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==== Soldiers ====
==== Soldiers ====
While the existence of a soldier caste is well known in [[ant]]s and [[termite]]s, the phenomenon was unknown among bees until 2012, when some stingless bees were found to have a similar caste of defensive specialists that help guard the nest entrance against intruders.<ref>{{harvtxt|Grüter|Menezes|Imperatriz-Fonseca|Ratnieks|2012}}</ref> To date, at least 10 species have been documented to possess such "soldiers", including ''[[Tetragonisca angustula]]'', ''[[Tetragonisca fiebrigi|T. fiebrigi]]'', and ''[[Frieseomelitta longipes]]'', with the guards not only larger, but also sometimes a different color from ordinary workers.<ref>{{harvtxt|Grüter|Segers|Menezes|Vollet-Neto|Falcón|von Zuben|Bitondi|Nascimento|Almeida|2017}}</ref><ref>{{harvtxt|Grüter|2020|p=258}}</ref>
While the existence of a soldier caste is well known in [[ant]]s and [[termite]]s, the phenomenon was unknown among bees until 2012, when some stingless bees were found to have a similar caste of defensive specialists that help guard the nest entrance against intruders.<ref>{{harvtxt|Grüter|Menezes|Imperatriz-Fonseca|Ratnieks|2012}}</ref> To date, at least 10 species have been documented to possess such "soldiers", including ''[[Tetragonisca angustula]]'', ''[[Tetragonisca fiebrigi|T. fiebrigi]]'', and ''[[Frieseomelitta longipes]]'', with the guards not only larger, but also sometimes a different color from ordinary workers.<ref>{{harvtxt|Grüter|Segers|Menezes|Vollet-Neto|Falcón|von Zuben|Bitondi|Nascimento|Almeida|2017}}</ref><ref>{{harvtxt|Grüter|2020|p=258}}</ref>


==== Division of labour ====
==== Division of labour ====
Vegetable resin, gathered from a variety of plant species in the wild, is an essential raw material brought back to the hive. Stored in small, sticky clumps in peripheral areas of the colony, it is often mistakenly treated as a synonym for [[propolis]]. However, in beekeeping terminology, propolis refers to a mixture of resin, wax, enzymes, and possibly other substances. Stingless bees go beyond the classic propolis by producing various derivatives from resins and wax, sometimes using pure resins for sealing or defense, a behavior not observed in ''Apis'' bees. Understanding these distinctions is vital for effective production and value addition to the meliponiculture activity.<ref name=":10" /><ref name=":11" /><ref name=":12" /><ref name=":13" />
Vegetable resin, gathered from a variety of plant species in the wild, is an essential raw material brought back to the hive. Stored in small, sticky clumps in peripheral areas of the colony, it is often mistakenly treated as a synonym for [[propolis]]. However, in beekeeping terminology, propolis refers to a mixture of resin, wax, enzymes, and possibly other substances. Stingless bees go beyond the classic propolis by producing various derivatives from resins and wax, sometimes using pure resins for sealing or defense, a behavior not observed in ''Apis'' bees. Understanding these distinctions is vital for effective production and value addition to the meliponiculture activity.<ref name=":10" /><ref name=":11" /><ref name=":12" /><ref name=":13" />


[[Honey]], a prized product of bee colonies, is crafted through the processing of [[nectar]]s, [[Honeydew (secretion)|honeydews]], and fruit juices by worker bees. They store these collected substances in an extension of their gut called a [[crop (anatomy)|crop]]. Back at the hive, the bees ripen or dehydrate the nectar droplets by spinning them inside their mouthparts until honey is formed. Ripening concentrates the nectar and increases the sugar content, though it is not nearly as concentrated as the honey from ''Apis mellifera''. Stored in food pots, meliponines' honey is often referred to as pot-honey due to its distinctive storage method. Stingless bee honeys differ from ''A. mellifera'' honey in terms of color, texture, and flavor, being more liquid with a higher water content. Rich in [[mineral]]s, [[amino acid]]s, and [[flavonoid]] compounds, the composition of honey varies among colonies of the same species, influenced by factors such as season, habitat, and collected resources.<ref name=":10" /><ref name=":11" /><ref name=":12" /><ref name=":13" />
[[Honey]], a prized product of bee colonies, is crafted through the processing of [[nectar]]s, [[Honeydew (secretion)|honeydews]], and fruit juices by worker bees. They store these collected substances in an extension of their gut called a [[crop (anatomy)|crop]]. Back at the hive, the bees ripen or dehydrate the nectar droplets by spinning them inside their mouthparts until honey is formed. Ripening concentrates the nectar and increases the sugar content, though it is not nearly as concentrated as the honey from ''Apis mellifera''. Stored in food pots, meliponines' honey is often referred to as pot-honey due to its distinctive storage method. Stingless bee honeys differ from ''A. mellifera'' honey in terms of color, texture, and flavor, being more liquid with a higher water content. Rich in [[mineral]]s, [[amino acid]]s, and [[flavonoid]] compounds, the composition of honey varies among colonies of the same species, influenced by factors such as season, habitat, and collected resources.<ref name=":10" /><ref name=":11" /><ref name=":12" /><ref name=":13" />


Special methods are being developed to harvest moderate amounts of honey from stingless bees in these areas without causing harm. For honey production, the bees need to be kept in a box specially designed to make the honey stores accessible without damaging the rest of the nest structure. Some recent box designs for honey production provide a separate compartment for the honey stores so the honey pots can be removed without spilling honey into other areas of the nest. Unlike a hive of commercial honeybees, which can produce 75 kg (165 lbs) of honey a year, a hive of Australian stingless bees produces less than 1 kg (2 lbs). Stingless bee honey has a distinctive "bush" taste—a mix of sweet and sour with a hint of fruit.<ref>{{harvtxt|Mduda|Hussein|Muruke|2023}}</ref><ref>{{harvtxt|Ferreira|Lencioni|Benassi|Barth|Bastos|2009}}</ref><ref>{{harvtxt|Sousa|Souza|Marques|Benassi|Gullón|Pintado|Magnani|2016}}</ref> The taste comes from plant resins—which the bees use to build their hives and honey pots—and varies at different times of year depending on the flowers and trees visited.
Special methods are being developed to harvest moderate amounts of honey from stingless bees in these areas without causing harm. For honey production, the bees need to be kept in a box specially designed to make the honey stores accessible without damaging the rest of the nest structure. Some recent box designs for honey production provide a separate compartment for the honey stores so the honey pots can be removed without spilling honey into other areas of the nest. Unlike a hive of commercial honeybees, which can produce 75 kg (165 lbs) of honey a year, a hive of Australian stingless bees produces less than 1 kg (2 lbs). Stingless bee honey has a distinctive "bush" taste—a mix of sweet and sour with a hint of fruit.<ref>{{harvtxt|Mduda|Hussein|Muruke|2023}}</ref><ref>{{harvtxt|Ferreira|Lencioni|Benassi|Barth|Bastos|2009}}</ref><ref>{{harvtxt|Sousa|Souza|Marques|Benassi|Gullón|Pintado|Magnani|2016}}</ref> The taste comes from plant resins—which the bees use to build their hives and honey pots—and varies at different times of year depending on the flowers and trees visited.
Stingless bees, as a collective group, display remarkable adaptability to diverse nesting sites. They can be found in exposed nests in trees, from living in ant and termite nests above and below ground to cavities in trees, trunks, branches, rocks, or even human constructions.<ref name=":14">{{harvtxt|Grüter|2020|pp=87-97}}</ref>
Stingless bees, as a collective group, display remarkable adaptability to diverse nesting sites. They can be found in exposed nests in trees, from living in ant and termite nests above and below ground to cavities in trees, trunks, branches, rocks, or even human constructions.<ref name=":14">{{harvtxt|Grüter|2020|pp=87-97}}</ref>


Many [[beekeeper]]s keep the bees in their original log [[beehive|hive]] or transfer them to a wooden box, as this makes controlling the hive easier. Some beekeepers put them in bamboos, flowerpots, coconut shells, and other recycling containers such as a water jug, a broken guitar, and other safe and closed containers.<ref>{{harvtxt|Venturieri|2004|pp=36-39}}</ref><ref>{{harvtxt|Contrera|Venturieri|2008}}</ref><ref name=":26" />
Many [[beekeeper]]s keep the bees in their original log [[beehive|hive]] or transfer them to a wooden box, as this makes controlling the hive easier. Some beekeepers put them in bamboos, flowerpots, coconut shells, and other recycling containers such as a water jug, a broken guitar, and other safe and closed containers.<ref>{{harvtxt|Venturieri|2004|pp=36-39}}</ref><ref>{{harvtxt|Contrera|Venturieri|2008}}</ref><ref name=":26" />


==== Exposed nests ====
==== Exposed nests ====


Some ''[[Partamona]]'' species exhibit a distinctive entrance architecture, where workers of ''[[Partamona helleri|P. helleri]]'' construct a large outer mud entrance leading to a smaller adjacent entrance. This unique design enables foragers to enter with high speed, bouncing off the ceiling of the outer entrance towards the smaller inner entrance. The peculiar appearance of this entrance has led to local names such as "toad mouth", highlighting the intriguing adaptations found in stingless bee nest entrances.<ref name=":15" />
Some ''[[Partamona]]'' species exhibit a distinctive entrance architecture, where workers of ''[[Partamona helleri|P. helleri]]'' construct a large outer mud entrance leading to a smaller adjacent entrance. This unique design enables foragers to enter with high speed, bouncing off the ceiling of the outer entrance towards the smaller inner entrance. The peculiar appearance of this entrance has led to local names such as "toad mouth", highlighting the intriguing adaptations found in stingless bee nest entrances.<ref name=":15" />
<gallery perrow="4" widths="160" caption="Different species nest entrances in Brazilian stingless bees">
<gallery perrow="4" widths="160" caption="Different species nest entrances in Brazilian stingless bees">
File:Abelha limão.jpg|''[[Lestrimelitta limao]]''
File:Abelha limão.jpg|''[[Lestrimelitta limao]]''
File:Frieseomelitta varia nest entrance.jpg|''[[Frieseomelitta varia]]''
File:Frieseomelitta varia nest entrance.jpg|''[[Frieseomelitta varia]]''
{{Main|Pollination}}
{{Main|Pollination}}
[[File:Bee pollinating a rose.jpg|thumb|Solitary bee visiting and pollinating a flower.]]
[[File:Bee pollinating a rose.jpg|thumb|Solitary bee visiting and pollinating a flower.]]
Bees play a critical role in the ecosystem, particularly in the pollination of natural vegetation. This activity is essential for the reproduction of various plant species, particularly in [[tropical forest]]s where most tree species rely on [[Entomophily|insect pollination]]. Even in temperate climates, where [[Anemophily|wind pollination]] is prevalent among forest trees, many bushes and herbaceous plants, rely on bees for pollination. The significance of bees extends to arid regions, such as [[Deserts and xeric shrublands|desertic and xeric shrublands]], where bee-pollinated plants are essential for preventing [[erosion]], supporting [[wildlife]], and ensuring [[Ecological stability|ecosystem stability]].<ref name=":23">{{harvtxt|Michener|2000|pp=4-5}}</ref>
Bees play a critical role in the ecosystem, particularly in the pollination of natural vegetation. This activity is essential for the reproduction of various plant species, particularly in [[tropical forest]]s where most tree species rely on [[Entomophily|insect pollination]]. Even in temperate climates, where [[Anemophily|wind pollination]] is prevalent among forest trees, many bushes and herbaceous plants, rely on bees for pollination. The significance of bees extends to arid regions, such as [[Deserts and xeric shrublands|desertic and xeric shrublands]], where bee-pollinated plants are essential for preventing [[erosion]], supporting [[wildlife]], and ensuring [[Ecological stability|ecosystem stability]].<ref name=":23">{{harvtxt|Michener|2000|pp=4-5}}</ref>


The impact of bee pollination on agriculture is substantial. In the late 1980s, certain plants were estimated to contribute between $4.6 to $18.9 billion to the U.S. economy, primarily through insect-pollinated crops. Although some bee-pollinated plants can self-pollinate in the absence of bees, the resulting crops often suffer from inbreeding depression. The quality and quantity of seeds or fruits are significantly enhanced when bees participate in the pollination process. Although estimates of crop pollination attributed to honey bees are uncertain, it is undeniable that bee pollination is a vital and economically valuable activity.<ref name=":23" />
The impact of bee pollination on agriculture is substantial. In the late 1980s, certain plants were estimated to contribute between $4.6 to $18.9 billion to the U.S. economy, primarily through insect-pollinated crops. Although some bee-pollinated plants can self-pollinate in the absence of bees, the resulting crops often suffer from inbreeding depression. The quality and quantity of seeds or fruits are significantly enhanced when bees participate in the pollination process. Although estimates of crop pollination attributed to honey bees are uncertain, it is undeniable that bee pollination is a vital and economically valuable activity.<ref name=":23" />
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== Relationship to humans ==
== Worldwide overview ==

=== Africa ===
Stingless bees also play a vital ecological role across [[Sub-Saharan Africa]] and [[Madagascar]]. To understand these insects on the African continent, it's important to consider the prevailing socio-economic and cultural contexts. Despite their ecological significance, the diversity, conservation, and behavior of these bees remain underexplored, particularly compared to better-studied regions such as [[South America]] and [[Southeast Asia]]. Also, honeybees were extensively researched, in contrast to native meliponines.<ref name=":31">{{harvtxt|Kajobe|2008}}</ref><ref name=":32">{{harvtxt|Byarugaba|2004}}</ref>

Africa is home to seven [[Biodiversity hotspot|biodiversity hotspots]], yet the recorded bee fauna is moderate relative to the continent's size. Madagascar stands out with exceptionally high levels of endemic species, though much of the bee diversity remains undocumented.<ref name=":30">{{harvtxt|Eardley|Gikungu|Schwarz|2009}}</ref>'''<ref name=":31" />''' Africa is home to aproximately 36 species of meliponines, including seven endemic to Madagascar. Most of these bees are found in equatorian regions (tropical forests and some savannahs).<ref>{{harvtxt|Grüter|2010|pp=47 & 49}}</ref>

Factors such as habitat destruction, pesticide use, and invasive species pose significant threats to these pollinators. Furthermore, high rates of nest mortality, driven by predation and human activity, exacerbate conservation challenges. Research indicates that stingless bees in Africa face greater pressures than their counterparts in the American and Asian tropics, underlining the urgency for targeted conservation measures.<ref name=":30" />'''<ref name=":31" />'''

Uganda's [[Bwindi Impenetrable National Park]] has shown the presence of at least five stingless bee species in, distributed across two genera: ''[[Meliponula]]'' and ''[[Hypotrigona]]''.<ref name=":31" />'''<ref name=":32" />''' In Madagascar, there is only one genus of stingless bees: ''[[Liotrigona]].''<ref name=":30" />

Meliponiculture, for example, is practised in Angola and Tanzania, and interest in managing stingless bees is growing in other African countries as well. <ref>{{harvtxt|Grüter|2010|p=27}}</ref>


=== Australia ===
=== Australia ===
"''Coot-tha''", which derives from "''ku-ta''", is one of the Aboriginal names for "wild stingless bee honey".<ref>{{harvtxt |Vit|Pedro|Roubik|2018}}</ref> Of the 1,600 species of wild bees native to Australia, about 14 are meliponines.<ref>{{cite news | url=http://www.abc.net.au/science/articles/2003/05/08/2045526.htm | title=Stingless bee rescue |author= Wendy Pyper |date=May 8, 2003 |work=ABC Science}}</ref> These species bear a variety of names, including Australian native honey bees, native bees, sugar-bag bees, and sweat bees (because they land on people's skin to collect sweat).<ref>{{Cite web |title=Australian Native Stingless Bees |url=https://www.aussiebee.com.au/australian-stingless-bees.html |access-date=2024-05-28 |website=www.aussiebee.com.au}}</ref> All are small and usually black in colour, with hairy extended hind legs for carrying [[nectar]] and [[pollen]]; because of the latter, they are sometimes mistaken for [[bumblebee]]s. The various stingless species look quite similar, with the two most common species, ''[[Tetragonula carbonaria]]'' and ''[[Austroplebeia australis]]'', displaying the greatest variation, as the latter is smaller and less active. Both of these inhabit the area around [[Brisbane]].<ref>{{Cite web |last=Steve |date=2015-05-25 |title=Native Stingless Bees - Tetragonula carbonaria |url=https://www.nativebeehives.com/native-stingless-bees-tetragonula-carbonaria/ |access-date=2024-05-28 |website=www.nativebeehives.com |language=en-AU}}</ref>
Of the 1,600 species of wild bees native to Australia, about 14 are meliponines.<ref>{{cite news | url=http://www.abc.net.au/science/articles/2003/05/08/2045526.htm | title=Stingless bee rescue |author= Wendy Pyper |date=May 8, 2003 |work=ABC Science}}</ref> "''Coot-tha''", which derives from "''ku-ta''", is one of the Aboriginal names for "wild stingless bee honey".<ref>{{harvtxt|Vit|Pedro|Roubik|2018}}</ref> These species bear a variety of names, including Australian native honey bees, native bees, sugar-bag bees, and sweat bees (because they land on people's skin to collect sweat).<ref>{{Cite web |title=Australian Native Stingless Bees |url=https://www.aussiebee.com.au/australian-stingless-bees.html |access-date=2024-05-28 |website=www.aussiebee.com.au}}</ref> The various stingless species look quite similar, with the two most common species, ''[[Tetragonula carbonaria]]'' and ''[[Austroplebeia australis]]'', displaying the greatest variation, as the latter is smaller and less active. Both of these inhabit the area around [[Brisbane]].<ref>{{Cite web |last=Steve |date=2015-05-25 |title=Native Stingless Bees - Tetragonula carbonaria |url=https://www.nativebeehives.com/native-stingless-bees-tetragonula-carbonaria/ |access-date=2024-05-28 |website=www.nativebeehives.com |language=en-AU}}</ref>


As stingless bees are usually harmless to humans, they have become an increasingly attractive addition to the suburban backyard. Most meliponine beekeepers do not keep the bees for [[honey]], but rather for the pleasure of conserving native species whose original [[habitat (ecology)|habitat]] is declining due to human development. In return, the bees [[pollinate]] crops, garden flowers, and bushland during their search for nectar and pollen. While a number of beekeepers fill a small niche market for bush honey, native meliponines only produce small amounts and the structure of their hives makes the honey difficult to extract. Only in warm areas of Australia such as [[Queensland]] and northern [[New South Wales]] are favorable for these bees to produce more honey than they need for their own survival. Most bees only come out of the hive when it is above about 18°C (64°F).<ref>{{Cite web|url=https://www.abc.net.au/news/2019-07-05/moves-underway-to-make-a-standard-for-native-bee-honey/11278346|title=Native bee honey set to have its own food standard|last=Thomas|first=Kerrin|date=2019-07-05|website=ABC News|language=en-AU|access-date=2019-07-10}}</ref> Harvesting honey from a nest in a cooler area could weaken or even kill the nest.
As stingless bees are usually harmless to humans, they have become an increasingly attractive addition to the suburban backyard. Most meliponine beekeepers do not keep the bees for [[honey]], but rather for the pleasure of conserving native species whose original [[habitat (ecology)|habitat]] is declining due to human development. In return, the bees [[pollinate]] crops, garden flowers, and bushland during their search for nectar and pollen. While a number of beekeepers fill a small niche market for bush honey, native meliponines only produce small amounts and the structure of their hives makes the honey difficult to extract. Only in warm areas of Australia such as [[Queensland]] and northern [[New South Wales]] are favorable for these bees to produce more honey than they need for their own survival. Most bees only come out of the hive when it is above about 18°C (64°F).<ref>{{Cite web|url=https://www.abc.net.au/news/2019-07-05/moves-underway-to-make-a-standard-for-native-bee-honey/11278346|title=Native bee honey set to have its own food standard|last=Thomas|first=Kerrin|date=2019-07-05|website=ABC News|language=en-AU|access-date=2019-07-10}}</ref> Harvesting honey from a nest in a cooler area could weaken or even kill the nest.
[[File:Pollinating bee covered with pollen.jpg|thumb|''[[Trigona spinipes]],'' covered with [[pollen]], visiting a flower of the vegetable sponge gourd (''Luffa cylindrica'') in [[Campinas]], [[Brazil]].]]
[[File:Pollinating bee covered with pollen.jpg|thumb|''[[Trigona spinipes]],'' covered with [[pollen]], visiting a flower of the vegetable sponge gourd (''Luffa cylindrica'') in [[Campinas]], [[Brazil]].]]


