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''Melipona beecheii'' is a species of
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 gen ...
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 ...
. It is native to Central America from the
Yucatán Peninsula The Yucatán Peninsula (, also , ; es, Península de Yucatán ) is a large peninsula in southeastern Mexico and adjacent portions of Belize and Guatemala. The peninsula extends towards the northeast, separating the Gulf of Mexico to the north ...
in the north to
Costa Rica Costa Rica (, ; ; literally "Rich Coast"), officially the Republic of Costa Rica ( es, República de Costa Rica), is a country in the Central American region of North America, bordered by Nicaragua to the north, the Caribbean Sea to the no ...
in the south. ''M. beecheii'' was cultivated in the
Yucatán Peninsula The Yucatán Peninsula (, also , ; es, Península de Yucatán ) is a large peninsula in southeastern Mexico and adjacent portions of Belize and Guatemala. The peninsula extends towards the northeast, separating the Gulf of Mexico to the north ...
starting in the
pre-Columbian era In the history of the Americas, the pre-Columbian era spans from the original settlement of North and South America in the Upper Paleolithic period through European colonization, which began with Christopher Columbus's voyage of 1492. Usually, ...
by the ancient
Maya civilization The Maya civilization () of the Mesoamerican people is known by its ancient temples and glyphs. Its Maya script is the most sophisticated and highly developed writing system in the pre-Columbian Americas. It is also noted for its art, ...
. The Mayan name for ''M. beecheii'' is ''xunan kab'', which translates roughly to "regal lady bee". ''M. beecheii'' serves as the subject of various Mayan religious ceremonies.


Taxonomy and phylogeny

''M. beecheii'' 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. 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 ''Melipona'' contains nearly 50 other species.


Description and identification

''M. beecheii'' has a golden-yellowish and brown striped body with translucent wings. ''M. beecheii'' has small, white hairs covering the head, thorax, and abdomen. Queens, workers, and drones are all roughly the same size. Moreover, all members of the hive regardless of their future role develop in identical, mass-provisioned, sealed cells. This allows for self-determination of roles, the root of caste conflict within ''M. beecheii''.


Distribution and habitat

''M. beecheii'' is a tropical
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 gen ...
bee that has been observed in a variety of geographic locations with tropical climates. ''M. beecheii'' can be found in Central America, especially in the southern Yucatán Peninsula of Mexico,
Guatemala Guatemala ( ; ), officially the Republic of Guatemala ( es, República de Guatemala, links=no), is a country in Central America. It is bordered to the north and west by Mexico; to the northeast by Belize and the Caribbean; to the east by Hon ...
,
El Salvador El Salvador (; , meaning " The Saviour"), officially the Republic of El Salvador ( es, República de El Salvador), is a country in Central America. It is bordered on the northeast by Honduras, on the northwest by Guatemala, and on the south ...
,
Nicaragua Nicaragua (; ), officially the Republic of Nicaragua (), is the largest country in Central America, bordered by Honduras to the north, the Caribbean to the east, Costa Rica to the south, and the Pacific Ocean to the west. Managua is the coun ...
, and Costa Rica.


Conservation status

This bee has been observed primarily in
tropical forests Tropical forests (a.k.a. jungle) are forested landscapes in tropical regions: ''i.e.'' land areas approximately bounded by the tropic of Cancer and Capricorn, but possibly affected by other factors such as prevailing winds. Some tropical for ...
, both humid and arid. ''M. beecheii'' builds its nests inside hollow cavities of trees, and as a result, has been seriously affected by deforestation. Moreover, use of insecticides in the forest is increasing and interest in the ancient Mayan practice of beekeeping is decreasing. Beekeepers who work with ''M. beecheii'' in the Mayan zone in
Quintana Roo Quintana Roo ( , ), officially the Free and Sovereign State of Quintana Roo ( es, Estado Libre y Soberano de Quintana Roo), is one of the 31 states which, with Mexico City, constitute the 32 federal entities of Mexico. It is divided into 11 mu ...
state, Mexico, have reported a 93% decrease in hives over the past 25 years.


