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''Trigona corvina'' ( Cockerell, 1913) is a species of stingless bee that lives primarily in
Central Central is an adjective usually referring to being in the center of some place or (mathematical) object. Central may also refer to: Directions and generalised locations * Central Africa, a region in the centre of Africa continent, also known a ...
and South America. In
Panama Panama ( , ; es, link=no, Panamá ), officially the Republic of Panama ( es, República de Panamá), is a transcontinental country spanning the southern part of North America and the northern part of South America. It is bordered by Cost ...
, they are sometimes known as zagañas. They live in protective nests high in the trees, but they can be extremely aggressive and territorial over their resources. They use their
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavi ...
s to protect their food sources and to signal their location to nest mates. This black stingless bees of the tribe
Meliponini Stingless bees, sometimes called stingless honey bees or simply meliponines, are a large group of bees (about 550 described species), comprising the tribe Meliponini (or subtribe Meliponina according to other authors). They belong in the family ...
can be parasitic toward citrus trees but also helpful for crop pollination.


Taxonomy and phylogeny

''Trigona corvina'' belong to ''
Trigona ''Trigona'' is one of the largest genera of stingless bees, comprising about 32 species, exclusively occurring in the New World, and formerly including many more subgenera than the present assemblage; many of these former subgenera have been ele ...
'', the largest genus of stingless bees, with over 80 species. ''T. corvina'' was once classified as a variety of ''Melipona ruficrus'' based on worker appearance. Fossil records of the Meliponini tribe have been discovered and it is now understood that they differentiated from other related wasps in the Late Cretaceous period. The tribe is distinct with regards to their reduced wing venation and their reduced sting, which led to their development as stingless bees. It is possible that the differentiation of the Meliponini occurred in parallel with the dominance of flowering plants.


Description and identification


Identification

Workers of ''T. corvina'' are primarily black with smokey wings; they have a thorax width of 2.34 mm. The typical body length for a ''T. corvina'' is 6mm. They have a wide facial quadrangle and five-toothed mandibles covered in reddish brown hairs. They also have black, erect hairs on their clypeus. The virgin queens are differentiated from the workers by their shiny head, wider thorax and longer wings. They are a larger insect overall than workers. Males have a narrower facial quadrangle than both the workers and the queen. They are also differentiated by their
mandibles In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
, which have red stripes at their base and also by their legs which are not uniformly black. ''
Trigona fuscipennis ''Trigona fuscipennis'' is a stingless bee species that originates in Mexico but is also found in Central and South America. They are an advanced eusocial group of bees and play a key role as pollinators in wet rainforests. The species has many co ...
'' bees and ''Trigona corvina'' are often mistaken for one another since they are similar in appearance. ''Trigona fuscipenni''s workers are also completely black with one narrow red band just before the apex of the mandibles. But unlike the ''T. corvina'' species, they are smaller, have a slightly different mandible color and do not have erect black bristles.


Nest structure

The nests of ''T. corvina'' are dark grey, ovoid nests build around small branches on trees in South America. Examples of their nests have weighed 69 pounds and are 22 inches high and 17-18 inches in diameter. The brood area is always built first; it is strong and highly protective. 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 b ...
encasement consists of pollen from emerged brood cells and pollen excrement from young adult bees. Within the brood area, there are about 80,000 brood cells, all interconnected for easy passage. The queen cells are much larger and paler than the others. The next layers of the nest contain filled honey pots; these layers are also strongly supported with pollen excrement and wax. The outer layers of the nest are designed to be easily broken in case of attack or a quick exit. The layers consist of sheaths of hardened and brittle resin or cerumen (wax) supported by columns. ''T. corvina'' are known for their uncleanliness, as most of the nest scutellum is built up on top of the nest.


Thermal constraints

It has been demonstrated that body size and coloration impacts a bee’s ability to survive in certain temperatures. The larger and darker the bee, the better it does in colder environments. The color of the bee also influences where it tends to forage, either in sunlight or under the forest canopy. ''Trigona corvina'' is a midsized, black bee with a passive cooling rate of .32 °C/s. This is important because their foraging and nesting locations are often based on thermal constraints. For example, ''T. corvina'' refrain from foraging during the hottest period of the day. It also explains why they prefer foraging areas in shady areas.


Heterozygosity

''Trigona corvina'' have high
heterozygosity Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism. Mo ...
due to their environment. They typically live in regions with mosaic crops and plants that also have a low agricultural output. It has been demonstrated that drones have the highest diversity because they often come from many different colonies to mate with the Queen. It is important to maintain their heterozygosity for their continued proliferations and it is of concern because there has been a decline in diversity and abundance of insect pollinators.