Brazil is home to several species bees belonging to Meliponini, with more than 300 species already identified and probably more yet to be discovered and described. They vary greatly in shape, size, and habits, and 20 to 30 of these species have good potential as honey producers. Although they are still quite unknown by most people, an increasing number of beekeepers (''meliponicultores'', in Portuguese) have been dedicated to these bees throughout the country.<ref name=":27" /><ref>{{harvtxt|Villas-Bôas|2018|p=17}}</ref> This activity has experienced significant growth since August 2004, when national laws were changed to allow native bee colonies to be freely marketed, which was previously forbidden in an unsuccessful attempt to protect these species. Nowadays the capture or destruction of existing colonies in nature is still forbidden, and only new colonies formed by the bees themselves in artificial traps can be collected from the wild.<ref>[https://conama.mma.gov.br/?option=com_sisconama&task=arquivo.download&id=440 CONAMA 2004 Resolution]</ref> Most colonies marketed are artificially produced by authorized beekeepers, through division of already existing captive colonies. Besides honey production, Brazilian stingless bees such as the jataí (''[[Tetragonisca angustula]]''), mandaguari (''[[Scaptotrigona postica]]''), and mandaçaia (''[[Melipona quadrifasciata]]'') serve as major pollinators of tropical plants and are considered the ecological equivalent of the honey bee.<ref name=":21" /><ref name=":22" />
Brazil is home to several species bees belonging to Meliponini, with more than 300 species already identified and probably more yet to be discovered and described. They vary greatly in shape, size, and habits, and 20 to 30 of these species have good potential as honey producers. Although they are still quite unknown by most people, an increasing number of beekeepers (''meliponicultores'', in Portuguese) have been dedicated to these bees throughout the country.<ref name=":27" /><ref>{{harvtxt|Villas-Bôas|2018|p=17}}</ref> This activity has experienced significant growth since August 2004, when national laws were changed to allow native bee colonies to be freely marketed, which was previously forbidden in an unsuccessful attempt to protect these species. Nowadays the capture or destruction of existing colonies in nature is still forbidden, and only new colonies formed by the bees themselves in artificial traps can be collected from the wild.<ref>[https://conama.mma.gov.br/?option=com_sisconama&task=arquivo.download&id=440 CONAMA 2004 Resolution]</ref> Most marketed colonies are artificially produced by authorized beekeepers, through division of already existing captive colonies. Besides honey production, Brazilian stingless bees such as the jataí (''[[Tetragonisca angustula]]''), mandaguari (''[[Scaptotrigona postica]]''), and mandaçaia (''[[Melipona quadrifasciata]]'') serve as major pollinators of tropical plants and are considered the ecological equivalent of the honey bee.<ref name=":21" /><ref name=":22" />


Also, much practical and academic work is being done about the best ways of keeping such bees, multiplying their colonies, and exploring the honey they produce. Among many others, species such as jandaíra (''[[Melipona subnitida]]'') and true ''uruçu'' (''[[Melipona scutellaris]]'') in the northeast of the country, mandaçaia (''[[Melipona quadrifasciata]]'') and yellow uruçu (''Melipona rufiventris'') in the south-southeast, tiúba or jupará (''[[Melipona interrupta]]''<ref>{{Cite web|url=http://moure.cria.org.br/catalogue?id=34747|title=+Moure's Bee Catalogue}}</ref>) and straw-bee (''[[Scaptotrigona polysticta]]'') in the north and jataí (''[[Tetragonisca angustula]]'') throughout the country are increasingly kept by small, medium, and large producers. Many other species as the mandaguari (''[[Scaptotrigona postica]]''), the guaraipo (''[[Melipona bicolor]]''), marmelada (''[[Frieseomelitta varia]]'') and the iraí (''[[Nannotrigona testaceicornis]]''), to mention a few, are also reared. Through the cultivation of honey or selling of colonies, keeping stingless bees is an increasingly profitable activity. A single colony of species like mandaçaia and true "uruçu" can be divided up to four times a year, and each of the new colonies obtained this way can be sold for about US$100.
Also, much practical and academic work is being done about the best ways of keeping such bees, multiplying their colonies, and exploring the honey they produce.<ref>{{Cite web |date=2024-07-26 |title=Arquivos Artigos - A.B.E.L.H.A. |url=https://abelha.org.br/artigos/ |access-date=2024-12-31 |website=abelha.org.br |language=pt-BR}}</ref> Among many others, species such as jandaíra (''[[Melipona subnitida]]'') and true ''uruçu'' (''[[Melipona scutellaris]]'') in the northeast of the country, mandaçaia (''[[Melipona quadrifasciata]]'') and yellow uruçu (''Melipona rufiventris'') in the south-southeast, tiúba or jupará (''[[Melipona interrupta]]'') and canudo (''[[Scaptotrigona polysticta]]'') in the north and jataí (''[[Tetragonisca angustula]]'') throughout the country are increasingly kept by small, medium, and large producers. Many other species as the mandaguari (''[[Scaptotrigona postica]]''), the guaraipo (''[[Melipona bicolor]]''), marmelada (''[[Frieseomelitta varia]]'') and the iraí (''[[Nannotrigona testaceicornis]]''), to mention a few, are also reared.<ref name=":33">{{Cite web |date=2022-02-21 |title=Fichas catalográficas das espécies relevantes para a meliponicultura - A.B.E.L.H.A. |url=https://abelha.org.br/fichas-catalograficas-das-especies-relevantes-para-a-meliponicultura-2/ |access-date=2024-12-31 |website=abelha.org.br |language=pt-BR}}</ref>


According to [[Chico Mendes Institute for Biodiversity Conservation|ICMBio]] and the [[Ministry of Environment and Climate Change (Brazil)|Ministry of the Environment]] there are presently four species of Meliponini listed in the [[IUCN Red List|National Red List of Threatened Species]] in Brazil. ''[[Melipona capixaba]]'', ''[[Melipona rufiventris]]'', ''[[Melipona scutellaris]]'', and ''[[Partamona littoralis]]'' all listed as Endangered (EN).<ref>{{Cite web |title=IBAMA |url=https://www.ibama.gov.br/component/legislacao/?view=legislacao&force=1&legislacao=134521#:~:text=Ementa:,Extin%C3%A7%C3%A3o%20-%20Peixes%20e%20Invertebrados%20Aqu%C3%A1ticos. |access-date=2024-02-26 |website=www.ibama.gov.br}}</ref><ref>[[Instituto Chico Mendes de Conservação da Biodiversidade|ICMBio]]. 2018. [https://www.gov.br/icmbio/pt-br/centrais-de-conteudo/publicacoes/publicacoes-diversas/livro_vermelho_2018_vol7.pdf Livro Vermelho da Fauna Brasileira Ameaçada de Extinção: Volume VII - Invertebrados]. ''In'': ICMBio. (Org.). [https://www.gov.br/icmbio/pt-br/centrais-de-conteudo/publicacoes/publicacoes-diversas/livro_vermelho_2018_vol1.pdf Livro Vermelho da Fauna Brasileira Ameaçada de Extinção]. Brasília: ICMBio. 727p.</ref>
According to [[Chico Mendes Institute for Biodiversity Conservation|ICMBio]] and the [[Ministry of Environment and Climate Change (Brazil)|Ministry of the Environment]] there are presently four species of Meliponini listed in the [[IUCN Red List|National Red List of Threatened Species]] in Brazil. ''[[Melipona capixaba]]'', ''[[Melipona rufiventris]]'', ''[[Melipona scutellaris]]'', and ''[[Partamona littoralis]]'' all listed as Endangered (EN).<ref>{{Cite web |title=IBAMA |url=https://www.ibama.gov.br/component/legislacao/?view=legislacao&force=1&legislacao=134521#:~:text=Ementa:,Extin%C3%A7%C3%A3o%20-%20Peixes%20e%20Invertebrados%20Aqu%C3%A1ticos. |access-date=2024-02-26 |website=www.ibama.gov.br}}</ref><ref>[[Instituto Chico Mendes de Conservação da Biodiversidade|ICMBio]]. 2018. [https://www.gov.br/icmbio/pt-br/centrais-de-conteudo/publicacoes/publicacoes-diversas/livro_vermelho_2018_vol7.pdf Livro Vermelho da Fauna Brasileira Ameaçada de Extinção: Volume VII - Invertebrados]. ''In'': ICMBio. (Org.). [https://www.gov.br/icmbio/pt-br/centrais-de-conteudo/publicacoes/publicacoes-diversas/livro_vermelho_2018_vol1.pdf Livro Vermelho da Fauna Brasileira Ameaçada de Extinção]. Brasília: ICMBio. 727p.</ref>


==== Honey production ====
==== Honey production ====
Although the colony population of most of these bees is much smaller than that of European bees, the productivity per bee can be quite high. Probably the world champions in honey productivity, the manduri (''[[Melipona marginata]]'') and the ''guaraipo'' (''M. bicolor''), live in swarms of only around 300 individuals, but can still produce up to 5 liters (.79 US gallon) of honey a year under the right conditions. The manduri is one of the smallest bees in the genus ''[[Melipona]]'', varying in length from 6 to 7 mm (15/64" to 9/32").<ref>{{harvtxt|Witter|Blochtein|Santos|2007|p=27}}</ref> Species of the genus ''Scaptotrigona'' have very large colonies, with up to 20,000 individuals, and can produce from 3 to 8 liters (2 US gallons) of honey a year.
Although the colony population of most of these bees is much smaller than that of European bees, the productivity per bee can be quite high. Interestingly, honey production is more connected to the body size, not the colony size. The manduri (''[[Melipona marginata]]''), jandaíra (''[[Melipona subnitida]]'') and the ''guaraipo'' (''[[Melipona bicolor|M. bicolor]]'') live in swarms of only around 300 individuals but can still produce up to 5 liters (1.3 US gallon) of honey a year under the right conditions.<ref>{{harvtxt|Grüter|2010|p=14}}</ref> In large bee farms, only the availability of flowers limits the honey production per colony. However, much larger numbers of beehives are required to produce amounts of honey comparable to that of European bees. Also, due to the fact of those bees storing honey in cerumen pots instead of standardized honeycombs as in the honeybee rearing makes extraction a lot more difficult and laborious.<ref>{{harvtxt|Fonseca|Sodré|Carvalho|Alves|Souza|Silva|Oliveira|Machado|Clarton|2007}}</ref>

Some large breeders have more than 3,000 hives of the tamer but still highly productive species in the genus ''Melipona'', such as the tiúba, the true ''uruçu'', and the jandaíra, each with 3,000 or more bees per colony. They can produce over 1.5 tons (3,000&nbsp;lbs) of honey every year.{{Citation needed|date=June 2024}} In large bee farms, only the availability of flowers limits the honey production per colony. Their honey is considered more palatable because it's not overly sweet, and it also is thought to have medicinal properties more pronounced than honey from bees of the genus ''Apis'' due to the higher level of antimicrobial substances. As a result, the honey from stingless bees returns very high profits, with prices greater than those for the more common honey produced by European bees. However, much larger numbers of beehives are required to produce amounts of honey comparable to that of European bees. Also, due to the fact of those bees storing honey in cerumen pots instead of standardized honeycombs as in the honeybee rearing makes extraction a lot more difficult and laborious.<ref>{{harvtxt|Fonseca|Sodré|Carvalho|Alves|Souza|Silva|Oliveira|Machado|Clarton|2007}}</ref>

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The honey from stingless bees has a lighter color and a higher water content, from 25% to 35%, compared to the honey from the genus ''Apis''. This contributes to its less cloying taste but also causes it to spoil more easily. Thus, for marketing, this honey needs to be processed through desiccation, fermentation or pasteurization. In its natural state, it should be kept under refrigeration.<ref>{{harvtxt|Grüter|2020|pp=12-14}}</ref>


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The honey from stingless bees has a higher water content, from 25% to 35%, compared to the honey from the genus ''Apis''. This contributes to its less cloying taste but also causes it to spoil more easily. Thus, for marketing, this honey needs to be processed through desiccation, fermentation or pasteurization. In its natural state, it should be kept under refrigeration.<ref>{{harvtxt|Grüter|2020|pp=12-14}}</ref>
==== Bees as pets ====
==== Bees as pets ====
[[Image:Mandaçaias.jpg|left|thumb|A swarm of mandaçaias bees around an artificial [[beehive]] installed in a house's backyard in [[Brazil]]]]
[[Image:Mandaçaias.jpg|left|thumb|A swarm of mandaçaias bees around an artificial [[beehive]] installed in a house's backyard in [[Brazil]]]]
Due to the lack of a functional stinger and characteristic nonaggressive behavior of many Brazilian species of stingless bees, they can be reared without problems in densely populated environments such as cities, provided enough flowers are at their disposal nearby. Some breeders (''meliponicultores'') can produce honey even in apartments up to the 12th floor.<ref>{{cite web|url=http://meliponariocapixaba.blogspot.com.br/2009/11/e-possivel-criar-abelhas-em.html|title=Meliponario Capixaba: É POSSÍVEL CRIAR ABELHAS EM APARTAMENTOS?|author=João Luiz|access-date=12 March 2016}}</ref>
Due to the lack of a functional stinger and characteristic nonaggressive behavior of many Brazilian species of stingless bees, they can be reared without problems in densely populated environments (residential buildings, schools, urban parks), provided enough flowers are at their disposal nearby. Some breeders (''meliponicultores'') can produce honey even in apartments up to the 12th floor.<ref>{{cite web|url=http://meliponariocapixaba.blogspot.com.br/2009/11/e-possivel-criar-abelhas-em.html|title=Meliponario Capixaba: É POSSÍVEL CRIAR ABELHAS EM APARTAMENTOS?|author=João Luiz|access-date=12 March 2016}}</ref>


The [[Melipona quadrifasciata|mandaçaias]] (''Melipona quadrifasciata'') are extremely tame, rarely attacking humans (only when their hives are opened for honey extraction or colony division). They form small, manageable colonies of only 400&ndash;600&nbsp;individuals. At the same time, a single rational beehive of ''mandaçaia'' can produce up to 4&nbsp;liters (1&nbsp;US gallon) of honey a year, making the species very attractive for home keepers. They are fairly large bees, up to 11&nbsp;mm (7/16") in length, and as a result have better body heat control, allowing them to live in regions where temperatures can drop a little lower than 0&nbsp;°C (32&nbsp;°F). However, they are somewhat selective about which flowers they will visit, preferring the flora that occurs in their natural environment. They are thus difficult to keep outside their region of origin (the northeast coast of Brazil).<ref>{{harvtxt|Brito|Faquinello|Paula-Leite|Carvalho|1997}}</ref>
The [[Melipona quadrifasciata|mandaçaias]] (''Melipona quadrifasciata'') are extremely tame, rarely attacking humans (only when their hives are opened for honey extraction or colony division). They form small, manageable colonies of only 400&ndash;600&nbsp;individuals. They are fairly large bees, up to 11&nbsp;mm (7/16") in length, and as a result have better body heat control, allowing them to live in regions where temperatures can drop a little lower than 0&nbsp;°C (32&nbsp;°F). However, they are somewhat selective about which flowers they will visit, preferring the flora that occurs in their natural environment. They are thus difficult to keep outside their region of origin (the eastern coast of Brazil). Once very common, the mandaçaia is now rather rare in nature, mainly due to the destruction of their native forests in the of Brazil.<ref>{{harvtxt|Brito|Faquinello|Paula-Leite|Carvalho|1997}}</ref><ref name=":33" />


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Other groups of Brazilian stingless bees, genera ''[[Plebeia]]'' and ''[[Leurotrigona]]'', are also very tame and much smaller, with one of them (''[[Plebeia minima]]'') reaching no more than 2.5&nbsp;mm (3/32") in length, and the ''[[lambe-olhos]]'' ("lick-eyes" bee, ''[[Leurotrigona muelleri]]'') being even smaller, at no more than 1.5&nbsp;mm (3/32"). Many of these species are known as mirim (meaning 'small' in the [[Tupi-Guarani languages]]). As a result, they can be kept in very small artificial hives, thus being of interest for keepers who want them as pollinators in small glasshouses or just for the pleasure of having a 'toy' bee colony at home.<ref name=":33" /><ref>{{Cite web |title=Lambe-olhos – Leurotrigona muelleri – Laboratório de Sistemática de Plantas |url=https://sites.usp.br/jardimdabotanicausprp/lambe-olhos-leurotrigona-muelleri/ |access-date=2024-12-31 |website=sites.usp.br}}</ref><ref>{{Cite web |title=Mirim – Plebeia droryana – Laboratório de Sistemática de Plantas |url=https://sites.usp.br/jardimdabotanicausprp/mirim-plebeia-droryana/ |access-date=2024-12-31 |website=sites.usp.br}}</ref> Being so tiny, these species produce only a very small amount of honey, typically less than 500&nbsp;ml (1/2 US pint) a year, so are not interesting for commercial honey production.
Other species, like the ''tiúba'' and the true ''uruçu'', are also very tame and highly productive. Their colonies have from 3,000&ndash;5,000 individuals (for comparison, the population of honeybee swarms can peak at 80,000&nbsp;individuals) and can produce up to 10 liters (2.6 US gallons) of honey a year.{{Citation needed|date=June 2024}} They can be easily kept at home but will survive only in regions with a warm climate, their larvae dying at temperatures lower than 12&nbsp;°C (54&nbsp;°F). The yellow ''uruçu'', however, can survive at temperatures lower than 0&nbsp;°C (32&nbsp;°F), and their colonies, bearing about 3,500&nbsp;individuals, can produce up to 6&nbsp;liters (1.5&nbsp;US gallons) of honey a year.{{Citation needed|date=June 2024}} But this species will react with powerful nipping if its nests are disturbed, and usually they are only kept by professional meliponicultors.


