuals. For instance, dominant individuals will spend much of their time resting and building the cells in the colony, while subordinate individuals will spend time outside or on the edge of the nest. The pre-emergence phase is typically just over 71 days long and is divided into three portions: * Egg subphase - the time between the laying of eggs to the appearance of the first larva (usually takes around 26.5 days) * Larval subphase - the time between the appearance of the first larva to the appearance of the first pupae (usually takes around 21.8 ± 7.1 days) * Pupal subphase - the time between the appearance of the first pupae to the appearance of the first adult (usually takes around 22.8 ± 8.7 days)


Post-emergence phase

The post-emergence phase is the time between the emergence of the first adult and the abandonment of the colony, and reproductive episodes usually occur during this phase as well. Females typically emerge first, meaning that the initiation of the post-emergence phase coincides with the appearance of the first female. Males begin to emerge after females, but any female that happens to emerge with these males are considered to be worker females. Between 77 and 196 days after the foundation of the colony, the initial dominant female will abandon the colony. This is followed by an increase in fighting behavior as other females will fight with each other to be the most dominant individual. The losing females will either remain in the colony as subordinates or leave the nest and start a new colony; either way, in many cases, the original nest is completely abandoned by the end of seven months.


Behavior

Studies have shown that ''B. j. juncea'' exhibit thirty-nine different types of behavior. These different behaviors fall into five different categories, with some overlap, and these categories are: foraging, building, feeding, inactivity, and reproduction. Some of the behaviors that fall into these five categories include: * Foraging: Behaviors include absence from the nest, landing on the nest with food such as liquid material or prey, and giving the prey to those who need it, such as the larvae * Building: Behaviors include landing with and/or malaxating pulp, enlargement of the cells in which the wasps live, reinforcement of the pedicel (an antenna segment), or rubbing the pedicel with abdominal sternites * Feeding: Behaviors include the reception or malaxation of prey, feeding the larvae, and obtaining larval secretions * Inactivity: Behaviors include keeping watch over the colony, resting, and self-cleaning * Reproduction: Behaviors include nest inspection for reproduction, cell initiation, and laying the eggs The frequency with which these behaviors occur and the way in which the behaviors are carried out also differs between colonies that are in the pre-emergence phase and those that are in the post-emergence phase.


Dominance and subordination acts

''B. j. juncea'' individuals will also exhibit acts of behavior towards other individuals, and these acts are classified as either dominance acts or subordination acts. Dominance acts include: * Falling flights: two adults will fight to the point where they will fall out of the nest, and one may die as a result *
Grappling Grappling, in hand-to-hand combat, describes sports that consist of gripping or seizing the opponent. Grappling is used at close range to gain a physical advantage over an opponent, either by imposing a position or causing injury. Grappling i ...
: two wasps climbing on top of each other and wrestling; a winner is decided when an individual puts its gaster on the thorax of the other (the loser). The winner is the dominant individual. This is the most prevalent dominance act and plays the largest role in defining the dominance structure. * Biting another nest-mate: one insect bites another on an areas including the head, thorax, wings, or abdomen. * Chasing: occurs when an individual raises its wings and quickly moves toward another * Antennal palpation: an individual touches a portion of another individual with its antennas Dominant females will also be the only ones laying eggs. Individuals who ultimately succumb to the dominance of another will then exhibit acts of subordination, which include: * Submission: giving in to behaviors such as grappling * Akinesis: a body posture indicated by both the antennae and body lying flat on the nest, usually occurs after submission * Avoidance: the individual who submitted will attempt to avoid the one that dominated it


Dominance hierarchies

Dominance exists in a
hierarchy 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 ...
within colonies of ''Belonogaster juncea juncea'', and the "dominance rank" of a particular individual may largely determine which of the five behavioral categories comprises the majority of an individual's behavior. For example, the most dominant individual will spend most of its time on reproductive behavior and inactive nest behavior. The amount of time that individuals will spend on building the nest and especially foraging activity will increase as one moves down the hierarchy. Because of this, it is possible to deduce the rank of an individual based on the amount of time spent in the nest versus foraging for food. Dominant females, along with being the largest individual in the nest, also exhibit the most well-developed
ovaries The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. T ...
, which may indicate that they are the ones who reproduce. Subordinate individuals have thread-like ovaries, with oocytes that are not fully developed.


Hierarchy structure and stability

The results of the Appleby Test, which measures the "dominance index" of a particular individual, determine that the hierarchy in ''B. j. juncea'' is linear, meaning that dominant individuals are distinct from subordinates of lower and lower ranks. It has also been noted that most of the time, an individual will more frequently perform dominance acts on individuals of a rank that is immediately below theirs versus individuals that are several ranks below theirs, but occasionally, the most dominant individual (i.e. the α individual) will perform dominance acts on other individuals that are many ranks below them. The hierarchy is also stable. A dominant female usually emerges in a colony within two days of its founding, and it typically remains dominant until it abandons the colony or dies. When this occurs, dominance acts occur much more frequently within the colony as the replacement female vies for dominance. Falling fights are most frequently observed during this time. Also, because falling fights do not regularly occur during queen replacement in other species of ''Belonogaster'', such as ''B. grisea'' or ''B. petiolata'', ''B. j. juncea'' dominance hierarchy establishment is considered to be more severe when compared to other members of the genus.