Colony cycle

In ''M. beecheii'', all females, even those designated to be workers, have the ability to develop as queens. As a result of this potential to self-determine, immature females may try to develop as queens to gain greater reproductive opportunities. When a colony's queen dies, or before the formation of new colonies by swarming, new queens are produced, only one of which is selected to serve in that function per colony. Workers kill the extra queens by biting off their heads, abdomens, and limbs. The average life expectancy of developing queens is 47 hours. Advantages to producing excess queens include the provision of spare queens in case of queen failure, and the ability to select the best queen from a pool of candidates.


Behavior


Division of labor

''M. beecheii'' exhibits
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 ...
specialization within the community. Single-foraging bees are responsible for harvesting one single commodity, such as
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 gametop ...
,
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 mutualist ...
, or
resin In polymer chemistry and materials science, resin is a solid or highly viscous substance of plant or synthetic origin that is typically convertible into polymers. Resins are usually mixtures of organic compounds. This article focuses on nat ...
, in a single day while multiforaging bees forage for two or three of those resources. About half of the bees are single-foraging and half are multiforaging. A switch in foraging specialization occurs very rapidly, often within a single day. No one style was found to be advantageous over the other in terms of feeding and resource-acquisition opportunities. However, activity patterns and longevity differed significantly between groups. Individuals responsible for foraging nectar were observed to be active all day and died, on average, after three full days of foraging. However, pollen foragers were only observed to be active for about 1–3 hours per day and lived for an average of 12 days. In spite of this contrast, both styles of foraging resulted in an equal number of flights in a lifetime. The way in which individual ''M. beecheii'' organisms' foraging style is designated within the community is still unknown.


Caste determination

Caste determination in
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 ...
is either trophogenic or genetic. In trophogenic caste determination, an organism's caste is dictated by the quality and quantity of food it received during development. In genetic caste determination, an organism's caste is inherent in its genetics. Trophogenic caste determination is much more common; genetic caste determination is quite rare. While there is some evidence for a nutritional influence on caste determination in ''Melipona'', in ''M. beecheii'' queens and workers have similar weights, supporting self-determination of caste.


Reproduction

Similar to most beehives, an ''M. beecheii'' colony has one fertile queen and numerous nonreproductive female worker bees. The queen that is fertile and lays eggs is known as the physogastric queen. However, in ''M. beecheii'' excess queens are produced, most of which are destined to remain virgins. These excess queens are almost always killed by workers before they are able to lay eggs.


Pre-discharge phase

The physogastric queen’s reproduction is dependent on the workers. The workers arrange the cells prior to the queen's arrival. When the queen arrives at a cell, the pre-discharge phase begins with a worker inserting themselves into a cell.


Discharge phase

The queen then performs
oviposition The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
, the process of depositing eggs into a cell. The queen taps the worker with her antennae and forelegs. This contact serves to signal the worker to insert larval food into the cell. Afterwards, the queen checks the cell and eats the trophic egg, an unfertilized egg that is made specifically for the queen's nutrition.


Abiotic factors

''Melipona beecheii'' reproduction depends on many abiotic factors, including the time of year, climate, and resources available. Extreme reproduction was observed at the end of the rainy season wherein an excess of resources were able to be stored. In contrast, due to poor resource obtainment, ''M. beecheii'' populations decreased when there were good foraging conditions and food storage had built up. When foraging conditions are good, less energy is expended on reproduction.


Interaction with other species


Diet

''Melipona beecheii'', like most other bee species, consumes nectar and pollen. ''M. beecheii'' exhibits foraging specialization within the community.


Predators

The typical predators of ''M. beecheii'' are many different species of birds, spiders, lizards, and other bugs. ''M. beecheii'' is an easier target because it is unable to sting as a form of defense.