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[[Image:Abelha-jatai.jpg|right|thumb|Entrance pipe of ''jataís´s'' colony: The ''jataí'' bees build a wax pipe in the entrance of their nest.]]
Another suitable species for keeping at home is the ''guaraipo'' (''[[Melipona bicolor]]''). It is also quite tame, never attacking the beekeeper, and their colonies have fewer than 600 individuals. They can withstand temperatures as low as &minus;10&nbsp;°C (14&nbsp;°F), and each colony can produce over 3&nbsp;liters ({{frac|3|4}}&nbsp;US gallons) of honey a year.{{Citation needed|date=June 2024}} Their colonies usually have more than one single queen at a time (usually two or three, but sometimes up to five), a phenomenon called [[Gyne|polygyne]], and thus are less sensitive to the death of one queen, which can cause the loss of a whole colony in other species. But the ''guaraipo'' is very sensitive to low levels of humidity, and their hives must be equipped with means to keep a high moisture content. Once very common, the ''guaraipo'' is now rather rare in nature, mainly due to the destruction of their native forests in the south-southeast of Brazil.
⚫
Belonging to the same group, the jataí (''[[Tetragonisca angustula]]''), the marmelada (''[[Frieseomelitta varia]]''), and the moça-branca (''[[Frieseomelitta doederleini]]'') are intermediate in size between those very small species and the European bee. They are very adaptable species; the jataí, and can be reared in many different regions and environments, being quite common in most Brazilian cities. The jataí can bite when disturbed, but its jaws are weak, and in practice they are harmless, while the marmelada and moça-branca usually deposit propolis on their aggressors. Jataí is one of the first species to be kept by home beekeepers. Their nests can be easily identified in trees or wall cavities by the yellow wax pipe they build at the entrance, usually guarded by some soldier bees, which are stronger than regular worker bees. The marmelada and moça-branca make a little less honey, but it is denser and sweeter than most from other stingless bees and is considered very tasty.<ref name=":33" /><ref>{{harvtxt|Castanheira|Contel|2005|}}</ref>


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=== Central America ===
⚫
Other groups of Brazilian stingless bees, genera ''[[Plebeia]]'' and ''[[Leurotrigona]]'', are also very tame and much smaller, with one of them (''[[Plebeia minima]]'') reaching no more than 2.5&nbsp;mm (3/32") in length, and the ''[[lambe-olhos]]'' ("lick-eyes" bee, ''[[Leurotrigona muelleri]]'') being even smaller, at no more than 1.5&nbsp;mm (3/32"). Many of these species are known as ''mirim'' (meaning 'small' in the [[Tupi-Guarani languages]]). As a result, they can be kept in very small artificial hives, thus being of interest for keepers who want them as pollinators in small glasshouses or just for the pleasure of having a 'toy' bee colony at home. Being so tiny, these species produce only a very small amount of honey, typically less than 500&nbsp;ml (1/2 US pint) a year, so are not interesting for commercial honey production.

⚫
[[Image:Abelha-jatai.jpg|right|thumb|Entrance pipe of ''jataís´s'' colony: The ''jataí'' bees build a wax pipe in the entrance of their nest.]]
⚫
Belonging to the same group, the ''jataí'' (''[[Tetragonisca angustula]]''), the ''marmelada ''(''[[Frieseomelitta varia]]''), and the ''moça-branca'' (''[[Frieseomelitta doederleini]]'') are intermediate in size between those very small species and the European bee. They are very adaptable species; the ''jataí'', and can be reared in many different regions and environments, being quite common in most Brazilian cities. The ''jataí'' can bite when disturbed, but its jaws are weak, and in practice they are harmless, while the ''marmelada'' and ''moça-branca'' usually deposit propolis on their aggressors. Producing up to 1.5&nbsp;liters (0.4&nbsp;US gallons) of honey a year, their honey is considered among the best from stingless bees. In fact, the ''jataí'' was one of the first species to be kept by home beekeepers. Their nests can be easily identified in trees or wall cavities by the yellow wax pipe they build at the entrance, usually guarded by some soldier bees, which are stronger than regular worker bees. The'' marmelada'' and ''moça-branca'' make a little less honey, but it is denser and sweeter than most from other stingless bees and is considered very tasty.

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=== Maya stingless bees of Central America ===
[[Image:Colmena melipona.jpg|right|thumb|A Maya stingless bee hive: A piece of hollow log provides a home for meliponine bees in [[Belize]].]]
[[Image:Colmena melipona.jpg|right|thumb|A Maya stingless bee hive: A piece of hollow log provides a home for meliponine bees in [[Belize]].]]
The stingless bees ''[[Melipona beecheii]]'' and ''[[Melipona yucatanica|M. yucatanica]]'' are the primary native bees cultured in Central America, though a few other species are reported as being occasionally managed (e.g., ''[[Trigona fulviventris]]'' and ''[[Scaptotrigona mexicana]]'').<ref>{{harvtxt|Kent|1984}}</ref> They were extensively cultured by the [[Maya civilization]] for honey, and regarded as sacred. They continue to be cultivated by the modern [[Maya peoples]], although these bees are endangered due to massive deforestation, altered [[agriculture|agricultural]] practices (especially overuse of [[insecticide]]s), and changing beekeeping practices with the arrival of the [[Africanized honey bee]], which produces much greater honey crops.<ref name=":26">{{harvtxt|Villanueva|Roubik|Colli-Ucán|1998}}</ref>
The stingless bees ''[[Melipona beecheii]]'' and ''[[Melipona yucatanica|M. yucatanica]]'' are the primary native bees cultured in Central America, though a few other species are reported as being occasionally managed (e.g., ''[[Trigona fulviventris]]'' and ''[[Scaptotrigona mexicana]]'').<ref>{{harvtxt|Kent|1984}}</ref> They were extensively cultured by the [[Maya civilization]] for honey, and regarded as sacred. They continue to be cultivated by the modern [[Maya peoples]], although these bees are endangered due to massive deforestation, altered [[agriculture|agricultural]] practices (especially overuse of [[insecticide]]s), and changing beekeeping practices with the arrival of the [[Africanized honey bee]], which produces much greater honey crops.<ref name=":26">{{harvtxt|Villanueva|Roubik|Colli-Ucán|1998}}</ref>


==== History ====
==== History ====
Native meliponines (''M. beecheii'' being the favorite) have been kept by the lowland Maya for thousands of years. The [[Yucatec Maya language]] name for this bee is ''xunan kab'', meaning "(royal, noble) lady bee".<ref>{{cite web|title=Diccionario Introductorio|url=http://www.uqroo.mx/libros/maya/diccionario.pdf|work=uqroo.mx|publisher=Universidad De Quintana Roo|access-date=27 March 2014|language=es}}</ref> The bees were once the subject of religious ceremonies and were a symbol of the bee-god [[Ah-Muzen-Cab]], known from the ''[[Madrid Codex (Maya)|Madrid Codex]]''.
Native meliponines (''M. beecheii'' being the most common) have been kept by the lowland Maya for thousands of years. The [[Yucatec Maya language]] name for this bee is ''xunan kab'', meaning "(royal, noble) lady bee".<ref>{{harvtxt|Grüter|2010|p=54}}</ref><ref>{{cite web|title=Diccionario Introductorio|url=http://www.uqroo.mx/libros/maya/diccionario.pdf|work=uqroo.mx|publisher=Universidad De Quintana Roo|access-date=27 March 2014|language=es}}</ref> The bees were once the subject of religious ceremonies and were a symbol of the bee-god [[Ah-Muzen-Cab]], known from the ''[[Madrid Codex (Maya)|Madrid Codex]]''.<ref name=":34">{{Cite web |date=2016-07-17 |title=People and bees. Mayan bees in the Madrid Codex |url=https://ecomercioagrario.com/en/people-and-bees-mayan-bees-in-the-madrid-codex/ |access-date=2024-12-31 |website=eComercio Agrario |language=en-US}}</ref>


The bees were, and still are, treated as pets. Families would have one or many log-hives hanging in and around their houses. Although they are stingless, the bees do bite and can leave welts similar to a mosquito bite. The traditional way to gather bees, still favored among the locals, is find a wild hive, then the branch is cut around the hive to create a portable log, enclosing the colony. This log is then capped on both ends with another piece of wood or pottery and sealed with mud. This clever method keeps the melipine bees from mixing their brood, pollen, and honey in the same comb as do the European bees. The brood is kept in the middle of the hive, and the honey is stored in vertical "pots" on the outer edges of the hive. A temporary, replaceable cap at the end of the log allows for easy access to the honey while doing minimal damage to the hive. However, inexperienced handlers can still do irreversible damage to a hive, causing the hive to swarm and abscond from the log. With proper maintenance, though, hives have been recorded as lasting over 80 years, being passed down through generations. In the archaeological record of Mesoamerica, stone discs have been found that are generally considered to be the caps of long-disintegrated logs that once housed the beehives.
The bees were, and still are, treated as pets. Families would have one or many log-hives hanging in and around their houses. Although they are stingless, the bees do bite and can leave welts similar to a mosquito bite. The traditional way to gather bees, still favored among the locals, is find a wild hive, then the branch is cut around the hive to create a portable log, enclosing the colony. With proper maintenance, hives have been recorded as lasting over 80 years, being passed down through generations. In the archaeological record of Mesoamerica, stone discs have been found that are generally considered to be the caps of long-disintegrated logs that once housed the beehives.<ref name=":34" /><ref name=":35">{{Cite web |last=Kennedy |first=Jennifer |last2=Arghiris |first2=Richard |date=2019-01-31 |title=House of the Royal Lady Bee: Maya revive native bees and ancient beekeeping |url=https://news.mongabay.com/2019/01/house-of-the-royal-lady-bee-maya-revive-native-bees-and-ancient-beekeeping/ |access-date=2024-12-31 |website=Mongabay Environmental News |language=en-US}}</ref>


==== Tulum ====
==== Tulum ====
[[Tulum]], the site of a [[pre-Columbian]] Maya city on the Caribbean coast 130&nbsp;km (81&nbsp;mi) south of Cancun, has a god depicted repeatedly all over the site. Upside down, he appears as a small figure over many doorways and entrances. One of the temples, the "Templo del Dios Descendente" or the Temple of the Descending God, stands just left of the central plaza. Speculation is that he may be the "Bee God", Ah Muzen Cab, as seen in the Madrid Codex. It is possible that this was a religious/trade center with emphasis on ''xunan kab'', the "royal lady".
[[Tulum]], the site of a [[pre-Columbian]] Maya city on the Caribbean coast 130&nbsp;km (81&nbsp;mi) south of Cancun, has a god depicted repeatedly all over the site. Upside down, he appears as a small figure over many doorways and entrances. One of the temples, the Temple of the Descending God (''Templo del Dios Descendente''), stands just left of the central plaza. Speculation is that he may be the "Bee God", Ah Muzen Cab, as seen in the ''Madrid Codex''. It is possible that this was a religious/trade center with emphasis on ''xunan kab'', the "royal lady".<ref name=":35" />


==== Economic uses ====
==== Economic uses ====
[[Balché]], a traditional [[Mesoamerica]]n alcoholic beverage similar to [[mead]], was made from fermented honey and the bark of the leguminous'' balché'' tree (''[[Lonchocarpus violaceus]]''), hence its name.<ref>{{harvtxt|Ott|1998}}</ref> It was traditionally brewed in a canoe. The drink was known to have [[entheogen]]ic properties, that is, to produce mystical experiences, and was consumed in medicinal and ritual practices. Beekeepers would place the nests near the psychoactive plant ''[[Turbina corymbosa]]'' and possibly near ''balché'' trees, forcing the bees to use nectar from these plants to make their honey.<ref>{{harvtxt|Ott|1998}}</ref> Additionally, brewers would add extracts of the bark of the ''balché'' tree to the honey mixture before fermentation. The resulting beverage is responsible for psychotropic effects when consumed, due to the ergoline compounds in the pollen of the ''T. corymbosa'', the ''Melipona'' nectar gathered from the ''balché'' flowers, or the hallucinogenic compounds of the ''balché'' tree bark.
[[Balché]], a traditional [[Mesoamerica]]n alcoholic beverage similar to [[mead]], was made from fermented honey and the bark of the leguminous'' balché'' tree (''[[Lonchocarpus violaceus]]''), hence its name. It was traditionally brewed in a canoe. The drink was known to have [[entheogen]]ic properties, that is, to produce mystical experiences, and was consumed in medicinal and ritual practices. Beekeepers would place the nests near the psychoactive plant ''[[Turbina corymbosa]]'' and possibly near ''balché'' trees, forcing the bees to use nectar from these plants to make their honey. Additionally, brewers would add extracts of the bark of the ''balché'' tree to the honey mixture before fermentation. The resulting beverage is responsible for psychotropic effects when consumed, due to the ergoline compounds in the pollen of the ''T. corymbosa'', the ''Melipona'' nectar gathered from the ''balché'' flowers, or the hallucinogenic compounds of the ''balché'' tree bark.<ref>{{harvtxt|Ott|1998}}</ref>


[[Lost-wax casting]], a common metalworking method typically found where the inhabitants keep bees, was also used by the Maya. The wax from ''Melipona'' is soft and easy to work, especially in the humid Maya lowland. This allowed the Maya to create smaller works of art, jewelry, and other metalsmithing that would be difficult to forge. It also makes use of the leftovers from honey extraction. If the hive was damaged beyond repair, the whole of the comb could be used, thus using all of the hive. With experienced keepers, though, only the honey pot could be removed, the honey extracted, and the wax used for casting or other purposes.
[[Lost-wax casting]], a common metalworking method typically found where the inhabitants keep bees, was also used by the Maya. The wax from ''Melipona'' is soft and easy to work, especially in the humid Maya lowland. This allowed the Maya to create smaller works of art, jewelry, and other metalsmithing that would be difficult to forge. It also makes use of the leftovers from honey extraction. If the hive was damaged beyond repair, the whole of the comb could be used, thus using all of the hive. With experienced keepers, though, only the honey pot could be removed, the honey extracted, and the wax used for casting or other purposes.<ref>{{harvtxt|Pitses|2018|}}</ref><ref>{{harvtxt|Schwarz|1945|}}</ref>


==== Future ====
==== Future ====
The outlook for meliponines in Mesoamerica is uncertain. The number of active Meliponini beekeepers is shy in comparison with the Africanized ''[[Apis mellifera]]'' breeders. The high honey yield, 100&nbsp;kg (220&nbsp;lbs) or more annually, along with the ease of hive care and ability to create new hives from existing stock, commonly outweighs the negative consequences of "killer bee" hive maintenance.
The outlook for meliponines in Mesoamerica is uncertain. The number of active Meliponini beekeepers is shy in comparison with the Africanized ''[[Apis mellifera]]'' breeders. The high honey yield, 100&nbsp;kg (220&nbsp;lbs) or more annually, along with the ease of hive care and ability to create new hives from existing stock, commonly outweighs the negative consequences of "killer bee" hive maintenance.<ref name=":36">{{harvtxt|Villanueva-G|Roubik|Colli-Ucán|2005|}}</ref>

Further complicating the issue, [[Africanized honey bee]]s do not visit some flora, such as those in the [[Solanaceae|tomato family]], and several forest trees and shrubs, which rely on the native stingless bees for pollination. A decline in populations of native flora has already occurred in areas where stingless bees have been displaced by Africanized honey bees.


An additional blow to the art of meliponine beekeeping is that many of the meliponine beekeepers are now elderly, and their hives may not be cared for once they die. The hives are considered similar to an old family collection, to be parted out once the collector dies or to be buried in whole or part along with the beekeeper upon death. In fact, a survey of a once-popular area of the Maya lowlands shows the rapid decline of beekeepers, down to around 70 in 2004 from thousands in the late 1980s. It is traditional in the Maya lowlands that the hive itself or parts of the hive be buried along with the beekeeper to ''volar al cielo'', "to fly to heaven".{{citation needed|date=March 2007}} Conservation efforts are underway in several parts of Mesoamerica.<ref>A comprehensive conservation guide can be found in the June 2005 issue of ''Bee World''.</ref>
An additional blow to the art of meliponine beekeeping is that many of the ''meliponicultores'' are now elderly, and their hives may not be cared for once they die. The hives are considered similar to an old family collection, to be parted out once the collector dies or to be buried in whole or part along with the beekeeper upon death. In fact, a survey of a once-popular area of the Maya lowlands shows the rapid decline of beekeepers, down to around 70 in 2004 from thousands in the late 1980s. Conservation efforts are underway in several parts of Mesoamerica.<ref name=":36" /><ref>A comprehensive conservation guide can be found in the June 2005 issue of ''Bee World''.</ref>


== References ==<!-- ActaAmazonica34 p333 -->
== References ==<!-- ActaAmazonica34 p333 -->
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* {{Cite journal |last1=Viana |first1=Blandina Felipe |last2=Coutinho |first2=Jeferson Gabriel da Encarnação |last3=Garibaldi |first3=Lucas Alejandro |last4=Castagnino |first4=Guido Laercio Bragança |last5=Gramacho |first5=Kátia Peres |last6=Silva |first6=Fabiana Oliveira |date=2014-10-09 |title=Stingless bees further improve apple pollination and production |url=https://pollinationecology.org/index.php/jpe/article/view/309 |journal=Journal of Pollination Ecology |language=en |volume=14 |pages=261–269 |doi=10.26786/1920-7603(2014)26 |issn=1920-7603|doi-access=free }}
* {{Cite journal |last1=Viana |first1=Blandina Felipe |last2=Coutinho |first2=Jeferson Gabriel da Encarnação |last3=Garibaldi |first3=Lucas Alejandro |last4=Castagnino |first4=Guido Laercio Bragança |last5=Gramacho |first5=Kátia Peres |last6=Silva |first6=Fabiana Oliveira |date=2014-10-09 |title=Stingless bees further improve apple pollination and production |url=https://pollinationecology.org/index.php/jpe/article/view/309 |journal=Journal of Pollination Ecology |language=en |volume=14 |pages=261–269 |doi=10.26786/1920-7603(2014)26 |issn=1920-7603|doi-access=free }}
* {{cite journal |author=Villanueva, Rogel |display-authors=et al. |year=2005 |title=Extinction of ''Melipona beecheii'' and traditional beekeeping in the Yucatán peninsula |journal=Bee World |volume=86 |issue=2 |pages=35–41 |doi=10.1080/0005772X.2005.11099651 |s2cid=31943555}}
* {{cite journal |author=Villanueva, Rogel |display-authors=et al. |year=2005 |title=Extinction of ''Melipona beecheii'' and traditional beekeeping in the Yucatán peninsula |journal=Bee World |volume=86 |issue=2 |pages=35–41 |doi=10.1080/0005772X.2005.11099651 |s2cid=31943555}}
* {{Cite journal |last=Villanueva-G |first=Rogel |last2=Roubik |first2=David W |last3=Colli-Ucán |first3=Wilberto |date=2005-01 |title=Extinction of Melipona beecheii and traditional beekeeping in the Yucatán peninsula |url=http://www.tandfonline.com/doi/full/10.1080/0005772X.2005.11099651 |journal=Bee World |language=en |volume=86 |issue=2 |pages=35–41 |doi=10.1080/0005772X.2005.11099651 |issn=0005-772X}}
* {{Cite journal |last1=Wenseleers |first1=T. |last2=Ratnieks |first2=F. L. W. |last3=Ribeiro |first3=M. D. F. |last4=Alves |first4=D. D. A. |last5=Imperatriz-Fonseca |first5=V. L. |date=June 2005 |title=Working-class royalty: Bees beat the caste system |journal=[[Biology Letters]] |volume=1 |issue=2 |pages=125–128 |doi=10.1098/rsbl.2004.0281 |pmc=1626201 |pmid=17148145}}{{open access}}
* {{Cite journal |last1=Wenseleers |first1=T. |last2=Ratnieks |first2=F. L. W. |last3=Ribeiro |first3=M. D. F. |last4=Alves |first4=D. D. A. |last5=Imperatriz-Fonseca |first5=V. L. |date=June 2005 |title=Working-class royalty: Bees beat the caste system |journal=[[Biology Letters]] |volume=1 |issue=2 |pages=125–128 |doi=10.1098/rsbl.2004.0281 |pmc=1626201 |pmid=17148145}}{{open access}}
* {{Cite journal |last1=Witter |first1=Sidia |last2=Blochtein |first2=Betina |last3=Santos |first3=Camila dos |date=August 2007 |title=Abelhas sem ferrão do Rio Grande do Sul: manejo e conservação |url=https://www.agricultura.rs.gov.br/upload/arquivos/202105/11142006-boletim-15.pdf |journal=Boletim FEPAGRO |issue=15}}
* {{Cite journal |last1=Witter |first1=Sidia |last2=Blochtein |first2=Betina |last3=Santos |first3=Camila dos |date=August 2007 |title=Abelhas sem ferrão do Rio Grande do Sul: manejo e conservação |url=https://www.agricultura.rs.gov.br/upload/arquivos/202105/11142006-boletim-15.pdf |journal=Boletim FEPAGRO |issue=15}}
* {{Cite book |last=Grüter |first=Christoph |url=http://link.springer.com/10.1007/978-3-030-60090-7 |title=Stingless Bees: Their Behaviour, Ecology and Evolution |date=2020 |publisher=Springer International Publishing |isbn=978-3-030-60089-1 |series=Fascinating Life Sciences |location=Cham |language=en |doi=10.1007/978-3-030-60090-7|s2cid=227250633 }}
* {{Cite book |last=Grüter |first=Christoph |url=http://link.springer.com/10.1007/978-3-030-60090-7 |title=Stingless Bees: Their Behaviour, Ecology and Evolution |date=2020 |publisher=Springer International Publishing |isbn=978-3-030-60089-1 |series=Fascinating Life Sciences |location=Cham |language=en |doi=10.1007/978-3-030-60090-7|s2cid=227250633 }}
* {{Cite book |last=Jalil |first=Abu Hassan |url=https://books.google.com/books?id=fUh9BAAAQBAJ |title=Beescape for Meliponines: Conservation of Indo-Malayan Stingless Bees |date=2014 |publisher=Partridge Publishing Singapore |isbn=978-1-4828-2362-2 |language=ar}}
* {{Cite book |last=Jalil |first=Abu Hassan |url=https://books.google.com/books?id=fUh9BAAAQBAJ |title=Beescape for Meliponines: Conservation of Indo-Malayan Stingless Bees |date=2014 |publisher=Partridge Publishing Singapore |isbn=978-1-4828-2362-2 |language=ar}}
* {{Cite book |last=Kajobe |first=R. |url=https://books.google.com.br/books/about/Foraging_Behaviour_of_Equatorial_Afrotro.html?id=bMuezQEACAAJ&redir_esc=y |title=Foraging Behaviour of Equatorial Afrotropical Stingless Bees: Habitat Selection and Competition for Resources |date=2008 |publisher=Utrecht University |language=en}}
* {{Cite book |last=Michener |first=Charles D. |url=https://books.google.com/books?id=bu_1gmY13FIC |title=The Bees of the World |date=2000 |publisher=Johns Hopkins Univ. Press |isbn=978-0-8018-6133-8 |location=Baltimore, Md.}}
* {{Cite book |last=Michener |first=Charles D. |url=https://books.google.com/books?id=bu_1gmY13FIC |title=The Bees of the World |date=2000 |publisher=Johns Hopkins Univ. Press |isbn=978-0-8018-6133-8 |location=Baltimore, Md.}}
* {{Cite book |last=Roubik |first=David W. |url=https://www.cambridge.org/core/books/ecology-and-natural-history-of-tropical-bees/01B9D0798E5CBA8A68865ED0CC166A4A |title=Ecology and natural history of tropical bees |date=1989 |publisher=Cambridge Univ. Pr |isbn=978-0-521-26236-1 |series=Cambridge tropical biology series |location=Cambridge}}
* {{Cite book |last=Roubik |first=David W. |url=https://www.cambridge.org/core/books/ecology-and-natural-history-of-tropical-bees/01B9D0798E5CBA8A68865ED0CC166A4A |title=Ecology and natural history of tropical bees |date=1989 |publisher=Cambridge Univ. Pr |isbn=978-0-521-26236-1 |series=Cambridge tropical biology series |location=Cambridge}}