Distribution and habitat

''T. corvina'' are native to Central and South America. They were originally discovered in Guatemala, Costa Rica and the Canal Zone of Panama. The species eventually spread to many more localities throughout Central and South America. They build their nests in trees from 8 feet to 40 feet into the air and they usually prefer areas with plentiful plant life. Since ''T. corvina'' are aggressive bees, their nests are regularly spaced to avoid unnecessary competition. It has been shown that their colonies have a density of 1.0 colonies/ha.


Colony cycle

A new colony is started when a virgin queen from one colony mates with a male of a neighboring colony. They will then create a new colony close to the nest of the virgin queen. New nests are created by the new queens but workers from the old nest must shuttle materials back and forth until the nest is complete. Additionally, many workers from the old nest must join the new queen in colonizing her nest until new generations of workers are born. Nests can exist for over 20 years, showing the extreme longevity of colonies. In a nest found in Panama, it was discovered that 91% of the bees in the nest were workers, 8% were males and <1% were virgin queens. Since nests are built around exposed branches, ''T. corvina'' nests are often damaged or knocked down in the absence of natural causes, indicating attack by large animals. This results in colony loss for the bees.


Behavior


Caste system

''Trigona corvina'' colonies are founded by a single virgin queen who rapidly mates with a single male and her
workers The workforce or labour force is a concept referring to the pool of human beings either in employment or in unemployment. It is generally used to describe those working for a single company or industry, but can also apply to a geographic regio ...
. Once the queen begins her colony, she grows in size and eventually loses the ability to fly. The queen lays all the eggs. The worker bees do not reproduce and they have a 3:1 sister to brother relatedness ratio as do all members of
Hymenoptera Hymenoptera is a large order of insects, comprising the sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are parasitic. Females typic ...
. Worker bees do the foraging, nest building, and raising of the brood. They are able to fly from colony to colony.


Communication


Pheromones

''T. corvina'' rely on
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavi ...
s for much of their communication with nest mates and rivals. They produce pheromones from their
labial glands The labial glands are minor salivary glands situated between the mucous membrane and the orbicularis oris around the orifice of the mouth. They are circular in form, and about the size of small peas; their ducts open by minute orifices upon the m ...
. The function of signaling depends on the profitability, but they commonly will scent mark a food source either for self-orientation, to deter rivals or to direct a nest mate to the resource. Once an individual finds a good food source, they will return to the same source for many days. If an individual detects the scent of a rival bee, they will avoid the plant in order to avoid conflict and to save time. It has also been shown that pheromones are a method of
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex (int ...
between male drones and queens. This form of communication differentiates
stingless bee Stingless bees, sometimes called stingless honey bees or simply meliponines, are a large group of bees (about 550 described species), comprising the tribe Meliponini (or subtribe Meliponina according to other authors). They belong in the family ...
s from
honey bee A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the current cosmop ...
s who use dances to indicate resource location.


Local Enhancement

''T. corvina'' have been shown to have local enhancement, which means that they prefer to land near other ''T. corvina'' individuals. They use both visual and epicuticular hydrocarbon cues to locate and monopolize food sources. ''Trigona corvina'' are able to recognize other members of their species through visual cues but they much prefer to use olfactory cues. This is because they can only see from 10–20 cm away but they are able to detect scent from 10–20 meters away. Color vision has emerged as an important characteristic for ''Trigona corvina'' since it enables them to identify other black bees from rival species.


Foraging

It has been shown that a few members of the colony act as trained foraging bees. These trained bees will go on pilot flights to search out new food sources. When the trained bee returns, they will be visually conspicuous about the food location by repeated hovering and landing behavior. They will also deposit pheromone clouds around the location of the food in case they are not present to show the way. The presence of the trained bee is not necessary for the colony to find the food source, but it is helpful overall. As more and more members of the colony arrive at the food source, the trained bee becomes even less important.


Interaction with other species


Diet

''T. corvina'' have a minimum
foraging Foraging is searching for wild food resources. It affects an animal's fitness because it plays an important role in an animal's ability to survive and reproduce. Foraging theory is a branch of behavioral ecology that studies the foraging behavi ...
range of 3.14 km and forage over 500 species of flowering plants, including over 100 tree species. Due to their large foraging range, they have access to exotic and forest-edge species. They are quick pollinators and they have an average plant visitation time of 25.6 seconds/ 3 flowers. This quick foraging combined with their wide range allows them to visit a huge number of plants during each foraging expedition. From sampling
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametophy ...
excrements, it is clear that '' Cavanillesia'' (Bombacaceae) is a primary food source for this species of bee. Due to their aggressive nature, ''T. corvina'' are very
territorial A territory is an area of land, sea, or space, particularly belonging or connected to a country, person, or animal. In international politics, a territory is usually either the total area from which a state may extract power resources or an ...
over their foraging areas and do not move without a fight. Their aggression increases when there are more bees in a foraging area.