Kin selection


Immature brood recognition

''Belonogaster juncea juncea'' have the ability to recognize their own immature brood and differentiate it from those of other, alien females. They most likely are able to do this through the use of odors and scents that are unique and characteristic to their own brood. Typically, only one female in the nest is responsible for this brood recognition.


Foundress size and colony success

Colonies of ''Belonogaster juncea juncea'' can be haplometrotic or pleometrotic, meaning that they are founded by one or several (usually two to eight) foundresses, respectively. Generally, pleometrotic colonies are more successful than haplometrotic colonies; pleometrotic colonies display increased total colony productivity and increased probability that an adult is produced when compared to haplometrotic colonies. Pleometrotic colonies are favored over haplometric colonies because ecological constraints on ''B. j. juncea'' are quite high, so the cost of a female attempting to found her own colony are typically greater than the costs of either joining an existing colony as a subordinate or even usurping an existing colony.


Co-foundresses

Co-foundresses are types of foundress females that, while they do not take part in initiating the nest, join a colony upon its initiation by the initial foundress. These females that were associated with each other came from the same nest approximately 86.7% of the time, which suggests that co-foundress relatedness is high. There is also a directly proportional relationship between colony survivability and number of foundresses, as it is much more common for pleometrotic colonies with co-foundresses to reach the stage where reproduction occurs than it is for haplometrotic colonies.


Nest usurpation

Rarely, a foreign female ''B. j. juncea'' will invade a colony and take the place of its dominant female. The invader will then eat the eggs already present in the nest and will destroy many of the existing cells. This act of usurpation has been observed primarily in the pre-emergence stages of colonies, specifically during the egg sub-phase and the pupal sub-phase. Usurpation most likely occurs when the costs to a female of founding a new nest are greater than the costs of usurping another, already existing nest. This process has only been effective on pleometrotic colonies in which at least one of the original foundresses remains in the colony after it is usurped, and this is most likely because if a usurper were to invade a haplometrotic colony, the original foundress could quickly abandon the nest, leaving it very prone to failure. It is hypothesized that this behavior is not very common due to the high degree of relatedness among foundresses and because the presence of multiple foundresses strengthens communal defense mechanisms that would keep the usurper out.


Serial polygyny

Occasionally, instead of completely abandoning the nest after seven months, a female may remain in the same nest and start another colony cycle. This female effectively replaces the previously dominant female as the most dominant individual in the nest, since the dominant female left before the original cycle ended. This succession of queens in the same nest is known as serial
polygyny Polygyny (; from Neoclassical Greek πολυγυνία (); ) is the most common and accepted form of polygamy around the world, entailing the marriage of a man with several women. Incidence Polygyny is more widespread in Africa than in any o ...
. This is not a very common occurrence when compared to complete nest abandonment, and in the periods between queens, egg and larvae quantity decreased as a result of no care being provided to them.


Interactions with other species


Diet

''Belonogaster juncea juncea'' will typically feed and provide food to their larvae in the forms of either liquid matter or prey. Liquid matter primarily consists of honeydew 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 ...
from various species of plants. In order to obtain this liquid manner, they exhibit a relationship with Aleyrodidae (whiteflies) and can parasitize the various trees or leaves in order to extract fluids. Their prey includes various species of insects such as caterpillars, winged ants, and
grasshopper Grasshoppers are a group of insects belonging to the suborder Caelifera. They are among what is possibly the most ancient living group of chewing herbivorous insects, dating back to the early Triassic around 250 million years ago. Grasshopp ...
s.


Predators and parasitism

Various species of ants,
praying mantises Mantises are an order (Mantodea) of insects that contains over 2,400 species in about 460 genera in 33 families. The largest family is the Mantidae ("mantids"). Mantises are distributed worldwide in temperate and tropical habitats. They ha ...
,
spiders Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species dive ...
, and sphecids have been known to attack nests of ''B. j. juncea''. However, when foundresses choose a nesting site, they typically choose places where predators cannot easily reach them, such as sides of buildings and stone rocks. Because of this, attacks on their nests are rare. ''B. j. juncea'' is also parasitized by ''Anacamtomyia'', a type of tachinid parasite. This parasitism and infestation of ''B. j. juncea'' nests often plays a role in nest abandonment.


Human importance


Traditional medicine

In certain African regions, especially Nigeria, ''B. j. juncea'' and other wasps of the genus ''Belonogaster'' are used in traditional forms of medicine. They are said to heal sicknesses in children if the whole wasp is cooked with the roots of plants and then later consumed. They are also occasionally used for various ceremonial purposes.O. A. Lawal and A. D. Banjo, 2007. Survey for the Usage of Arthropods in Traditional Medicine in Southwestern Nigeria. Journal of Entomology, 4: 104–112.


References


External links


Jean-Luc Renneson's studies of Belonogaster juncea
{{Taxonbar, from=Q18162797 Vespidae Arthropods of the Middle East