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'removed unreferenced and dubious sections | added refs | organized last sections | added africa section | improvements disscussed at [https://en.wikipedia.org/wiki/Wikipedia:Peer_review/Stingless_bee/archive1 Wikipedia:Peer review/Stingless bee/archive1]'
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'{{Short description|Bee tribe, reduced stingers, strong bites}} {{Automatic taxobox | name = Stingless bees | fossil_range = {{fossil_range|70.6|0|Maastrichtian-Present}} | image = Meliponula ferruginea.jpg | image_caption = ''Meliponula ferruginea'' | taxon = Meliponini | authority = [[Amédée Louis Michel le Peletier, comte de Saint-Fargeau|Lepeletier]], 1836 | subdivision_ranks = Genera | subdivision = {{ubl| ''[[Aparatrigona]]''| ''[[Apotrigona]]''| ''[[Asperplebeia]]''| ''[[Austroplebeia]]''| ''[[Axestotrigona]]''| ''[[Camargoia]]''| ''[[Celetrigona]]''| ''[[Cephalotrigona]]''| ''[[Cleptotrigona]]''| [[Extinction|†]]''[[Cretotrigona]]''| ''[[Dactylurina]]''| ''[[Dolichotrigona]]''| ''[[Duckeola]]''| ''[[Ebaiotrigona]]''| †''[[Exebotrigona]]''| ''[[Friesella]]''| ''[[Frieseomelitta]]''| ''[[Geniotrigona]]''| ''[[Geotrigona]]''| ''[[Heterotrigona]]''| ''[[Homotrigona]]''| ''[[Hypotrigona]]''| †''[[Kelneriapis]]''| ''[[Lepidotrigona]]''| ''[[Lestrimelitta]]''| ''[[Leurotrigona]]''| ''[[Liotrigona]]''| †''[[Liotrigonopsis]]''| ''[[Lisotrigona]]''| ''[[Lophotrigona]]''| ''[[Meliplebeia]]''| ''[[Melipona]]''| †''[[Meliponorytes]]''| ''[[Meliponula]]''| ''[[Meliwillea]]''| ''[[Mourella]]''| ''[[Nannotrigona]]''| ''[[Nanoplebeia]]''| ''[[Nogueirapis]]''| ''[[Odontotrigona]]''| ''[[Oxytrigona]]''| ''[[Papuatrigona]]''| ''[[Parapartamona]]''| ''[[Paratrigona]]''| ''[[Pariotrigona]]''| ''[[Paratrigonoides]]''| ''[[Partamona]]''| ''[[Platytrigona]]''| ''[[Plebeia]]''| ''[[Plebeiella]]''| ''[[Plebeina]]''| ''[[Plectoplebeia]]''| †''[[Proplebeia]]''| ''[[Ptilotrigona]]''| ''[[Sahulotrigona]]''| ''[[Scaptotrigona]]''| ''[[Scaura]]''| ''[[Schwarziana]]''| ''[[Sundatrigona]]''| ''[[Tetragona]]''| ''[[Tetragonilla]]''| ''[[Tetragonisca]]''| ''[[Tetragonula]]''| ''[[Tetrigona]]''| ''[[Trichotrigona]]''| ''[[Trigona (genus)|Trigona]]''| ''[[Trigonisca]]''| ''[[Wallacetrigona]]''}} }} '''Stingless bees (SB)''', sometimes called '''stingless honey bees''' or simply '''meliponines''', are a large group of bees (from about 462 to 552 described species),<ref>{{harvtxt|Grüter|2020|pp=46-47}}</ref><ref name=":27">{{harvtxt|Nogueira|2023}}</ref> comprising the [[Tribe (biology)|tribe]] '''Meliponini'''<ref>{{harvtxt|Michener|2000|p=803}}</ref><ref>{{harvtxt|Grüter|2020|p=1}}</ref> (or [[subtribe]] Meliponina according to other authors).<ref>{{harvtxt|Silveira|2002|p=253}}</ref> They belong in the [[Family (biology)|family]] [[Apidae]] ([[subfamily]] [[Apinae]]), and are closely related to common [[honey bee]]s (HB, tribe [[Apini]]), orchid bees (tribe [[Euglossini]]), and [[bumblebee]]s (tribe [[Bombini]]). These four bee tribes belong to the [[Pollen basket|corbiculate]] bees [[Monophyly|monophyletic]] group.<ref>{{harvtxt|Grüter|2020|p=43}}</ref><ref>{{harvtxt|Roubik|1989|p=8}}</ref> Meliponines have stingers, but they are highly reduced and cannot be used for defense, though these bees exhibit other defensive behaviors and mechanisms.<ref>{{harvtxt|Kajobe|2006}}</ref><ref name=":1">{{harvtxt|Chakuya|Gandiwa|Muboko|Muposhi|2022}}</ref> Meliponines are not the only type of bee incapable of stinging: all male bees and many female bees of several other families, such as [[Andrenidae]] and [[Megachilidae]] (tribe [[Dioxyini]]), also cannot sting.<ref>{{harvtxt|Michener|2000|p=111}}</ref> Some stingless bees have powerful mandibles and can inflict painful bites.<ref>{{harvtxt|Sarchet|2014}}</ref><ref>{{harvtxt|Grüter|2020|pp=7 & 16}}</ref> Some species can present large mandibular glands for the secretion of caustic defense substances, secrete unpleasant smells or use sticky materials to immobilise enemies.<ref name=":3">{{harvtxt|Grüter|2020|p=65}}</ref><ref>{{harvtxt|Grüter|2020|p=4}}</ref> The main honey-producing bees of this group generally belong to the genera ''[[Scaptotrigona]]'', ''[[Tetragonisca]]'', ''[[Melipona]]'' and ''[[Austroplebeia]]'', although there are other genera containing species that produce some usable honey. They are farmed in [[meliponiculture]] in the same way that European honey bees (genus ''[[Apis (genus)|Apis]]'') are cultivated in [[apiculture]]. Throughout [[Mesoamerica]], the [[Maya peoples|Mayans]] have engaged in extensive meliponiculture on a large scale since before the arrival of Columbus. Meliponiculture played a significant role in [[Maya society]], influencing their social, economic, and religious activities. The practice of maintaining '''stingless bees''' in man-made structures is prevalent across the Americas, with notable instances in countries such as [[Brazil]], [[Peru]], and [[Mexico]].<ref>{{harvtxt|Grüter|2020|pp=25-27}}</ref><ref>{{harvtxt|Villas-Bôas|2018|pp=14-15}}</ref> == Geographical distribution == Stingless bees can be found in most tropical or subtropical regions of the world, such as the [[Africa|African continent]] ([[Afrotropical realm|Afrotropical]] region), [[Southeast Asia]] and [[Australia]] ([[Indomalayan realm|Indo-Malayan]] and [[Australasian realm|Australasian]] region), and tropical [[Americas|America]] ([[Neotropical realm|Neotropical]] region).<ref>{{harvtxt|Michener|2000|p=803}}</ref><ref name=":0">{{harvtxt|Grüter|2020|pp=47-49}}</ref><ref>{{harvtxt|Silveira|2002|p=31}}</ref><ref>{{harvtxt|Rasmussen|Thomas|Engel|2017}}</ref> The majority of [[Indigenous (ecology)|native]] [[eusocial]] bees of [[Central America|Central]] and [[South America]] are SB, although only a few of them produce honey on a scale such that they are farmed by humans.<ref>{{harvtxt|Cortopassi-Laurino|Imperatriz-Fonseca|Roubik|2006}}</ref><ref>{{harvtxt|Venturieri|Raiol|Pareira|2003}}</ref> The Neotropics, with approximately 426 species, boast the highest abundance and species richness, ranging from Cuba and Mexico in the north to Argentina in the south.<ref name=":0" /> They are also quite diverse in [[Africa]], including [[Madagascar]],<ref>{{harvtxt|Koch|2010}}</ref> and are farmed there also. Around 36 species exist on the continent. The equatorial regions harbor the greatest diversity, with the [[Sahara|Sahara Desert]] acting as a natural barrier to the north. The range extends southward to [[South Africa]] and southern Madagascar, with most African species inhabiting [[tropical forest]]s or both tropical forests and [[savanna]]hs.<ref name=":0" /> Meliponine honey is prized as a medicine in many African communities, as well as in South America. Some cultures use SB honey against [[Human digestive system|digestive]], [[Respiratory system|respiratory]], [[Eye|ocular]] and [[Reproductive system|reproductive]] problems, although more research is needed to disclose evidence that supports these practices.<ref name=":1" /><ref>{{harvtxt|Grüter|2020|p=|pp=28-29}}</ref><ref>{{harvtxt|Souza|Yuyama|Aguiar|Oliveira|2004}}</ref> In Asia and Australia, approximately 90 species of stingless bees span from [[India]] in the west to the [[Solomon Islands]] in the east, and from [[Nepal]], China ([[Yunnan]], [[Hainan]]), and [[Taiwan]] in the north to Australia in the south. [[File:Geographic distribution of stingless bees.png|center|thumb|399x399px|Geographic distribution of stingless bees.]] === Origin and dispersion === Phylogenetic analyses reveal three distinct groups in the evolutionary history of '''Meliponini''': the Afrotropical, the Indo-Malay/Australasia, and the Neotropical lineages. The evolutionary origin of the Meliponini is Neotropical. Studies observing contemporary [[species richness]] show that it remains highest in the Neotropics.<ref name=":2">{{harvtxt|Roubik|2023a}}</ref> The hypothesis proposes the potential dispersion of stingless bees from what is now [[North America]]. According to this scenario, these bees would have then traveled to Asia by crossing the [[Bering Strait]] ([[Beringia]] route) and reached [[Europe]] through [[Greenland]] ([[Thule Land Bridge|Thulean route]]).<ref>{{harvtxt|Grüter|2020|pp=54-56}}</ref><ref name=":2" /><ref>{{harvtxt|Rasmussen|Cameron|2009}}</ref><ref>{{harvtxt|Jalil|2014|p=126}}</ref> == Evolution and phylogeny == Meliponines form a [[clade]] within the corbiculate bees, characterized by unique pollen-carrying structures known as corbiculae ([[Pollen basket|pollen baskets]]) located on their hind legs. This group also includes other three tribes: honey bees ([[Apini]]), bumble bees ([[Bombini]]), and orchid bees ([[Euglossini]]). The concept of higher [[eusociality]], defined by the presence of distinct queen and worker castes and characterized by features such as perennial colony lifestyles and extensive food sharing among adults, is particularly relevant in understanding the social structure of these tribes. Both Meliponini and Apini tribes are considered higher eusocial, while Bombini is considered to be primitively eusocial.<ref name=":29">{{harvtxt|Grüter|2020|pp=43-44}}</ref> The phylogenetic relationships among the four tribes of corbiculate bees have been a topic of considerable debate within the scientific community. Two primary questions arise: the relationship of stingless bees to honey bees and bumble bees, and whether their eusocial behavior evolved independently or from a [[Common descent|common ancestor]]. Morphological and behavioral studies have suggested that Meliponini and Apini are sister groups, indicating a single origin of higher eusociality. In contrast, molecular studies often support a relationship between Meliponini and Bombini, proposing independent origins of higher eusociality in both Apini and Meliponini.<ref name=":29" /> A morphological, behavioral, and molecular data analysis provided strong support for the latter hypothesis of dual origins of higher eusociality. Subsequent research has reinforced the idea that stingless bees and honey bees evolved their eusocial lifestyles independently, resulting in distinct adaptive strategies for colony reproduction, brood rearing, foraging communication, and colony defense. This divergence helps explain the varied ecological and social solutions developed by these two groups of bees, such as [[foraging]] communication, colony defense/reproduction and brood rearing.<ref name=":29" /><ref>{{harvtxt|Payne|2014}}</ref> {{clade| {{clade |1={{clade |1=[[Melittidae]] |2={{clade |label1={{nowrap|long-tongued bees}} |1={{clade |label1=&nbsp;[[Apidae]]&nbsp; |1={{clade |label2={{nowrap|eusocial}} |1=[[Euglossini]] |2={{clade |label1={{nowrap|h. eusocial}} |1=[[Apini]] |2={{clade |1=[[Bombini]] |label1={{nowrap|p. eusocial}} |2=[[Meliponini]] |label2={{nowrap|h. eusocial}} |style2=background-color:#f9a6e8 }} }}}} |3=[[Megachilidae]] (mason, leafcutter bees) }} |label2={{nowrap|short-tongued bees}} |2={{clade |1=[[Andrenidae]] (mining bees) |2={{clade |1=[[Halictidae]] (sweat bees) |2={{clade |2=[[Colletidae]] (plasterer bees) |1=[[Stenotritidae]] (large Australian bees) }} }} }} }} }} }} |style=font-size:85%;line-height:85%|label1=[[Anthophila]] (bees)}} === Fossil history === The [[fossil]] record for stingless bees is notably robust compared to that of many other bee groups, with twelve extinct species currently identified. Fossils of these bees are primarily found in [[amber]] and [[copal]], where excellent preservation typically occurs. This favorable fossilization process may be attributed to the behaviors of stingless bees, which collect significant amounts of tree [[resin]] for building nests and defense, increasing the likelihood of entrapment.<ref name=":24">{{harvtxt|Grüter|2010|pp=49-51}}</ref> Despite this relatively good fossil record, the evolutionary history of stingless bees remains poorly understood, particularly regarding their widespread distribution across various ecological niches around the globe. The oldest known fossil stingless bee is ''[[Cretotrigona prisca]]'', a small worker bee approximately 5 mm in body length, discovered in New Jersey amber. This species is believed to have existed during the [[Late Cretaceous]] period, around 65–70 million years ago, marking it as the oldest confirmed fossil of an apid bee and the earliest fossil evidence of a eusocial bee. ''C. prisca'' exhibits striking similarities to extant stingless bees, indicating that the evolutionary lineage of meliponines may date back to this period.<ref name=":24" /> Some researchers suggest that stingless bees likely evolved in the Late Cretaceous, approximately 70–87 million years ago.<ref name=":24" /><ref>{{harvtxt|Rasmussen|Cameron|2009}}</ref><ref>{{harvtxt|Cardinal|Danforth|2011}}</ref> According to recent studies, corbiculate bees, which include stingless bees, are thought to have appeared around 84–87 million years ago, further supporting the notion of their evolution during this dynamic period in Earth's history.<ref name=":24" /><ref>{{harvtxt|Cardinal|Danforth|2013}}</ref><ref>{{harvtxt|Martins|Melo|Renner|2014}}</ref> == Behaviour, biology and ecology == === Overview === Meliponines, considered highly eusocial insects, exhibit a remarkable caste division. The colonies typically consist of a [[Queen bee|queen]], [[Worker bee|workers]], and sometimes male [[Drone (bee)|drones]].<ref name=":4">{{harvtxt|Nogueira-Neto|1997|p=78}}</ref> The queen is responsible for reproduction, while the workers perform various tasks such as foraging, nursing, and defending the colony. Individuals work together with a well-defined division of labor for the overall benefit.<ref name=":5">{{harvtxt|Nogueira-Neto|1997|p=82}}</ref> Stingless bees are valuable pollinators and contribute to ecosystem health by producing essential products. These insects collect and store honey, pollen, resin, propolis, and cerumen. Honey serves as their primary [[carbohydrate]] source, while pollen provides essential [[protein]]s. Resin, propolis, and cerumen are used in nest construction and maintenance.<ref>{{harvtxt|Nogueira-Neto|1997|pp=40-46}}</ref><ref>{{harvtxt|Grüter|2020|pp=12-15}}</ref> Nesting behavior varies among species and may involve hollow tree trunks, external hives, the soil, termite nest or even urban structures. This adaptability underscores their resilience and ability to coexist with human activities.<ref>{{harvtxt|Grüter|2020|pp=87-121}}</ref> === Castes === ==== Workers ==== {{Main|Worker bee}} In a SB colony, workers constitute the predominant segment of the population, serving as the colony's primary workforce. They undertake a multitude of responsibilities crucial for the colony's well-being, including defense, cleaning, handling building materials, and the collection and processing of food. Recognizable by the corbicula - a distinctive structure on their hind legs resembling a small basket - workers efficiently carry pollen, resin, clay, and other materials gathered from the environment. Given their abundance and unique physical feature, workers play a central role in sustaining the colony.<ref>{{harvtxt|Grüter|2020|pp=7-11}}</ref><ref name=":9">{{harvtxt|Villas-Bôas|2018|p=23}}</ref><ref>{{harvtxt|Nogueira-Neto|1997|pp=82-83}}</ref> ==== Queens ==== {{Main|Queen bee}} The principal egg layer in SB colonies is the queen, distinguished from the workers by differences in both size and shape. Stingless bee queens - except in the case of the ''[[Melipona]]'' genus, where queens and workers receive similar amounts of food and thus exhibit similar sizes - are generally larger and weigh more than workers (approximately 2–6 times). Post-mating, meliponine queens undergo [[Physogastrism|physogastry]], developing a distended abdomen. This physical transformation sets them apart from honey bee queens, and even ''Melipona'' queens can be easily identified by their enlarged abdomen after mating.<ref name=":4" /><ref name=":6">{{harvtxt|Villas-Bôas|2018|p=19}}</ref><ref name=":7">{{harvtxt|Grüter|2020|pp=7 & 5}}</ref> Stingless bee colonies typically follow a monogynous structure, featuring a single egg-laying queen. An exception is noted in ''[[Melipona bicolor]]'' colonies, which are often polygynous (large populations may have as many as 5 physogastric queens simultaneously involved in oviposition).<ref name=":7" /><ref name=":5" /><ref name=":8">{{harvtxt|Imperatriz-Fonseca|Zucchi|1995}}</ref> Depending on the species, queens can lay varying quantities of eggs daily, ranging from a dozen (e.g., ''[[Plebeia julianii]]'') to several hundred (e.g., ''[[Trigona recursa]]''). While information on queen lifespans is limited, available data suggest that queens generally outlive workers, with lifespans usually falling between 1 and 3 years, although some queens may live up to 7 years.<ref name=":7" /><ref name=":8" /> The laying queen assumes the crucial role of producing eggs that give rise to all castes within the colony. Additionally, she plays a pivotal role in organizing the colony, overseeing a complex communication system primarily reliant on the use of [[pheromone]]s.<ref name=":6" /> ==== Males (drones) ==== {{Main|Drone (bee)}} The primary function of males, or drones, is to mate with queens, performing limited tasks within the nest and leaving at around 2–3 weeks old, never to return. The production of males can vary, occurring continuously, sparsely or in large spurts when numerous drones emerge from brood combs for brief periods. Identifying a male can be challenging due to its similar body size to workers, but distinctive features such as the absence of a corbicula, larger eyes, slightly smaller [[Mandible (insect mouthpart)|mandibles]], slightly longer and v-shaped [[Antenna (biology)|antennae]], and often a lighter face color distinguish them. Clusters of males, numbering in the hundreds, can be observed outside colonies, awaiting the opportunity to mate with virgin queens.<ref name=":9" /><ref>{{harvtxt|Nogueira-Neto|1997|pp=83-85}}</ref><ref>{{harvtxt|Grüter|2020|p=11}}</ref> Males in a stingless bee colony, either produced mainly by the laying queen or primarily by the workers, play an important role in reproduction. Workers can produce males by laying unfertilized eggs, enabled by the [[haplodiploidy]] system, where males are [[Ploidy|haploid]], having only one set of [[chromosome]]s, while workers are diploid and incapable of producing female eggs due to their inability to mate. This sex determination system is common to all [[hymenoptera]]ns.