Nannotrigona Testaceicornis
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''Nannotrigona testaceicornis'' is a
eusocial Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generat ...
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 A ...
species of the order
Hymenoptera Hymenoptera is a large order (biology), order of insects, comprising the sawfly, 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 Par ...
and the genus ''Nannotrigona''. Its local common name is ''abelhas iraí''. This species has a large geographic distribution and occupies different biomes, including urban areas, around Neotropical America. The bees of this species nest in trees or artificial cavities because of this broad distribution. ''N. testaceicornis'' is important for agriculture because it will pollinate a vast number of plant species year round.


Taxonomy

''Nannotrigona testaceicornis'' is a member of the family
Apidae Apidae is the largest family within the superfamily Apoidea, containing at least 5700 species of bees. The family includes some of the most commonly seen bees, including bumblebees and honey bees, but also includes stingless bees (also used for ...
of eusocial bees within the order
Hymenoptera Hymenoptera is a large order (biology), order of insects, comprising the sawfly, 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 Par ...
. The subfamily
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 A ...
is commonly referred to as "stingless bees." The genus ''Nannotrigonna'' has about nine known species and ranges from
Sonora Sonora (), officially Estado Libre y Soberano de Sonora ( en, Free and Sovereign State of Sonora), is one of the 31 states which, along with Mexico City, comprise the Administrative divisions of Mexico, Federal Entities of Mexico. The state is d ...
,
Chihuahua Chihuahua may refer to: Places *Chihuahua (state), a Mexican state **Chihuahua (dog), a breed of dog named after the state **Chihuahua cheese, a type of cheese originating in the state **Chihuahua City, the capital city of the state **Chihuahua Mun ...
, and
San Luis Potosí San Luis Potosí (), officially the Free and Sovereign State of San Luis Potosí ( es, Estado Libre y Soberano de San Luis Potosí), is one of the 32 states which compose the Federal Entities of Mexico. It is divided in 58 municipalities and i ...
,
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
, to Santa Catarina,
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
, and
Paraguay Paraguay (; ), officially the Republic of Paraguay ( es, República del Paraguay, links=no; gn, Tavakuairetã Paraguái, links=si), is a landlocked country in South America. It is bordered by Argentina to the south and southwest, Brazil to th ...
.


Description

''Nannotrigona testaceicornis'' bees are black and generally have grey hairs with a coarse and wrinkled
thorax The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the cre ...
. Along with the rest of the genus ''Nannotrigonis'', they are about 3 to 5 millimeters long. There is much variation within the species genotype, leading to variation in the venation of the wings. Colonies consist of about 2000 to 3000 individuals. Nests are made in tree cavities or in artificial sites, and the brood cells are in
combs Combs may refer to: Places France * Combs-la-Ville, a commune in the southeastern suburbs of Paris United Kingdom *Combs, Derbyshire, England *Combs, Suffolk, England United States *Combs, Arkansas, a community *Combs, Kentucky, a com ...
. The nest entrances are rather large for such small bees and are usually lined by workers looking outward. These openings are in contrast to the entrances of many other small
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 A ...
which are so small that one or a few workers can block them.


Distribution and habitat

''Nannotrigona testaceicornis'' is a neotropical eusocial bee that has a large geographic distribution, primarily throughout Brazil. It can occupy various different
biomes A biome () is a biogeographical unit consisting of a biological community that has formed in response to the physical environment in which they are found and a shared regional climate. Biomes may span more than one continent. Biome is a broader ...
, including urban areas. Due to this variability, different genomes can occur within the species according to the conditions of the biome. This can be seen mostly in the ''N. testaceicornis'' patterns of
wing venation Insect wings are adult outgrowths of the insect exoskeleton that enable insects to flight, fly. They are found on the second and third Thorax (insect anatomy), thoracic segments (the mesothorax and metathorax), and the two pairs are often referre ...
. Because of their occupation of urban areas, ''N. testaceicornis'' can live in self-made and artificial hives. Most
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 A ...
s construct their nests in pre-existing cavities using several types of materials. ''N. testaceicornis'' often use artificial cavities from human constructions to build their nests. The constant presence of ''N. testaceicornis'' in the human environment promotes competition for nesting places, leading to intensified aggressive interactions between colonies within the species.


Colony cycle

Caste Caste is a form of social stratification characterised by endogamy, hereditary transmission of a style of life which often includes an occupation, ritual status in a hierarchy, and customary social interaction and exclusion based on cultura ...
determination in
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 A ...
s is more variable than in
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 co ...
s. In
Melipona ''Melipona'' is a genus of stingless bees, widespread in warm areas of the Neotropics, from Sinaloa and Tamaulipas (México) to Tucumán and Misiones (Argentina). About 70 species are known.Grüter, C. 2020. ''Stingless Bees: Their Behaviour, Ec ...
,
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 ...
may be reared in both royal cells and small cells. Ones reared in small cells are smaller than normal queens, since they emerge from brood cells in which normally only workers and males are reared. Just like normal queens, miniature queens can successfully mate and head colonies. Queens and workers are produced in identically sized cells, and are typically produced in great excess of colony needs. ''Nannotrigona testaceicornis'' have low investments in queens similar to those of species that rear their queens only in royal cells. The production of miniature queens can be explained by the caste conflict hypothesis. It assumes that selfish females reared in normal-sized cells become queens, instead of workers, because they have direct benefits in reproduction. If the mated queen dies during reproductive
diapause In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press It ...
, there are new queens in the nest available to be selected by workers. The selected miniature queen can mate and replace the dead one, assuring the future of the colony.