Parasitism

''Trigona corvina'' have been known to act as parasites to citrus plantations. They typically collect the sticky
propolis Propolis or bee glue is a resinous mixture that honey bees produce by mixing saliva and beeswax with exudate gathered from tree buds, sap flows, or other botanical sources. It is used as a sealant for unwanted open spaces in the beehive. Pro ...
from the surfaces of young orange leaves and then they collect the liquid that emerges from the leaf margins after they destroy them with their mandibles. They are so effective in their destruction of the citrus plants that they are able to prevent its growth.


Defense

''T. corvina'' are highly aggressive bees. Although this provides them a competitive advantage over less aggressive bees, it also results in fights to the death. They have four levels of aggression for different situations. Level 1 is a low intensity threat that may occur when a rival bee approaches an occupied food source. The bee will spread its mandibles and tilt its head up so that its mandibles are pointing at the rival. They will then lift their abdomen and hold their wings at a wide angle; level 1 is only a display of aggression. Level 2 involves brief bodily contact and the goal is to knock the intruder off of the plant or to the ground if the fight occurs in the air. Occasionally, the bees will nip at each other’s legs, but injury is not intended. Level 3 fighting involves biting with the intent of injury. They pull on each other’s mandibles and legs, often interlocking their mandibles. It is common for more than two bees to be involved. Level 4 aggressions frequently results in a fight to the death. Bees involved will release an alarm pheromone, which will bring nest mates to their assistance. Sometimes fighting can result in battles between entire colonies, if colonies are all ''T. corvina'', it will usually result in a splitting of resources to avoid excessive deaths. There is a trade-off for ''Trigona corvina'' between effort spent defending their resource from intruders and foraging. They have to find an
Evolutionarily stable strategy An evolutionarily stable strategy (ESS) is a strategy (or set of strategies) that is ''impermeable'' when adopted by a population in adaptation to a specific environment, that is to say it cannot be displaced by an alternative strategy (or set of ...
(ESS) between the two important activities in order to gain the most benefit. It has evolved that ''Trigona corvina'' foragers do best with interactions with other aggressive bees or with bees who are easily scared off.


Competition

The
African honey bee The East African lowland honey bee (''Apis mellifera scutellata'') is a subspecies of the western honey bee. It is native to central, southern and eastern Africa, though at the southern extreme it is replaced by the Cape honey bee (''Apis mellif ...
is a common competitor of ''T. corvina''. When introduced to the same habitat, the ''T. corvina'' loses resources, since both forage in open spaces. Additionally, if a ''T. corvina'' nest is vacated, an African honey bee colony may move in. Honeybees can be very detrimental to ''T. corvina'' foraging because they are twice the size and are much more effective at collecting pollen. They also are able to forage more rapidly, so they have a much higher overall yield and their large size makes it difficult for the ''T. corvina'' to defend their territories.


Parasites

''T. corvina'' workers have been found with mites attached to the outer face of their hind tibiae. Males and queens are usually free of parasites.


Role in agriculture

''T. corvina'', as a species of stingless bees, are important crop
pollinators A pollinator is an animal that moves pollen from the male anther of a flower to the female carpel, stigma of a flower. This helps to bring about fertilization of the ovules in the flower by the male gametes from the pollen grains. Insects are ...
. Species native to Costa Rica are known to visit
chayote Chayote (''Sechium edule''), also known as mirliton and choko, is an edible plant belonging to the gourd family, Cucurbitaceae. This fruit was first cultivated in Mesoamerica between southern Mexico and Honduras, with the most genetic diversity ...
flowers, which become much more fruitful when visited by ''T. corvina''. They are also known to pollinate Panama hat plants (''
Carludovica palmata ''Carludovica palmata'' (Panama hat plant or toquilla palm) is a palm-like monocot plant. It is not a true palm. Its leaves are different from the leaves of true palms, and unlike true palms it does not develop a woody trunk. Its female flowers ...
''). In general, stingless bees are effective pollinators because they are less harmful to humans than
honeybees A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the current cosm ...
, and they are resistant to the common diseases and parasites of
honeybees A honey bee (also spelled honeybee) is a eusocial flying insect within the genus ''Apis'' of the bee clade, all native to Afro-Eurasia. After bees spread naturally throughout Africa and Eurasia, humans became responsible for the current cosm ...
.


References

{{Taxonbar, from=Q1913036
corvina Corvina is an Italian wine grape variety that is sometimes also referred to as Corvina Veronese or Cruina. The total global wine-growing area in 2010 was , all of which is grown in the Veneto region of northeast Italy, except for planted in A ...
Hymenoptera of North America Hymenoptera of South America Insects of Central America Insects described in 1913