<ref>{{harvtxt|Grüter|2020|p=11}}</ref> ==== Soldiers ==== While the existence of a soldier caste is well known in [[ant]]s and [[termite]]s, the phenomenon was unknown among bees until 2012, when some stingless bees were found to have a similar caste of defensive specialists that help guard the nest entrance against intruders.<ref>{{harvtxt|Grüter|Menezes|Imperatriz-Fonseca|Ratnieks|2012}}</ref> To date, at least 10 species have been documented to possess such "soldiers", including ''[[Tetragonisca angustula]]'', ''[[Tetragonisca fiebrigi|T. fiebrigi]]'', and ''[[Frieseomelitta longipes]]'', with the guards not only larger, but also sometimes a different color from ordinary workers.<ref>{{harvtxt|Grüter|Segers|Menezes|Vollet-Neto|Falcón|von Zuben|Bitondi|Nascimento|Almeida|2017}}</ref><ref>{{harvtxt|Grüter|2020|p=258}}</ref> ==== Division of labour ==== When the young worker bees emerge from their cells, they tend to initially remain inside the hive, performing different jobs. As workers age, they become guards or foragers. Unlike the [[larva]]e of [[honey bee]]s and many social wasps, meliponine larvae are not actively fed by adults ([[progressive provisioning]]). Pollen and nectar are placed in a cell, within which an [[egg]] is laid, and the cell is sealed until the adult bee emerges after [[pupa]]tion ([[mass provisioning]]). At any one time, hives can contain from 300 to more than 100,000 workers (with some authors claiming to calculate more than 150,000 workers, but with no methodology explanation), depending on species.<ref>{{harvtxt|Grüter|2020|pp=17-24}}</ref> === Products and materials === The industrious nature of stingless bees extends to their building activities. Unlike honey bees, they do not use pure wax for construction but combine it with resin to create cerumen, a material employed in constructing nest structures such as brood cells, food pots, and the protective involucrum. [[Beeswax|Wax]] is secreted by young bees through glands located on the top of their abdomen and this mixture not only provides structural strength but also offers antimicrobial properties, inhibiting the growth of fungi and bacteria. The creation of batumen involves combining cerumen with additional resin, mud, plant material, and sometimes even animal feces. Batumen, a stronger material, forms protective layers covering the walls of the nesting space, ensuring the safety of the colony.<ref name=":10">{{harvtxt|Nogueira-Neto|1997|pp=40-46}}</ref><ref name=":11">{{harvtxt|Grüter|2020|pp=12-15}}</ref><ref name=":12">{{harvtxt|Villas-Bôas|2018|pp=24-26}}</ref><ref name=":13">{{harvtxt|Venturieri|2004|pp=23-30}}</ref> On the other hand, [[clay]], sourced from the wild and exhibiting diverse colors based on its mineral origin, serves as another essential raw material for SB. While it can be used in its pure form, it is more common to combine clay with vegetable [[resin]]s to produce geopropolis. The inclusion of clay in this mixture enhances the durability and structural integrity of the resulting substance.<ref name=":10" /><ref name=":11" /><ref name=":12" /><ref name=":13" /> Vegetable resin, gathered from a variety of plant species in the wild, is an essential raw material brought back to the hive. Stored in small, sticky clumps in peripheral areas of the colony, it is often mistakenly treated as a synonym for [[propolis]]. However, in beekeeping terminology, propolis refers to a mixture of resin, wax, enzymes, and possibly other substances. Stingless bees go beyond the classic propolis by producing various derivatives from resins and wax, sometimes using pure resins for sealing or defense, a behavior not observed in ''Apis'' bees. Understanding these distinctions is vital for effective production and value addition to the meliponiculture activity.<ref name=":10" /><ref name=":11" /><ref name=":12" /><ref name=":13" /> [[Honey]], a prized product of bee colonies, is crafted through the processing of [[nectar]]s, [[Honeydew (secretion)|honeydews]], and fruit juices by worker bees. They store these collected substances in an extension of their gut called a [[crop (anatomy)|crop]]. Back at the hive, the bees ripen or dehydrate the nectar droplets by spinning them inside their mouthparts until honey is formed. Ripening concentrates the nectar and increases the sugar content, though it is not nearly as concentrated as the honey from ''Apis mellifera''. Stored in food pots, meliponines' honey is often referred to as pot-honey due to its distinctive storage method. Stingless bee honeys differ from ''A. mellifera'' honey in terms of color, texture, and flavor, being more liquid with a higher water content. Rich in [[mineral]]s, [[amino acid]]s, and [[flavonoid]] compounds, the composition of honey varies among colonies of the same species, influenced by factors such as season, habitat, and collected resources.<ref name=":10" /><ref name=":11" /><ref name=":12" /><ref name=":13" /> Special methods are being developed to harvest moderate amounts of honey from stingless bees in these areas without causing harm. For honey production, the bees need to be kept in a box specially designed to make the honey stores accessible without damaging the rest of the nest structure. Some recent box designs for honey production provide a separate compartment for the honey stores so the honey pots can be removed without spilling honey into other areas of the nest. Unlike a hive of commercial honeybees, which can produce 75 kg (165 lbs) of honey a year, a hive of Australian stingless bees produces less than 1 kg (2 lbs). Stingless bee honey has a distinctive "bush" taste—a mix of sweet and sour with a hint of fruit.<ref>{{harvtxt|Mduda|Hussein|Muruke|2023}}</ref><ref>{{harvtxt|Ferreira|Lencioni|Benassi|Barth|Bastos|2009}}</ref><ref>{{harvtxt|Sousa|Souza|Marques|Benassi|Gullón|Pintado|Magnani|2016}}</ref> The taste comes from plant resins—which the bees use to build their hives and honey pots—and varies at different times of year depending on the flowers and trees visited. In 2020 researchers at the [[University of Queensland]] found that some species of stingless bee in Australia, Malaysia, and Brazil produce honey that has [[trehalulose]]—a sugar with an unusually low [[Glycemic index|glycaemic index]] (GI) compared to that of glucose and fructose, the main sugars composing conventional honey.<ref>{{harvtxt|Mokaya|Nkoba|Ndunda|Vereecken|2002}}</ref><ref name=":25">{{harvtxt|Fletcher|Hungerford|Webber|Carpinelli de Jesus|Zhang|Stone|Blanchfield|Zawawi|2020a}}</ref> Such low glycaemic index honey is beneficial for humans because its consumption does not cause [[blood sugar level|blood sugar]] to spike, forcing the body to make more insulin in response. Honey with trehalulose is also beneficial as it this sugar cannot nourish the lactic acid-producing bacteria that cause [[tooth decay]]. The university's findings supported the long-standing claims of [[Indigenous Australian people]] that native honey is beneficial to human health.<ref name=":25" /><ref>{{Cite web |last=Layt |first=Stuart |date=2020-07-23 |title=Scientists say native stingless bee honey hits the sweet spot |url=https://www.brisbanetimes.com.au/national/queensland/scientists-say-native-stingless-bee-honey-hits-the-sweet-spot-20200723-p55eud.html |access-date=2020-07-27 |website=Brisbane Times |language=en}}</ref> This type of honey is scientifically supported as providing therapeutic value to humans as well.<ref name=":25" /><ref>{{harvtxt|Mduda|Hussein|Muruke|2023}}</ref><ref>{{harvtxt|Rodríguez-Malaver|Rasmussen|Gutiérrez|Gil|Nieves|Vit|2009}}</ref><ref>{{harvtxt|Nweze|Okafor|Nweze|Nweze|2017}}</ref><ref>{{harvtxt|Zulkhairi Amin|Sabri|Mohammad|Ismail|Chan|Ismail|Norhaizan|Norhaizan|2018}}</ref> === Nest === [[File:Natural cavity stingless bee nest.png|thumb|General main elements of stingless bees cavity nest colonies.]] Stingless bees, as a collective group, display remarkable adaptability to diverse nesting sites. They can be found in exposed nests in trees, from living in ant and termite nests above and below ground to cavities in trees, trunks, branches, rocks, or even human constructions.<ref name=":14">{{harvtxt|Grüter|2020|pp=87-97}}</ref> Many [[beekeeper]]s keep the bees in their original log [[beehive|hive]] or transfer them to a wooden box, as this makes controlling the hive easier. Some beekeepers put them in bamboos, flowerpots, coconut shells, and other recycling containers such as a water jug, a broken guitar, and other safe and closed containers.<ref>{{harvtxt|Venturieri|2004|pp=36-39}}</ref><ref>{{harvtxt|Contrera|Venturieri|2008}}</ref><ref name=":26" /> ==== Exposed nests ==== [[File:Arboreal stingless bee nest (Trigona sp.) Flores.jpg|thumb|''Trigona'' sp. exposed nest in a tree fork near [[Flores, Petén|Flores]], [[Guatemala]].]] Notably, certain species, such as the African ''Dactylurina'', construct hanging nests from the undersides of large branches for protection against adverse weather conditions. Additionally, some American ''[[Trigona]]'' species, including ''[[Trigona corvina|T. corvina]]'', ''[[Trigona spinipes|T. spinipes]]'', and T. nigerrima, as well as ''Tetragonisca weyrauchi'', build fully exposed nests.<ref name=":14" /> ==== Ground nests ==== [[File:Paratrigona ground nest.jpg|left|thumb|''Paratrigona'' sp. ground nest in [[Brasília]], [[Brazil]].]] A significant minority of meliponine species, belonging to genera like ''[[Camargoia]]'', ''[[Geotrigona]]'', ''[[Melipona]]'', ''[[Mourella]]'', ''[[Nogueirapis]]'', ''[[Paratrigona]]'', ''[[Partamona]]'', ''[[Schwarziana]]'', and others, opt for ground nests. These species take advantage of cavities in the ground, often utilizing abandoned nests of ants, termites, or rodents. Unlike some other cavity-nesting bees, stingless bees in this category do not excavate their own cavities but may enlarge existing ones.<ref name=":14" /> ==== Termite and ant shared nests ==== Numerous stingless bee species have evolved to coexist with termites. They inhabit parts of ant or termite nests, both above and below ground. These nests are often associated with various ant species, such as ''[[Azteca (ant)|Azteca]]'', ''[[Carpenter ant|Camponotus]]'', or ''[[Crematogaster]]'', and termite species like ''[[Nasutitermes]]'', ''[[Constrictotermes]]'', ''[[Macrotermes]]'', ''Microcerotermes'', ''[[Odontotermes]]'', or ''Pseudocanthotermes''. This strategy allows SB to utilize pre-existing cavities without the need for extensive excavation.<ref name=":14" /> ==== Cavity nests ==== The majority of stingless bees favor nesting in pre-existing cavities within tree trunks or branches. Nesting heights vary, with some colonies positioned close to the ground, typically below 5 meters, while others, like ''Trigona'' and ''[[Oxytrigona]]'', may nest at higher elevations, ranging from 10 to 25 meters. Some species, such as ''[[Melipona nigra]]'', exhibit unique nesting habits at the foot of a tree in root cavities or between roots. The choice of nesting height has implications for predation pressure and the microclimate experienced by the colony.<ref name=":14" /> The majority of stingless bee species exhibit a non-specific preference when it comes to selecting tree species for nesting. Instead, they opportunistically exploit whatever nesting sites are available This adaptability underscores the versatility of SB in adapting to various arboreal environments. Furthermore, cavity-nesting species can opportunistically utilize human constructions, nesting under roofs, in hollow spaces in walls, electricity boxes, or even metal tubes. In few cases, specific tree species, like ''[[Caryocar brasiliense]]'', may be preferred by certain stingless bee species (''[[Melipona quadrifasciata]]''), illustrating a degree of selectivity in nesting choices among different groups.<ref name=":14" /><ref>{{harvtxt|Antonini|Martins|2003}}</ref> ==== Entrances ==== Entrance tubes showcase a spectrum of characteristics, from being hard and brittle to soft and flexible. In many situations, the portion near the opening remains soft and flexible, aiding workers in sealing the entrance during the night. The tubes may also feature perforations and a coating of resin droplets, adding to the complexity of their design.<ref name=":15">{{harvtxt|Grüter|2020|pp=102-109}}</ref> The entrances serve as essential visual landmarks for returning bees, and they are often the first structures constructed at a new nest site. The diversity in entrance size influences foraging traffic, with larger entrances facilitating smoother traffic but potentially necessitating more entrance guards to ensure adequate defense.<ref name=":15" /> Some ''[[Partamona]]'' species exhibit a distinctive entrance architecture, where workers of ''[[Partamona helleri|P. helleri]]'' construct a large outer mud entrance leading to a smaller adjacent entrance. This unique design enables foragers to enter with high speed, bouncing off the ceiling of the outer entrance towards the smaller inner entrance. The peculiar appearance of this entrance has led to local names such as "toad mouth", highlighting the intriguing adaptations found in stingless bee nest entrances.<ref name=":15" /> <gallery perrow="4" widths="160" caption="Different species nest entrances in Brazilian stingless bees"> File:Abelha limão.jpg|''[[Lestrimelitta limao]]'' File:Frieseomelitta varia nest entrance.jpg|''[[Frieseomelitta varia]]'' File:Partamona helleri colmeia.jpg|''[[Partamona helleri]]'' File:Entrada de ninho de Scaptotrigona postica.jpg|''[[Scaptotrigona postica]]'' File:Parque Nacional da Serra do Itajaí - Liu Idárraga Orozco (86).jpg|''[[Oxytrigona tataira]]'' File:Melipona quadrifasciata nest guard.jpg|''[[Melipona quadrifasciata]]'' File:Tetragonisca angustula nest entrance.jpg|''[[Tetragonisca angustula]]'' File:Nannotrigona testaceicornis nest entrance.jpg|''[[Nannotrigona testaceicornis]]'' </gallery> [[File:Entrant tubes.png|thumb|Variation of entrant tube in some native stingless bee species found in Southeast Asia: (1) ''[[Geniotrigona thoracica]]'', (2) ''[[Homotrigona fimbriata]]'', (3) ''[[Lophotrigona canifrons]]'', (4) ''[[Tetragonilla collina]]'', (5) ''[[Pariotrigona klossi]]'', (6) ''[[Heterotrigona itama]]'', (7) ''[[Tetragonula fuscobalteata]]'', (8) ''[[Lepidotriogona terminata]]'', (9) ''[[Tetrigona melanoleuca]]'', (10) ''[[Tetrigona apicalis]]'', and (11) ''[[Tetragonula pagdeni]]''.]] ==== Brood cell arrangement ==== Stingless bee colonies exhibit a diversity of construction patterns of brood cells, primarily composed of soft cerumen, a mixture of wax and resin. Each crafted cell is designed to rear a single individual bee, emphasizing the precision and efficiency of their nest architecture.<ref name=":15" /> The quantity of brood cells within a nest displays significant variation across different stingless bee species. Nest size can range from a few brood cells, as observed in the Asian ''[[Lisotrigona carpenteri]]'', to remarkably expansive colonies with over 80,000 brood cells, particularly in some American ''[[Trigona]]'' species.<ref name=":15" /> Meliponine colonies exhibit diverse brood cell arrangements, primarily categorized into three main types: horizontal combs, vertical combs, and clustered cells. Despite these primary types, variations and intermediate forms are prevalent, contributing to the flexibility of nest structures.<ref name=":15" /><ref>{{harvtxt|Villas-Bôas|2018|pp=27-29}}</ref> The first type involves horizontal combs, often characterized by a spiral pattern or layers of cells. The presence of spirals may not be consistent within a species, varying among colonies or even within the same colony. Some species, such as ''[[Melipona]]'', ''[[Plebeia]]'', ''Plebeina'', ''[[Nannotrigona]]'', ''Trigona'', and ''Tetragona'', may occasionally build spirals alongside other comb structures, as observed in ''[[Oxytrigona mellicolor]]''. As space diminishes for upward construction, workers initiate the creation of a new comb at the bottom of the brood chamber. This innovative approach optimizes the available space when emerging bees vacate older, lower brood combs.<ref name=":15" />[[image:Sarang lebah kelulut.jpeg|thumb|250px|left|Hive box containing colony of ''[[Heterotrigona itama]]'']] The second prevalent brood cell arrangement involves clusters of cells held together with thin cerumen connections. This clustered style is observed in various distantly related genera, such as the American ''[[Trigonisca]]'', ''[[Frieseomelitta]]'', ''[[Leurotrigona]]'', the Australian ''[[Austroplebeia]]'', and the African ''[[Hypotrigona]]''. This arrangement is particularly useful for colonies in irregular cavities unsuitable for traditional comb building.<ref name=":15" /> The construction of vertical combs is a distinctive trait found in only two stingless bee species: the African ''[[Dactylurina]]'' and the American ''[[Scaura longula]]''. This vertical arrangement sets these species apart from the more commonly observed horizontal comb structures in other stingless bee genera.<ref name=":15" /> ==== Brood rearing ==== Stingless bee brood rearing is a sophisticated and intricately coordinated process involving various tasks performed by worker bees, closely synchronized with the queen's activities. The sequence begins with the completion of a new brood cell, marking the initiation of mass provisioning.<ref name=":16">{{harvtxt|Grüter|2020|p=161}}</ref> Upon finishing a brood cell, several workers engage in mass provisioning, regurgitating larval food into the cell. This collective effort is swiftly followed by the queen laying her egg on top of the provided larval food. The immediate sealing of the cell ensues shortly afterward, culminating this important phase of the brood rearing process.<ref name=":16" /> The practice of mass provisioning, oviposition, and cell sealing is considered an ancestral trait, shared with solitary wasps and bees. However, in the context of stingless bees, these actions represent distinct stages of a highly integrated social process. Notably, the queen plays a central role in orchestrating these activities, acting as a pacemaker for the entire colony.<ref name=":16" /> This process diverges significantly from brood rearing in ''Apis'' spp. In honeybee colonies, queens lay eggs into reusable empty cells, which are then progressively provisioned over several days before final sealing. The contrasting approaches in brood rearing highlight the unique social dynamics and adaptations within stingless bee colonies.<ref name=":16" /> === Swarming === {{See also|Swarming (honey bee)}} Stingless bees and honey bees, despite encountering a common challenge in establishing daughter colonies, employ contrasting strategies. There are three key differences: reproductive status and age of the queen that leaves the nest, temporal aspects of colony foundation, and communication processes for nest site selection.