Parasites

''Melipona beecheii'' is parasitized primarily by the parasitic phorid fly, ''Pseudohypocera kerteszi''. Other than this organism, there are not many parasites or disease-causing organisms that affect ''M. beecheii''. In general, American foulbrood, which is caused by '' Paenibacillus larvae'', is a harmful disease for all honey-producing bees, but there is not much data regarding its specific effect on ''M. beecheii''.


Mutualism

''Melipona beecheii'' often has mutualisms with flowering plants. ''M. beecheii'' pollinates the flowers by carrying both nectar and pollen between plants, allowing them to collect food for themselves and their colony in the process. While it is not yet clear which specific flowering plants ''M. beecheii'' forms mutualistic relationships with, the patterns and habits have been studied so a mutualistic relationship has been confirmed.


Competitors

''Melipona beecheiis main competitors are other pollinating insects. For ''M. beecheii'' in its tropical environment, its other pollinating, stingless bee competitors include '' Melipona fasciata'' and other nests of ''M. beecheii''.


Defense

Though ''M. beecheii'' is a stingless bee, it has the ability to bite other organisms when it feels threatened or is under attack. When in a communal setting, ''M. beecheii'' coordinates attack signals via secretions from their mandibular glands. The main pheromone involved in eliciting a communal attack response in ''M. beecheii'' is farnesyl acetate. When secreted, all individuals in the nest are stimulated to attack. An additional strategy that ''M. beecheii'' uses to protect itself from predation is remaining inside its nest. Because ''M. beecheii'' primarily builds its nest inside trees, remaining inside the nest proves to be a rather sturdy and safe environment to protect each individual organism from predation.


Human importance

Beekeepers in the Yucatán peninsula have harvested honey from the nests of ''M. beecheii'' until African honeybees arrived. As a result, ''M. beecheii'' is facing loss of habitat and the ancient tradition of stingless beekeeping is on the verge of dying out.


Agriculture

Because ''M. beecheii'' is a stingless bee and eats nectar, its
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
does not involve hunting pest insects. However, ''M. beecheii'' plays an extremely important role in honey production, as they have been kept for centuries almost exclusively in log nests primarily by people who live in the Yucatán Peninsula of Mexico. ''M. beecheii'' is an important source of honey both in the region and around the world. Due to the concern that stingless bee keeping is going extinct, the possibility of negative effects resulting from the impact of competition from feral African Apis mellifera, over-harvesting, failure to transfer colonies to hives or divide them, deforestation, hurricane damage and lack of instruction and incentive for new stingless bee keepers has been postulated and explored.


Stings

''Melipona beecheii'' is a stingless bee and, as a result, cannot sting humans or other organisms. This characteristic makes them ideal for
beekeeping Beekeeping (or apiculture) is the maintenance of bee colonies, commonly in man-made beehives. Honey bees in the genus ''Apis (insect), Apis'' are the most-commonly-kept species but other honey-producing bees such as ''Melipona'' stingless bees ar ...
. That being said, when agitated, ''M. beecheii'' does have the capacity to bite other organisms. However, there is no medical hazard or allergy factor involved.


History and art

In accordance with Mayan history, ''M. beecheii'' are the embodiment of a link to the spirit world, an endowment of the god Ah Muzen Cab. ''M. beecheii'' serves as the subject of various Mayan religious ceremonies. ''M. beecheii'' has been greatly revered by Mayan culture throughout history, leading to its keeping by the Mayan people as a means of attaining a higher-being, one which is closer to God. According to the Maya tradition, a priest would harvest ''M. beecheii'' honey during a religious ceremony that would take place twice a year. As a means of increasing the number of nests and honey production, beekeepers would regularly divide existing nests. This would also help to alleviate the predicament of excess queens that occurs with ''M. beecheii''.


References

{{Authority control beecheii Hymenoptera of North America Insects of Central America Insects of Mexico Insects described in 1831