Abdominal extracts

Abdominal extracts are differentiated between males and workers, but both come from the
Dufour's gland Dufour's gland is an abdominal gland of certain insects, part of the anatomy of the ovipositor or sting apparatus in female members of Apocrita. The diversification of Hymenoptera took place in the Cretaceous and the gland may have developed at a ...
, which is a large, wide, pear-shaped sac. The main chemical component of workers' abdominal extract is geranylgeranyl acetate while the extract from males' abdomens is (Z)-9-nonacosene. Experiments have shown that (S)-(+)-2-heptanol and (S)-(+)-2heptanol/(S)-(+)-2-nonanol (1:1) attract ''N. testaceicornis'' workers. However, males do not respond to these chemicals, implying that these compounds do not function as alarm or recruitment
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 behavio ...
s.


Behavior


Communication

A zigzag run and the jostling of nestmates inside the nest as well as airborne sounds and vibrational signals are observed as communication methods for ''N. testaceicornis'', rather than a scent trail. Both the vibrations and the jostling by the workers stimulate the nestmates to leave the nest and search for food. Depending on the profitability of the food source, workers will adjust their activity to recruit more or less nestmates. When a worker has been successful in finding a food source, it will return to the nest and begin bumping into nestmates and will stop for
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 ...
emitting thorax vibrations and airborne sounds, which can even be heard by the human ear.


Role differentiation

Female ''N. testaceicornis'' will have two sets of chromosomes (
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
), receiving one set from the
queen Queen or QUEEN may refer to: Monarchy * Queen regnant, a female monarch of a Kingdom ** List of queens regnant * Queen consort, the wife of a reigning king * Queen dowager, the widow of a king * Queen mother, a queen dowager who is the mother ...
and the other from a male
drone Drone most commonly refers to: * Drone (bee), a male bee, from an unfertilized egg * Unmanned aerial vehicle * Unmanned surface vehicle, watercraft * Unmanned underwater vehicle or underwater drone Drone, drones or The Drones may also refer to: ...
. Meanwhile, drones have one set of chromosomes (
haploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
), resulting from an unfertilized egg. The worker ''N. testaceicornis'' will primarily act as guards of the nest and search for food while the drones and queen stay inside the nest.


Caste determination

Caste Caste is a form of social stratification characterised by endogamy, hereditary transmission of a style of life which often includes an occupation, ritual status in a hierarchy, and customary social interaction and exclusion based on cultura ...
determination in stingless bees is still not fully understood. There are two main theories of determination, and there is not enough conclusive data to establish which one is correct for ''N. testaceicornis''. One theory stipulates that female larvae have the potential of following diverse pathways in development. Workers build royal cells for
queen Queen or QUEEN may refer to: Monarchy * Queen regnant, a female monarch of a Kingdom ** List of queens regnant * Queen consort, the wife of a reigning king * Queen dowager, the widow of a king * Queen mother, a queen dowager who is the mother ...
, but queens may also emerge from normal cells where males and workers emerge from. The queens in royal cells contain more food for the larvae, while the normal cells do not. In the normal cells, the larvae ingest less food and are therefore of smaller size and labeled as “miniature” or “dwarf” queens. The other theory formulates that during development, female larvae have the decision of becoming a worker or a queen and therefore have the power of self-determination. The miniature queens would still gain a higher pay-off being a small queen rather than a worker since they then have the potential to be selected by the workers as the next queen when the active queen dies.


Interspecies aggression

During a dispute, a swarm from one colony of ''N. testaceicornis'' will form a cloud of individuals near the active colony. A few individuals in the swarm will fly in the direction of the tube entrance of the active colony's nest, where they are met by the active colony’s guards. The attempted invasion promotes the formation of groups of two to four swarm and resident bees to fall to the ground and continue to fight until death. The active colony’s nest entrance will then be closed off by the residents. The failing swarm of ''N. testaceicornis'' will change their strategy and occupy an abandoned nest.


Sociality

''Nannotrigona testaceicornis'' defend by biting their predators and gather in large colonies of 2000 to 3000 individuals due to the lack of a toxic stinger. The more individuals in a colony, the more guards the colony has to watch the nest. In the day time, these guards will take shifts to block the entrance of the nest from unrelated individuals. At night, the guards will then seal the
cerumen Earwax, also known by the medical term cerumen, is a brown, orange, red, yellowish or gray waxy substance secreted in the ear canal of humans and other mammals. It protects the skin of the human ear canal, assists in cleaning and lubrication, a ...
tube entrance.


Importance to humans


Historical significance

''Nannotrigona testaceicornis'' was the first
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 A ...
whose communication about food sources was proven. Numerous people had noted the sudden appearance of large numbers of stingless bees on sweets that had been exposed without attracting bees, and therefore assumed communication. In 1953, W. E. Kerr put a dish of sugar syrup a few meters from a series of nests of ''N. testaceicornis''. No bees found it until one worker was taken to the dish and allowed to return to its hive, more and more bees arrived at the dish over time. These bees were also found to be inhabitants of a single hive.


Agriculture

''Nannotrigona testaceicornis'', along with other stingless bees, are used as crop
pollinators A pollinator is an animal that moves pollen from the male anther of a flower to the female 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 the maj ...
in greenhouses, especially during the winter season. They have been discovered to be polylectic, collecting pollen from the flowers of a variety of unrelated plants, making then great
pollinators A pollinator is an animal that moves pollen from the male anther of a flower to the female 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 the maj ...
.


References

{{Taxonbar, from=Q2050520 Meliponini Hymenoptera of South America Insects described in 1836 Taxa named by Amédée Louis Michel le Peletier