<ref name=":17">{{harvtxt|Grüter|2020|pp=131-152}}</ref> In HB (''Apis mellifera''), the mother queen, accompanied by a swarm of numerous workers, embarks on relocation to a new home once replacement queens have been reared. Conversely, in SB (meliponines), the departure is orchestrated by the unmated ("virgin") queen, leaving the mother queen in the original nest. Mated stingless bee cannot leave the hive due to damaged wings and increased abdominal size post-mating ([[physogastrism]]). The queen's weight in species like ''[[Scaptotrigona postica]]'' increases, for example, about 250%.<ref name=":17" /><ref>{{harvtxt|Engels|1987}}</ref> Unlike honey bees, stingless bee colonies are unable to perform absconding - a term denoting the abandonment of the nest and migration to a new location - making them reliant on alternative strategies to cope with challenges. Meliponines progressive found new colonies without abandonning their nest abruptly.<ref name=":17" /> These are the stages of stingless bees swarming:<ref name=":17" /> # Reconnaissance and preparation: Scouts inspect potential new nest sites for suitability, considering factors such as cavity size, entrance characteristics, and potential threats. The criteria for determining suitability remain largely unexplored. Some colonies engage in simultaneous preparation of multiple cavities before making a final decision and some others make the initial reconnaissance but do not move into the cavity; # Transport of building material and food: workers seal cracks in the chosen cavity using materials like resin, batumen, or mud. They construct an entrance tube, possibly serving as a visual beacon for nestmate workers. Early food pots are built and filled with honey, requiring a growing number of workers to transport cerumen and honey from the mother nest. # Progressive establishment and social link: the mother and daughter colony maintain a social link through workers traveling between the two nests. The duration of this link varies among species, ranging from a few days to several months. Stingless bee colonies display a preference for cavities previously used by other colonies, containing remnants of building material and nest structures. # Arrival of the queen: after initial preparations, an unmated queen, accompanied by additional workers, arrives at the new nest site. # Drone arrival: males (drones) aggregate outside the newly established nest. They often arrive shortly after swarming initiation, even before the completion of nest structures. Males can be observed near the entrance, awaiting further events. # Mating flight: males in aggregations do not enter the colony but await the queen's emergence for a mating flight. Although rarely observed, it is assumed that unmated stingless bee queens embark on a single mating flight, utilizing acquired sperm for the entirety of their reproductive life. === Natural enemies === In meliponiculture, beekeepers need to be aware of the presence of animals that can harm stingless bee colonies. There are several potential enemies, but the most damaging ones to meliponaries are listed below.<ref>{{harvtxt|Nogueira-Neto|1997|pp=367-390}}</ref> ==== Invertebrates ==== [[File:Forídeo e armadilha.png|thumb|Phorid fly and vinegar trap.]] [[Phoridae|Phorid]] flies in the genus ''[[Pseudohypocera]]'' pose a significant threat to stingless bee colonies, causing problems for beekeepers. These parasites lay eggs in open cells of pollen and honey, leading to potential extinction if not addressed. Early detection is crucial for manual removal or using [[vinegar]] traps. It's important never to leave an infested box unattended to prevent the cycle from restarting and avoid contaminating other colonies. Careful handling of food jars, especially during swarms transfers, is essential. Prompt removal of broken jars, sealing gaps with wax or tape, and maintaining vigilance during the rainy season for heightened phorid activity are recommended. Combatting these flies usually is a priority, particularly during increased reproductive periods.<ref name=":72">Embrapa. ''[https://www.embrapa.br/documents/1355163/39571288/Roteiro+Teórica+-+Cuidados+Especiais.pdf/4fd9e18e-34ea-638c-250b-73c2241e6f07 Inimigos Naturais & Cuidados Especiais].'' Curso Básico de Abelhas Sem Ferrão.</ref><ref name=":82">{{harvtxt|Villas-Bôas|2018|p=|pp=103-104}}</ref><ref>{{harvtxt|Grüter|2020|pp=235-236}}</ref> [[Termite]]s usually do not attack bees or their food pots. However, they can cause damage to the structure of hive boxes as there are many [[Xylophagy|xylophagous]] species. While termites do not usually pose major problems for beekeepers, they should still be monitored closely.<ref>{{harvtxt|Nogueira-Neto|1997|pp=368-370}}</ref><ref>{{harvtxt|Grüter|2020|p=236}}</ref>[[File:Hermetia illucens Black soldier fly edit1.jpg|thumb|[[Hermetia illucens|Black soldier fly.]]]] [[File:Caixa racional individual.png|thumb|166x166px|Rational box support with burnt oil to prevent ant attacks.]] [[Ant]]s are attracted to bee colonies by the smell of food. To prevent ant attacks, it's important to handle the hive boxes carefully and avoid exposing jars of pollen and honey. Although rare, when attacks do occur, there are intense conflicts between ants and bees. Stingless bees usually manage to defend themselves, but the damage to the bee population can be significant. To prevent ant infestations in meliponaries with individual supports, a useful strategy is to impregnate the box supports with burnt oil.<ref name=":92">{{harvtxt|Villas-Bôas|2018|p=106}}</ref><ref>{{harvtxt|Grüter|2020|p=235}}</ref> Another group of enemy flies are the [[Hermetia illucens|black soldier flies]] (''Hermetia illucens''). They lay their eggs in crevices of boxes and can extend the tip of their abdomen during laying, facilitating access to the inside of the hive. Larvae of this species feed on pollen, feces, and other materials found in colonies. In general, healthy bee colonies can coexist peacefully with soldier flies. However, in areas where these insects are prevalent, beekeepers must remain vigilant and protect the gaps in the colonies to prevent potential issues.<ref>{{harvtxt|Villas-Bôas|2018|p=105}}</ref> [[File:Entrada de ninho de Lestrimelitta tropica.jpg|left|thumb|''Lestrimelitta limao'' hive entrance.]] Cleptobiosis, also known as [[Kleptoparasitism|cleptoparasitism]], is a behaviour observed in various species of stingless bees, with over 30 identified species engaging in nest attacks, including honey bee nests. This behaviour serves the purpose of either resource theft or usurping the nest by swarming into an already occupied cavity and these bees are called robber bees. The Neotropical genus ''[[Lestrimelitta]]'' and the African genus ''[[Cleptotrigona]]'' represent bees with an obligate cleptobiotic lifestyle since they do not visit flowers for nectar or pollen. Furthermore, other species such as ''[[Melipona fuliginosa]]'', ''[[Oxytrigona tataira]],'' ''[[Trigona hyalinata]]'', ''[[Trigona spinipes|T. spinipes]]'', and ''[[Tetragona clavipes]]'' are reported to have comparable habits of pillaging and invading, which emphasises the variety of strategies employed by stingless bees in acquiring resources. Other enemies include: jumping spiders ([[Salticidae]]), moths, assassin bugs ([[Reduviidae]]), beetles, parasitoid wasps, predatory mites (''[[Amblyseius]]''), mantises ([[Mantodea]]), robber flies ([[Asilidae]]), etc.<ref>{{harvtxt|Grüter|2020|pp=236-238}}</ref> ==== Vertebrates ==== Human activities pose the most significant threat to stingless bees, whether through honey and nest removal, [[habitat destruction]], [[pesticide]] use or introduction of non-native competitors. Large-scale environmental alterations, particularly the conversion of natural habitats into urban or intensively [[Agriculture|farmed land]], are the most dramatic threats leading to habitat loss, reduced nest densities, and [[Extinction|species disappearance]].<ref name=":18">{{harvtxt|Grüter|2020|pp=238-239}}</ref> [[Primate]]s, including [[chimpanzee]]s, [[gorilla]]s, [[baboon]]s, and various monkey species, are known to threaten stingless bee colonies. [[Elephant]]s, [[honey badger]]s, [[sun bear]]s, [[spectacled bear]]s, [[anteater]]s, [[hog-nosed skunk]]s, [[armadillo]]s, [[tayra]]s, [[eyra cat]]s, [[kinkajou]]s, [[grisons]], and [[coyote]]s are among the mammals that consume or destroy stingless bee nests. Some, like the tayra and eyra cat, have specific preferences for stealing honey. [[Gecko]]s, [[lizard]]s, and [[toad]]s also pose threats by hunting adult bees or consuming workers at nest entrances. [[Woodpecker]]s and various bird species, including [[bee-eater]]s, [[woodcreeper]]s, [[drongo]]s, [[jacamar]]s, [[heron]]s, [[kingbird]]s, [[Flycatcher-shrike|flycatchers]], [[Swift (bird)|swifts]], and [[honeyeater]]s, occasionally prey on stingless bees. African [[honeyguide]]s have developed a mutualism with human honey-hunters, actively guiding them to bee nests for honey extraction and then consuming leftover wax and larvae.<ref name=":18" /> ==== Defense ==== Being [[tropics|tropical]], stingless bees are active all year round, although they are less active in cooler weather, with some species presenting [[diapause]].<ref name=":19">{{harvtxt|Grüter|2020|pp=248-260}}</ref><ref>{{harvtxt|Ribeiro|2002}}</ref><ref>{{harvtxt|Alves|Imperatriz-Fonseca|Santos-Filho|2009}}</ref> Unlike other eusocial bees, they do not sting, but will defend by biting if their nest is disturbed. In addition, a few (in the genus ''[[Oxytrigona]]'') have mandibular secretions, including [[formic acid]], that cause painful blisters.<ref name=":3" /><ref>{{harvtxt|Roubik|Smith|Carlson|1987}}</ref> Despite their lack of a [[sting (biology)|sting]], stingless bees, being eusocial, may have very large colonies made formidable by the number of defenders.<ref>{{harvtxt|Roubik|2006}}</ref><ref>{{harvtxt|Sarchet|2014}}</ref> Stingless bees use other sophisticated defence tactics to protect their colonies and ensure their survival. One important strategy is to choose nesting habitats with fewer natural enemies to reduce the risk of attacks. In addition, they use [[camouflage]] and [[mimicry]] to blend into their surroundings or imitate other animals to avoid detection. An effective strategy is to nest near colonies that provide protection, using collective strength to defend against potential invaders.<ref name=":19" /> Nest entrance guards play a vital role in colony defense by actively preventing unauthorized entry through attacking intruders and releasing alarm [[pheromone]]s to recruit additional defenders. It is worth noting that nest guards often carry sticky substances, such as resins and wax, in their corbiculae or mandibles. Stingless bees apply substances to attackers to immobilise them, thus thwarting potential threats to the colony. Some species (''Tetragonisca angustula'' and ''Nannotrigona testaceicornis'', for example) also close their nest entrances with a soft and porous layer of cerumen at night, further enhancing colony security during vulnerable periods. These intricate defence mechanisms demonstrate the adaptability and resilience of stingless bees in safeguarding their nests and resources.<ref name=":19" /> === Role differentiation === In a simplified sense, the sex of each bee depends on the number of [[chromosome]]s it receives. Female bees have two sets of chromosomes ([[diploid]])—one set from the [[queen (bee)|queen]] and another from one of the male bees or [[drone (bee)|drone]]s. Drones have only one set of chromosomes ([[haploid]]), and are the result of unfertilized eggs, though inbreeding can result in diploid drones. Unlike true honey bees, whose female bees may become workers or queens strictly depending on what kind of food they receive as larvae (queens are fed [[royal jelly]] and workers are fed pollen), the caste system in meliponines is variable, and commonly based simply on the amount of pollen consumed; larger amounts of pollen yield queens in the genus ''[[Melipona]]''. Also, a genetic component occurs, however, and as much as 25%<ref>{{harvtxt|Kerr|1950}}</ref> (typically 5–14%) of the female brood may be queens. Queen cells in the former case can be distinguished from others by their larger size, as they are stocked with more pollen, but in the latter case, the cells are identical to worker cells, and scattered among the worker brood. When the new queens emerge, they typically leave to mate, and most die.<ref name=":28">{{harvtxt |Wenseleers|Ratnieks|Ribeiro|Alves|Imperatriz-Fonseca|2005}}</ref> New nests are not established by swarms, but by a procession of workers that gradually construct a new nest at a secondary location. The nest is then joined by a newly mated queen, at which point many workers take up permanent residence and help the new queen raise her own workers. If a ruling queen is herself weak or dying, then a new queen can replace her. For ''[[Plebeia quadripunctata|Schwarziana quadripunctata]]'', although fewer than 1% of female worker cells produce dwarf queens, they comprise six of seven queen bees, and one of five proceed to head colonies of their own. They are reproductively active, but less fecund than large queens.<ref name=":28" /> == Interaction with humans == === Pollination === {{Main|Pollination}} [[File:Bee pollinating a rose.jpg|thumb|Solitary bee visiting and pollinating a flower.]] Bees play a critical role in the ecosystem, particularly in the pollination of natural vegetation. This activity is essential for the reproduction of various plant species, particularly in [[tropical forest]]s where most tree species rely on [[Entomophily|insect pollination]]. Even in temperate climates, where [[Anemophily|wind pollination]] is prevalent among forest trees, many bushes and herbaceous plants, rely on bees for pollination. The significance of bees extends to arid regions, such as [[Deserts and xeric shrublands|desertic and xeric shrublands]], where bee-pollinated plants are essential for preventing [[erosion]], supporting [[wildlife]], and ensuring [[Ecological stability|ecosystem stability]].<ref name=":23">{{harvtxt|Michener|2000|pp=4-5}}</ref> The impact of bee pollination on agriculture is substantial. In the late 1980s, certain plants were estimated to contribute between $4.6 to $18.9 billion to the U.S. economy, primarily through insect-pollinated crops. Although some bee-pollinated plants can self-pollinate in the absence of bees, the resulting crops often suffer from inbreeding depression. The quality and quantity of seeds or fruits are significantly enhanced when bees participate in the pollination process. Although estimates of crop pollination attributed to honey bees are uncertain, it is undeniable that bee pollination is a vital and economically valuable activity.<ref name=":23" /> Ramalho (2004) demonstrates that stingless bees amount to approximately 70% of all bees foraging on flowers in the Brazilian Tropical [[Atlantic Forest|Atlantic Rainforest]] even though they represented only 7% of all bee species.<ref>{{harvtxt|Ramalho|2004}}</ref> In a habitat in Costa Rica, stingless bees accounted for 50% of the observed foraging bees, despite representing only 16% of the recorded bee species.<ref>{{harvtxt|Brosi|Daily|Shih|Oviedo|Durán|2008}}</ref> Following this pattern, Cairns ''et al''. (2005) found that 52% of all bees visiting flowers in Mexican habitats were meliponines.<ref>{{harvtxt|Cairns|Villanueva-Gutiérrez|Koptur|Bray|2005}}</ref> Meliponine bees play a crucial role in tropical environments due to their high population rate, morphological diversity, diverse foraging strategies, generalist foraging habits ([[Polylectic|polylecty]]), and flower constancy during foraging trips. Nest densities and colony sizes can result in over a million individual stingless bees inhabiting a square kilometre of tropical habitat. Due to their diverse morphology and behaviour, bees are capable of collecting pollen and nectar from a wide range of flowering plants. Key plant families are reported as most visited by meliponines: [[Fabaceae]], [[Euphorbiaceae]], [[Asteraceae]] and [[Myrtaceae]].<ref name=":20">{{harvtxt|Grüter|2020|pp=323-330}}</ref> Grüter compiled some studies about twenty crops that substantially benefit from SB pollination (following table) and also lists seventy-four crops that are at least occasionally or potentially pollinated by stingless bees.<ref name=":20" /> {| class="wikitable mw-collapsible" |+ !Common name !Scientific name !Family !Pollinator genus !Reference |- |Annatto, achiote |''[[Bixa orellana]]'' |[[Bixaceae]] |''[[Melipona]]'' |<ref name=":21">{{harvtxt|Heard|1999}}</ref> |- |Aubergine |''[[Solanum melongena|Solanum melogena]]'' |[[Solanaceae]] |''Melipona'' |<ref>{{harvtxt|Nunes-Silva|Hrncir|da Silva|Roldão|Imperatriz-Fonseca|2013}}</ref> |- |Avocado |''[[Persea americana]]'' |[[Lauraceae]] |''[[Nannotrigona]], [[Trigona]]'' |<ref name=":21" /> |- |Camu-camu |''[[Myrciaria dubia]]'' |[[Myrtaceae]] |''Melipona, [[Scaptotrigona]]'' |<ref name=":21" /> |- |Carambola |''[[Averrhoa carambola]]'' |[[Oxalidaceae]] |''Trigona'' |<ref name=":21" /> |- |Chayote, choko |''[[Sechium edule]]'' |[[Cucurbitaceae|Cucubitaceae]] |''Trigona, [[Partamona]]'' |<ref name=":21" /> |- |Coconut |''[[Cocos nucifera]]'' |[[Arecaceae]] |various genera |<ref name=":21" /> |- |Coffee |''[[Coffea arabica]]'' |[[Rubiaceae]] |''[[Lepidotrigona]], Trigona'' |<ref name=":21" /><ref name=":22">{{harvtxt|Slaa|Chaves|Malagodi-Braga|Hofstede|2006}}</ref> |- |Coffee |''[[Coffea canephora]]'' |Rubiaceae |''Lepidotrigona, Trigona'' |<ref name=":22" /> |- |Cucumber |''[[Cucumis sativus]]'' |Curcubitaceae |''Nannotrigona, Scaptotrigona'' |<ref name=":21" /><ref name=":22" /> |- |Cupuaçu |''[[Theobroma grandiflorum]]'' |[[Malvaceae]] |''Trigona'' |<ref name=":21" /> |- |Macadamia |''[[Macadamia integrifolia]]'' |[[Proteaceae]] |''Trigona'' |<ref name=":21" /> |- |Mango |''[[Mangifera indica]]'' |[[Anacardiaceae]] |''Trigona'' |<ref name=":21" /> |- |Mapati, uvilla |''[[Pourouma cecropiifolia]]'' |[[Moraceae]] |''[[Oxytrigona]], Trigona'' |<ref name=":21" /> |- |Mealy sage |''[[Salvia farinacea]]'' |[[Lamiaceae]] |''Nannotrigona, [[Tetragonisca]]'' |<ref name=":22" /> |- |Rambutan |''[[Nephelium lappaceum]]'' |[[Sapindaceae]] |''Scaptotrigona'' |<ref name=":21" /><ref name=":22" /> |- |Rockmelon |''[[Cucumis melo]]'' |Curcubitaceae |''[[Heterotrigona]]'' |<ref>{{harvtxt|Azmi|Wan Sembok|Yusuf|Mohd Hatta|Salleh|Hamzah|Ramli|2019}}</ref> |- |Strawberry |''[[Fragaria]]'' sp. |[[Rosaceae]] |various genera |<ref name=":21" /><ref name=":22" /> |- |Sweet pepper |''[[Capsicum annuum]]'' |Solanaceae |''[[Austroplebeia]], Melipona, [[Tetragonula]]'' |<ref name=":21" /><ref name=":22" /><ref>{{harvtxt|Putra|Salmah|Swasti|2017}}</ref> |- |Tomato |''[[Lycopersicon esculentum|Solanum lycopersicum]]'' |Solanaceae |''Melipona, Nannotrigona'' |<ref name=":22" /> |} == Relationship to humans == === Australia === "''Coot-tha''", which derives from "''ku-ta''", is one of the Aboriginal names for "wild stingless bee honey".<ref>{{harvtxt |Vit|Pedro|Roubik|2018}}</ref> Of the 1,600 species of wild bees native to Australia, about 14 are meliponines.<ref>{{cite news | url=http://www.abc.net.au/science/articles/2003/05/08/2045526.htm | title=Stingless bee rescue |author= Wendy Pyper |date=May 8, 2003 |work=ABC Science}}</ref> These species bear a variety of names, including Australian native honey bees, native bees, sugar-bag bees, and sweat bees (because they land on people's skin to collect sweat).<ref>{{Cite web |title=Australian Native Stingless Bees |url=https://www.aussiebee.com.au/australian-stingless-bees.html |access-date=2024-05-28 |website=www.aussiebee.com.au}}</ref> All are small and usually black in colour, with hairy extended hind legs for carrying [[nectar]] and [[pollen]]; because of the latter, they are sometimes mistaken for [[bumblebee]]s. The various stingless species look quite similar, with the two most common species, ''[[Tetragonula carbonaria]]'' and ''[[Austroplebeia australis]]'', displaying the greatest variation, as the latter is smaller and less active. Both of these inhabit the area around [[Brisbane]].<ref>{{Cite web |last=Steve |date=2015-05-25 |title=Native Stingless Bees - Tetragonula carbonaria |url=https://www.nativebeehives.com/native-stingless-bees-tetragonula-carbonaria/ |access-date=2024-05-28 |website=www.nativebeehives.com |language=en-AU}}</ref> As stingless bees are usually harmless to humans, they have become an increasingly attractive addition to the suburban backyard. Most meliponine beekeepers do not keep the bees for [[honey]], but rather for the pleasure of conserving native species whose original [[habitat (ecology)|habitat]] is declining due to human development. In return, the bees [[pollinate]] crops, garden flowers, and bushland during their search for nectar and pollen. While a number of beekeepers fill a small niche market for bush honey, native meliponines only produce small amounts and the structure of their hives makes the honey difficult to extract. Only in warm areas of Australia such as [[Queensland]] and northern [[New South Wales]] are favorable for these bees to produce more honey than they need for their own survival. Most bees only come out of the hive when it is above about 18°C (64°F).<ref>{{Cite web|url=https://www.abc.net.au/news/2019-07-05/moves-underway-to-make-a-standard-for-native-bee-honey/11278346|title=Native bee honey set to have its own food standard|last=Thomas|first=Kerrin|date=2019-07-05|website=ABC News|language=en-AU|access-date=2019-07-10}}</ref> Harvesting honey from a nest in a cooler area could weaken or even kill the nest. ==== Pollination ==== Australian farmers rely almost exclusively on the introduced [[western honey bee]] to pollinate their crops. However, native bees may be better pollinators for certain agricultural crops. Stingless bees have been shown to be valuable pollinators of tropical plants such as [[macadamia]]s and [[mango]]s.<ref name=":21" /> Their foraging may also benefit strawberries, watermelons, citrus, avocados, [[lychee]]s, and many others.<ref name=":21" /><ref name=":22" /> Research into the use of stingless bees for crop pollination in Australia is still in its very early stages, but these bees show great potential. Studies at the [[University of Western Sydney]] have shown these bees are effective pollinators even in confined areas, such as glasshouses.<ref>{{cite journal |date=May 1999 |title=New Greenhouse Pollination Study With ''Trigona'' |url=http://www.aussiebee.com.au/ab10.html |journal=Aussie Bee Bulletin |issue=10 |quote=Pablo Occhiuzzi of the University of Western Sydney is studying the greenhouse pollination of capsicum with ''Trigona carbonaria''.}}</ref> === Brazil === {{see also|List of stingless bees of Brazil}} [[File:Abelhas sem ferrao brasil.png|thumb|Number of valid species of stingless bees in Brazil. State and region cut-outs. Study published in 2023.<ref name=":27" />]] [[File:Pollinating bee covered with pollen.jpg|thumb|''[[Trigona spinipes]],'' covered with [[pollen]], visiting a flower of the vegetable sponge gourd (''Luffa cylindrica'') in [[Campinas]], [[Brazil]].]] Brazil is home to several species bees belonging to Meliponini, with more than 300 species already identified and probably more yet to be discovered and described. They vary greatly in shape, size, and habits, and 20 to 30 of these species have good potential as honey producers. Although they are still quite unknown by most people, an increasing number of beekeepers (''meliponicultores'', in Portuguese) have been dedicated to these bees throughout the country.<ref name=":27" /><ref>{{harvtxt|Villas-Bôas|2018|p=17}}</ref> This activity has experienced significant growth since August 2004, when national laws were changed to allow native bee colonies to be freely marketed, which was previously forbidden in an unsuccessful attempt to protect these species. Nowadays the capture or destruction of existing colonies in nature is still forbidden, and only new colonies formed by the bees themselves in artificial traps can be collected from the wild.<ref>[https://conama.mma.gov.br/?option=com_sisconama&task=arquivo.download&id=440 CONAMA 2004 Resolution]</ref> Most colonies marketed are artificially produced by authorized beekeepers, through division of already existing captive colonies. Besides honey production, Brazilian stingless bees such as the jataí (''[[Tetragonisca angustula]]''), mandaguari (''[[Scaptotrigona postica]]''), and mandaçaia (''[[Melipona quadrifasciata]]'') serve as major pollinators of tropical plants and are considered the ecological equivalent of the honey bee.<ref name=":21" /><ref name=":22" /> Also, much practical and academic work is being done about the best ways of keeping such bees, multiplying their colonies, and exploring the honey they produce. Among many others, species such as jandaíra (''[[Melipona subnitida]]'') and true ''uruçu'' (''[[Melipona scutellaris]]'') in the northeast of the country, mandaçaia (''[[Melipona quadrifasciata]]'') and yellow uruçu (''Melipona rufiventris'') in the south-southeast, tiúba or jupará (''[[Melipona interrupta]]''<ref>{{Cite web|url=http://moure.cria.org.br/catalogue?id=34747|title=+Moure's Bee Catalogue}}</ref>) and straw-bee (''[[Scaptotrigona polysticta]]'') in the north and jataí (''[[Tetragonisca angustula]]'') throughout the country are increasingly kept by small, medium, and large producers. Many other species as the mandaguari (''[[Scaptotrigona postica]]''), the guaraipo (''[[Melipona bicolor]]''), marmelada (''[[Frieseomelitta varia]]'') and the iraí (''[[Nannotrigona testaceicornis]]''), to mention a few, are also reared. Through the cultivation of honey or selling of colonies, keeping stingless bees is an increasingly profitable activity. A single colony of species like mandaçaia and true "uruçu" can be divided up to four times a year, and each of the new colonies obtained this way can be sold for about US$100. According to [[Chico Mendes Institute for Biodiversity Conservation|ICMBio]] and the [[Ministry of Environment and Climate Change (Brazil)|Ministry of the Environment]] there are presently four species of Meliponini listed in the [[IUCN Red List|National Red List of Threatened Species]] in Brazil. ''[[Melipona capixaba]]'', ''[[Melipona rufiventris]]'', ''[[Melipona scutellaris]]'', and ''[[Partamona littoralis]]'' all listed as Endangered (EN).<ref>{{Cite web |title=IBAMA |url=https://www.ibama.gov.br/component/legislacao/?view=legislacao&force=1&legislacao=134521#:~:text=Ementa:,Extin%C3%A7%C3%A3o%20-%20Peixes%20e%20Invertebrados%20Aqu%C3%A1ticos. |access-date=2024-02-26 |website=www.ibama.gov.br}}</ref><ref>[[Instituto Chico Mendes de Conservação da Biodiversidade|ICMBio]]. 2018. [https://www.gov.br/icmbio/pt-br/centrais-de-conteudo/publicacoes/publicacoes-diversas/livro_vermelho_2018_vol7.pdf Livro Vermelho da Fauna Brasileira Ameaçada de Extinção: Volume VII - Invertebrados]. ''In'': ICMBio. (Org.). [https://www.gov.br/icmbio/pt-br/centrais-de-conteudo/publicacoes/publicacoes-diversas/livro_vermelho_2018_vol1.pdf Livro Vermelho da Fauna Brasileira Ameaçada de Extinção]. Brasília: ICMBio. 727p.</ref> ==== Honey production ==== Although the colony population of most of these bees is much smaller than that of European bees, the productivity per bee can be quite high. Probably the world champions in honey productivity, the manduri (''[[Melipona marginata]]'') and the ''guaraipo'' (''M. bicolor''), live in swarms of only around 300 individuals, but can still produce up to 5 liters (.79 US gallon) of honey a year under the right conditions. The manduri is one of the smallest bees in the genus ''[[Melipona]]'', varying in length from 6 to 7 mm (15/64" to 9/32").<ref>{{harvtxt|Witter|Blochtein|Santos|2007|p=27}}</ref> Species of the genus ''Scaptotrigona'' have very large colonies, with up to 20,000 individuals, and can produce from 3 to 8 liters (2 US gallons) of honey a year. Some large breeders have more than 3,000 hives of the tamer but still highly productive species in the genus ''Melipona'', such as the tiúba, the true ''uruçu'', and the jandaíra, each with 3,000 or more bees per colony. They can produce over 1.5 tons (3,000&nbsp;lbs) of honey every year.{{Citation needed|date=June 2024}} In large bee farms, only the availability of flowers limits the honey production per colony. Their honey is considered more palatable because it's not overly sweet, and it also is thought to have medicinal properties more pronounced than honey from bees of the genus ''Apis'' due to the higher level of antimicrobial substances. As a result, the honey from stingless bees returns very high profits, with prices greater than those for the more common honey produced by European bees. However, much larger numbers of beehives are required to produce amounts of honey comparable to that of European bees. Also, due to the fact of those bees storing honey in cerumen pots instead of standardized honeycombs as in the honeybee rearing makes extraction a lot more difficult and laborious.<ref>{{harvtxt|Fonseca|Sodré|Carvalho|Alves|Souza|Silva|Oliveira|Machado|Clarton|2007}}</ref> The honey from stingless bees has a lighter color and a higher water content, from 25% to 35%, compared to the honey from the genus ''Apis''. This contributes to its less cloying taste but also causes it to spoil more easily. Thus, for marketing, this honey needs to be processed through desiccation, fermentation or pasteurization. In its natural state, it should be kept under refrigeration.<ref>{{harvtxt|Grüter|2020|pp=12-14}}</ref> ==== Bees as pets ==== [[Image:Mandaçaias.jpg|left|thumb|A swarm of mandaçaias bees around an artificial [[beehive]] installed in a house's backyard in [[Brazil]]]] Due to the lack of a functional stinger and characteristic nonaggressive behavior of many Brazilian species of stingless bees, they can be reared without problems in densely populated environments such as cities, provided enough flowers are at their disposal nearby. Some breeders (''meliponicultores'') can produce honey even in apartments up to the 12th floor.<ref>{{cite web|url=http://meliponariocapixaba.blogspot.com.br/2009/11/e-possivel-criar-abelhas-em.html|title=Meliponario Capixaba: É POSSÍVEL CRIAR ABELHAS EM APARTAMENTOS?|author=João Luiz|access-date=12 March 2016}}</ref> The [[Melipona quadrifasciata|mandaçaias]] (''Melipona quadrifasciata'') are extremely tame, rarely attacking humans (only when their hives are opened for honey extraction or colony division). They form small, manageable colonies of only 400&ndash;600&nbsp;individuals. At the same time, a single rational beehive of ''mandaçaia'' can produce up to 4&nbsp;liters (1&nbsp;US gallon) of honey a year, making the species very attractive for home keepers. They are fairly large bees, up to 11&nbsp;mm (7/16") in length, and as a result have better body heat control, allowing them to live in regions where temperatures can drop a little lower than 0&nbsp;°C (32&nbsp;°F). However, they are somewhat selective about which flowers they will visit, preferring the flora that occurs in their natural environment. They are thus difficult to keep outside their region of origin (the northeast coast of Brazil).<ref>{{harvtxt|Brito|Faquinello|Paula-Leite|Carvalho|1997}}</ref> Other species, like the ''tiúba'' and the true ''uruçu'', are also very tame and highly productive. Their colonies have from 3,000&ndash;5,000 individuals (for comparison, the population of honeybee swarms can peak at 80,000&nbsp;individuals) and can produce up to 10 liters (2.6 US gallons) of honey a year.{{Citation needed|date=June 2024}} They can be easily kept at home but will survive only in regions with a warm climate, their larvae dying at temperatures lower than 12&nbsp;°C (54&nbsp;°F). The yellow ''uruçu'', however, can survive at temperatures lower than 0&nbsp;°C (32&nbsp;°F), and their colonies, bearing about 3,500&nbsp;individuals, can produce up to 6&nbsp;liters (1.5&nbsp;US gallons) of honey a year.{{Citation needed|date=June 2024}} But this species will react with powerful nipping if its nests are disturbed, and usually they are only kept by professional meliponicultors. Another suitable species for keeping at home is the ''guaraipo'' (''[[Melipona bicolor]]''). It is also quite tame, never attacking the beekeeper, and their colonies have fewer than 600 individuals. They can withstand temperatures as low as &minus;10&nbsp;°C (14&nbsp;°F), and each colony can produce over 3&nbsp;liters ({{frac|3|4}}&nbsp;US gallons) of honey a year.{{Citation needed|date=June 2024}} Their colonies usually have more than one single queen at a time (usually two or three, but sometimes up to five), a phenomenon called [[Gyne|polygyne]], and thus are less sensitive to the death of one queen, which can cause the loss of a whole colony in other species. But the ''guaraipo'' is very sensitive to low levels of humidity, and their hives must be equipped with means to keep a high moisture content. Once very common, the ''guaraipo'' is now rather rare in nature, mainly due to the destruction of their native forests in the south-southeast of Brazil. Other groups of Brazilian stingless bees, genera ''[[Plebeia]]'' and ''[[Leurotrigona]]'', are also very tame and much smaller, with one of them (''[[Plebeia minima]]'') reaching no more than 2.5&nbsp;mm (3/32") in length, and the ''[[lambe-olhos]]'' ("lick-eyes" bee, ''[[Leurotrigona muelleri]]'') being even smaller, at no more than 1.5&nbsp;mm (3/32"). Many of these species are known as ''mirim'' (meaning 'small' in the [[Tupi-Guarani languages]]). As a result, they can be kept in very small artificial hives, thus being of interest for keepers who want them as pollinators in small glasshouses or just for the pleasure of having a 'toy' bee colony at home. Being so tiny, these species produce only a very small amount of honey, typically less than 500&nbsp;ml (1/2 US pint) a year, so are not interesting for commercial honey production. [[Image:Abelha-jatai.jpg|right|thumb|Entrance pipe of ''jataís´s'' colony: The ''jataí'' bees build a wax pipe in the entrance of their nest.]] Belonging to the same group, the ''jataí'' (''[[Tetragonisca angustula]]''), the ''marmelada ''(''[[Frieseomelitta varia]]''), and the ''moça-branca'' (''[[Frieseomelitta doederleini]]'') are intermediate in size between those very small species and the European bee. They are very adaptable species; the ''jataí'', and can be reared in many different regions and environments, being quite common in most Brazilian cities. The ''jataí'' can bite when disturbed, but its jaws are weak, and in practice they are harmless, while the ''marmelada'' and ''moça-branca'' usually deposit propolis on their aggressors. Producing up to 1.5&nbsp;liters (0.4&nbsp;US gallons) of honey a year, their honey is considered among the best from stingless bees. In fact, the ''jataí'' was one of the first species to be kept by home beekeepers. Their nests can be easily identified in trees or wall cavities by the yellow wax pipe they build at the entrance, usually guarded by some soldier bees, which are stronger than regular worker bees. The'' marmelada'' and ''moça-branca'' make a little less honey, but it is denser and sweeter than most from other stingless bees and is considered very tasty. === Maya stingless bees of Central America === [[Image:Colmena melipona.jpg|right|thumb|A Maya stingless bee hive: A piece of hollow log provides a home for meliponine bees in [[Belize]].]] The stingless bees ''[[Melipona beecheii]]'' and ''[[Melipona yucatanica|M. yucatanica]]'' are the primary native bees cultured in Central America, though a few other species are reported as being occasionally managed (e.g., ''[[Trigona fulviventris]]'' and ''[[Scaptotrigona mexicana]]'').<ref>{{harvtxt|Kent|1984}}</ref> They were extensively cultured by the [[Maya civilization]] for honey, and regarded as sacred. They continue to be cultivated by the modern [[Maya peoples]], although these bees are endangered due to massive deforestation, altered [[agriculture|agricultural]] practices (especially overuse of [[insecticide]]s), and changing beekeeping practices with the arrival of the [[Africanized honey bee]], which produces much greater honey crops.<ref name=":26">{{harvtxt|Villanueva|Roubik|Colli-Ucán|1998}}</ref> ==== History ==== Native meliponines (''M. beecheii'' being the favorite) have been kept by the lowland Maya for thousands of years. The [[Yucatec Maya language]] name for this bee is ''xunan kab'', meaning "(royal, noble) lady bee".<ref>{{cite web|title=Diccionario Introductorio|url=http://www.uqroo.mx/libros/maya/diccionario.pdf|work=uqroo.mx|publisher=Universidad De Quintana Roo|access-date=27 March 2014|language=es}}</ref> The bees were once the subject of religious ceremonies and were a symbol of the bee-god [[Ah-Muzen-Cab]], known from the ''[[Madrid Codex (Maya)|Madrid Codex]]''. The bees were, and still are, treated as pets. Families would have one or many log-hives hanging in and around their houses. Although they are stingless, the bees do bite and can leave welts similar to a mosquito bite. The traditional way to gather bees, still favored among the locals, is find a wild hive, then the branch is cut around the hive to create a portable log, enclosing the colony. This log is then capped on both ends with another piece of wood or pottery and sealed with mud. This clever method keeps the melipine bees from mixing their brood, pollen, and honey in the same comb as do the European bees. The brood is kept in the middle of the hive, and the honey is stored in vertical "pots" on the outer edges of the hive. A temporary, replaceable cap at the end of the log allows for easy access to the honey while doing minimal damage to the hive. However, inexperienced handlers can still do irreversible damage to a hive, causing the hive to swarm and abscond from the log. With proper maintenance, though, hives have been recorded as lasting over 80 years, being passed down through generations. In the archaeological record of Mesoamerica, stone discs have been found that are generally considered to be the caps of long-disintegrated logs that once housed the beehives. ==== Tulum ==== [[Tulum]], the site of a [[pre-Columbian]] Maya city on the Caribbean coast 130&nbsp;km (81&nbsp;mi) south of Cancun, has a god depicted repeatedly all over the site. Upside down, he appears as a small figure over many doorways and entrances. One of the temples, the "Templo del Dios Descendente" or the Temple of the Descending God, stands just left of the central plaza. Speculation is that he may be the "Bee God", Ah Muzen Cab, as seen in the Madrid Codex. It is possible that this was a religious/trade center with emphasis on ''xunan kab'', the "royal lady". ==== Economic uses ==== [[Balché]], a traditional [[Mesoamerica]]n alcoholic beverage similar to [[mead]], was made from fermented honey and the bark of the leguminous'' balché'' tree (''[[Lonchocarpus violaceus]]''), hence its name.<ref>{{harvtxt|Ott|1998}}</ref> It was traditionally brewed in a canoe. The drink was known to have [[entheogen]]ic properties, that is, to produce mystical experiences, and was consumed in medicinal and ritual practices. Beekeepers would place the nests near the psychoactive plant ''[[Turbina corymbosa]]'' and possibly near ''balché'' trees, forcing the bees to use nectar from these plants to make their honey.<ref>{{harvtxt|Ott|1998}}</ref> Additionally, brewers would add extracts of the bark of the ''balché'' tree to the honey mixture before fermentation. The resulting beverage is responsible for psychotropic effects when consumed, due to the ergoline compounds in the pollen of the ''T. corymbosa'', the ''Melipona'' nectar gathered from the ''balché'' flowers, or the hallucinogenic compounds of the ''balché'' tree bark. [[Lost-wax casting]], a common metalworking method typically found where the inhabitants keep bees, was also used by the Maya. The wax from ''Melipona'' is soft and easy to work, especially in the humid Maya lowland. This allowed the Maya to create smaller works of art, jewelry, and other metalsmithing that would be difficult to forge. It also makes use of the leftovers from honey extraction. If the hive was damaged beyond repair, the whole of the comb could be used, thus using all of the hive. With experienced keepers, though, only the honey pot could be removed, the honey extracted, and the wax used for casting or other purposes. ==== Future ==== The outlook for meliponines in Mesoamerica is uncertain. The number of active Meliponini beekeepers is shy in comparison with the Africanized ''[[Apis mellifera]]'' breeders. The high honey yield, 100&nbsp;kg (220&nbsp;lbs) or more annually, along with the ease of hive care and ability to create new hives from existing stock, commonly outweighs the negative consequences of "killer bee" hive maintenance. Further complicating the issue, [[Africanized honey bee]]s do not visit some flora, such as those in the [[Solanaceae|tomato family]], and several forest trees and shrubs, which rely on the native stingless bees for pollination. A decline in populations of native flora has already occurred in areas where stingless bees have been displaced by Africanized honey bees. An additional blow to the art of meliponine beekeeping is that many of the meliponine beekeepers are now elderly, and their hives may not be cared for once they die. The hives are considered similar to an old family collection, to be parted out once the collector dies or to be buried in whole or part along with the beekeeper upon death. In fact, a survey of a once-popular area of the Maya lowlands shows the rapid decline of beekeepers, down to around 70 in 2004 from thousands in the late 1980s. It is traditional in the Maya lowlands that the hive itself or parts of the hive be buried along with the beekeeper to ''volar al cielo'', "to fly to heaven".{{citation needed|date=March 2007}} Conservation efforts are underway in several parts of Mesoamerica.<ref>A comprehensive conservation guide can be found in the June 2005 issue of ''Bee World''.</ref> == References ==<!-- ActaAmazonica34 p333 --> {{Reflist|2}} ==Bibliography== === Articles and publications === * {{cite journal |last1=Alves |first1=D A |last2=Imperatriz-Fonseca |first2=V L |last3=Santos-Filho |first3=P S. |year=2009 |title=Production of workers, queens and males in ''Plebeia remota'' colonies (Hymenoptera, Apidae, Meliponini), a stingless bee with reproductive diapause |journal=Genetics and Molecular Research |volume=8 |issue=2 |pages=672–683 |doi=10.4238/vol8-2kerr030 |pmid=19554766 |doi-access=free}} * {{Cite journal |last1=Antonini |first1=Yasmine |last2=Martins |first2=Rogério P. |date=2003 |title=The value of a tree species (''Caryocar brasiliense'') for a stingless bee ''Melipona quadrifasciata quadrifasciata'' |url=http://link.springer.com/10.1023/A:1027378306119 |journal=Journal of Insect Conservation |volume=7 |issue=3 |pages=167–174 |doi=10.1023/A:1027378306119|bibcode=2003JICon...7..167A |s2cid=6080884 }} * {{Cite journal |last1=Azmi |first1=Wahizatul Afzan |last2=Wan Sembok |first2=W Z |last3=Yusuf |first3=N |last4=Mohd. 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W |last2=Smith |first2=B. H |last3=Carlson |first3=R. G |year=1987 |title=Formic acid in caustic cephalic secretions of stingless bee, ''Oxytrigona'' (Hymenoptera: Apidae) |journal=Journal of Chemical Ecology |volume=13 |issue=5 |pages=1079–86 |doi=10.1007/BF01020539 |pmid=24302133 |bibcode=1987JCEco..13.1079R |s2cid=30511107}} * {{cite journal |last1=Roubik |first1=D W. |year=2006 |title=Stingless bee nesting biology |journal=Apidologie |volume=37 |issue=2 |pages=124–143 |doi=10.1051/apido:2006026 |doi-access=free}} * {{Cite web |last=Sarchet |first=Penny |date=14 November 2014 |title=Zoologger: Stingless suicidal bees bite until they die |url=https://www.newscientist.com/article/dn26562-zoologger-stingless-suicidal-bees-bite-until-they-die/ |access-date=2020-01-24 |website=New Scientist |language=en-US}} * {{Cite journal |last1=Slaa |first1=Ester Judith |last2=Chaves |first2=Luis Alejandro Sánchez |last3=Malagodi-Braga |first3=Katia Sampaio |last4=Hofstede |first4=Frouke Elisabeth |date=2006-03-01 |title=Stingless bees in applied pollination: practice and perspectives |url=http://dx.doi.org/10.1051/apido:2006022 |journal=Apidologie |language=en |volume=37 |issue=2 |pages=293–315 |doi=10.1051/apido:2006022 |s2cid=55280074 |issn=0044-8435}} * {{Cite journal |last1=Sousa |first1=Janaína Maria Batista de |last2=Souza |first2=Evandro Leite de |last3=Marques |first3=Gilmardes |last4=Benassi |first4=Marta de Toledo |last5=Gullón |first5=Beatriz |last6=Pintado |first6=Maria Manuela |last7=Magnani |first7=Marciane |date=2016-01-01 |title=Sugar profile, physicochemical and sensory aspects of monofloral honeys produced by different stingless bee species in Brazilian semi-arid region |journal=LWT - Food Science and Technology |volume=65 |pages=645–651 |doi=10.1016/j.lwt.2015.08.058 |issn=0023-6438 |doi-access=free}} * {{cite journal |last1=Souza |first1=R C S |last2=Yuyama |first2=L K O |last3=Aguiar |first3=J P L |last4=Oliveira |first4=F P M. |year=2004 |title=Valor nutricional do mel e pólen de abelhas sem ferrão da região amazônica |journal=Acta Amazonica |volume=34 |issue=2 |pages=333–336 |doi=10.1590/s0044-59672004000200021 |doi-access=free}} * {{cite journal |last1=Venturieri |first1=G C |last2=Raiol |first2=V F O |last3=Pareira |first3=C A B |year=2003 |title=Avaliação da introfução da criação racional de ''Melipona fasciculata'' (Apidae: Meliponina), entre os agricultores familiares de Bragança - PA, Brasil |journal=Biota Neotropica |volume=3 |issue=2 |pages=1–7 |doi=10.1590/s1676-06032003000200003 |doi-access=free}} * {{Cite journal |last1=Viana |first1=Blandina Felipe |last2=Coutinho |first2=Jeferson Gabriel da Encarnação |last3=Garibaldi |first3=Lucas Alejandro |last4=Castagnino |first4=Guido Laercio Bragança |last5=Gramacho |first5=Kátia Peres |last6=Silva |first6=Fabiana Oliveira |date=2014-10-09 |title=Stingless bees further improve apple pollination and production |url=https://pollinationecology.org/index.php/jpe/article/view/309 |journal=Journal of Pollination Ecology |language=en |volume=14 |pages=261–269 |doi=10.26786/1920-7603(2014)26 |issn=1920-7603|doi-access=free }} * {{cite journal |author=Villanueva, Rogel |display-authors=et al. |year=2005 |title=Extinction of ''Melipona beecheii'' and traditional beekeeping in the Yucatán peninsula |journal=Bee World |volume=86 |issue=2 |pages=35–41 |doi=10.1080/0005772X.2005.11099651 |s2cid=31943555}} * {{Cite journal |last1=Wenseleers |first1=T. |last2=Ratnieks |first2=F. L. W. |last3=Ribeiro |first3=M. D. F. |last4=Alves |first4=D. D. A. |last5=Imperatriz-Fonseca |first5=V. L. |date=June 2005 |title=Working-class royalty: Bees beat the caste system |journal=[[Biology Letters]] |volume=1 |issue=2 |pages=125–128 |doi=10.1098/rsbl.2004.0281 |pmc=1626201 |pmid=17148145}}{{open access}} * {{Cite journal |last1=Witter |first1=Sidia |last2=Blochtein |first2=Betina |last3=Santos |first3=Camila dos |date=August 2007 |title=Abelhas sem ferrão do Rio Grande do Sul: manejo e conservação |url=https://www.agricultura.rs.gov.br/upload/arquivos/202105/11142006-boletim-15.pdf |journal=Boletim FEPAGRO |issue=15}} * {{Cite journal |last1=Zulkhairi Amin |first1=Fatin Aina |last2=Sabri |first2=Suriana |last3=Mohammad |first3=Salma Malihah |last4=Ismail |first4=Maznah |last5=Chan |first5=Kim Wei |last6=Ismail |first6=Norsharina |last7=Norhaizan |first7=Mohd Esa |last8=Zawawi |first8=Norhasnida |date=2018-12-26 |title=Therapeutic Properties of Stingless Bee Honey in Comparison with European Bee Honey |journal=Advances in Pharmacological and Pharmaceutical Sciences |language=en |volume=2018 |pages=e6179596 |doi=10.1155/2018/6179596 |issn=2633-4682 |pmc=6327266 |pmid=30687402 |doi-access=free}} === Books === * {{Cite book |last=Grüter |first=Christoph |url=http://link.springer.com/10.1007/978-3-030-60090-7 |title=Stingless Bees: Their Behaviour, Ecology and Evolution |date=2020 |publisher=Springer International Publishing |isbn=978-3-030-60089-1 |series=Fascinating Life Sciences |location=Cham |language=en |doi=10.1007/978-3-030-60090-7|s2cid=227250633 }} * {{Cite book |last=Jalil |first=Abu Hassan |url=https://books.google.com/books?id=fUh9BAAAQBAJ |title=Beescape for Meliponines: Conservation of Indo-Malayan Stingless Bees |date=2014 |publisher=Partridge Publishing Singapore |isbn=978-1-4828-2362-2 |language=ar}} * {{Cite book |last=Michener |first=Charles D. |url=https://books.google.com/books?id=bu_1gmY13FIC |title=The Bees of the World |date=2000 |publisher=Johns Hopkins Univ. Press |isbn=978-0-8018-6133-8 |location=Baltimore, Md.}} * {{Cite book |last=Roubik |first=David W. |url=https://www.cambridge.org/core/books/ecology-and-natural-history-of-tropical-bees/01B9D0798E5CBA8A68865ED0CC166A4A |title=Ecology and natural history of tropical bees |date=1989 |publisher=Cambridge Univ. Pr |isbn=978-0-521-26236-1 |series=Cambridge tropical biology series |location=Cambridge}} * {{Cite book |last1=Silveira |first1=Fernando A. |url=https://books.google.com/books?id=2-JMAAAAYAAJ&q=Abelhas+brasileiras:+sistem%C3%A1tica+e+identifica%C3%A7%C3%A3o |title=Abelhas brasileiras: sistemática e identificação |last2=Melo |first2=Gabriel A. R. |last3=Almeida |first3=Eduardo A. B. |date=2002 |publisher=Silveira |isbn=978-85-903034-1-1 |edition=1st |location=Belo Horizonte}} * {{cite journal |last1=Venturieri |first1=G C. |year=2004 |title=Meliponicultura: Criação de Abelhas Indígenas Sem Ferrão |url=https://www.infoteca.cnptia.embrapa.br/infoteca/bitstream/doc/410121/1/livroabelha2014.pdf |journal=Comunicado Técnico Embrapa Amazônia Oriental |volume=118 |pages=1–4}} * Villas-Bôas, Jerônimo (2018). ''[https://ispn.org.br/site/wp-content/uploads/2018/10/ManualTecnologicoMel.pdf Manual Tecnológico de Aproveitamento Integral dos Produtos das Abelhas Nativas sem Ferrão]''. Brasília: Instituto Sociedade, População e Natureza (ISPN), v. 2. * {{Cite book |last1=Vit |first1=Patricia |url=https://books.google.com/books?id=NJxPDwAAQBAJ&dq=%22australia%22+%22native%22+%22stingless+bee%22&pg=PA401 |title=Pot-Pollen in Stingless Bee Melittology |last2=Pedro |first2=Silvia R. M. |last3=Roubik |first3=David W. |date=2018-03-05 |publisher=Springer |isbn=978-3-319-61839-5 |language=en}}< ==External links== {{Commons category|Meliponini}} {{Wikispecies|Meliponini|''Meliponini''}} *[http://www.ibra.org.uk/ International Bee Research Association] *[https://web.archive.org/web/20140215065345/http://www.ufv.br/dbg/bee/Versao2/stinglessbees.htm Stingless honey bees of Brazil] *[https://www.sciencedaily.com/releases/2005/06/050615062105.htm Mayan Stingless Bee Keeping: Going, Going, Gone?] *[http://green.blogs.nytimes.com/2012/02/17/a-different-kind-of-beekeeping-takes-flight/?_r=0 A Different Kind of Beekeeping Takes Flight] {{Taxonbar|from=Q127768}} {{Authority control}} [[Category:Meliponini| ]] [[Category:Entomology]] [[Category:Bees]]'
New page wikitext, after the edit (new_wikitext)
'{{Short description|Bee tribe, reduced stingers, strong bites}} {{Automatic taxobox | name = Stingless bees | fossil_range = {{fossil_range|70.6|0|Maastrichtian-Present}} | image = Meliponula ferruginea.jpg | image_caption = ''Meliponula ferruginea'' | taxon = Meliponini | authority = [[Amédée Louis Michel le Peletier, comte de Saint-Fargeau|Lepeletier]], 1836 | subdivision_ranks = Genera | subdivision = {{ubl| ''[[Aparatrigona]]''| ''[[Apotrigona]]''| ''[[Asperplebeia]]''| ''[[Austroplebeia]]''| ''[[Axestotrigona]]''| ''[[Camargoia]]''| ''[[Celetrigona]]''| ''[[Cephalotrigona]]''| ''[[Cleptotrigona]]''| [[Extinction|†]]''[[Cretotrigona]]''| ''[[Dactylurina]]''| ''[[Dolichotrigona]]''| ''[[Duckeola]]''| ''[[Ebaiotrigona]]''| †''[[Exebotrigona]]''| ''[[Friesella]]''| ''[[Frieseomelitta]]''| ''[[Geniotrigona]]''| ''[[Geotrigona]]''| ''[[Heterotrigona]]''| ''[[Homotrigona]]''| ''[[Hypotrigona]]''| †''[[Kelneriapis]]''| ''[[Lepidotrigona]]''| ''[[Lestrimelitta]]''| ''[[Leurotrigona]]''| ''[[Liotrigona]]''| †''[[Liotrigonopsis]]''| ''[[Lisotrigona]]''| ''[[Lophotrigona]]''| ''[[Meliplebeia]]''| ''[[Melipona]]''| †''[[Meliponorytes]]''| ''[[Meliponula]]''| ''[[Meliwillea]]''| ''[[Mourella]]''| ''[[Nannotrigona]]''| ''[[Nanoplebeia]]''| ''[[Nogueirapis]]''| ''[[Odontotrigona]]''| ''[[Oxytrigona]]''| ''[[Papuatrigona]]''| ''[[Parapartamona]]''| ''[[Paratrigona]]''| ''[[Pariotrigona]]''| ''[[Paratrigonoides]]''| ''[[Partamona]]''| ''[[Platytrigona]]''| ''[[Plebeia]]''| ''[[Plebeiella]]''| ''[[Plebeina]]''| ''[[Plectoplebeia]]''| †''[[Proplebeia]]''| ''[[Ptilotrigona]]''| ''[[Sahulotrigona]]''| ''[[Scaptotrigona]]''| ''[[Scaura]]''| ''[[Schwarziana]]''| ''[[Sundatrigona]]''| ''[[Tetragona]]''| ''[[Tetragonilla]]''| ''[[Tetragonisca]]''| ''[[Tetragonula]]''| ''[[Tetrigona]]''| ''[[Trichotrigona]]''| ''[[Trigona (genus)|Trigona]]''| ''[[Trigonisca]]''| ''[[Wallacetrigona]]''}} }} '''Stingless bees (SB)''', sometimes called '''stingless honey bees''' or simply '''meliponines''', are a large group of bees (from about 462 to 552 described species),<ref>{{harvtxt|Grüter|2020|pp=46-47}}</ref><ref name=":27">{{harvtxt|Nogueira|2023}}</ref> comprising the [[Tribe (biology)|tribe]] '''Meliponini'''<ref>{{harvtxt|Michener|2000|p=803}}</ref><ref>{{harvtxt|Grüter|2020|p=1}}</ref> (or [[subtribe]] Meliponina according to other authors).<ref>{{harvtxt|Silveira|2002|p=253}}</ref> They belong in the [[Family (biology)|family]] [[Apidae]] ([[subfamily]] [[Apinae]]), and are closely related to common [[honey bee]]s (HB, tribe [[Apini]]), orchid bees (tribe [[Euglossini]]), and [[bumblebee]]s (tribe [[Bombini]]). These four bee tribes belong to the [[Pollen basket|corbiculate]] bees [[Monophyly|monophyletic]] group.<ref>{{harvtxt|Grüter|2020|p=43}}</ref><ref>{{harvtxt|Roubik|1989|p=8}}</ref> Meliponines have stingers, but they are highly reduced and cannot be used for defense, though these bees exhibit other defensive behaviors and mechanisms.<ref>{{harvtxt|Kajobe|2006}}</ref><ref name=":1">{{harvtxt|Chakuya|Gandiwa|Muboko|Muposhi|2022}}</ref> Meliponines are not the only type of bee incapable of stinging: all male bees and many female bees of several other families, such as [[Andrenidae]] and [[Megachilidae]] (tribe [[Dioxyini]]), also cannot sting.<ref>{{harvtxt|Michener|2000|p=111}}</ref> Some stingless bees have powerful mandibles and can inflict painful bites.<ref>{{harvtxt|Sarchet|2014}}</ref><ref>{{harvtxt|Grüter|2020|pp=7 & 16}}</ref> Some species can present large mandibular glands for the secretion of caustic defense substances, secrete unpleasant smells or use sticky materials to immobilise enemies.<ref name=":3">{{harvtxt|Grüter|2020|p=65}}</ref><ref>{{harvtxt|Grüter|2020|p=4}}</ref> The main honey-producing bees of this group generally belong to the genera ''[[Scaptotrigona]]'', ''[[Tetragonisca]]'', ''[[Melipona]]'' and ''[[Austroplebeia]]'', although there are other genera containing species that produce some usable honey. They are farmed in [[meliponiculture]] in the same way that European honey bees (genus ''[[Apis (genus)|Apis]]'') are cultivated in [[apiculture]]. Throughout [[Mesoamerica]], the [[Maya peoples|Mayans]] have engaged in extensive meliponiculture on a large scale since before the arrival of Columbus. Meliponiculture played a significant role in [[Maya society]], influencing their social, economic, and religious activities. The practice of maintaining '''stingless bees''' in man-made structures is prevalent across the Americas, with notable instances in countries such as [[Brazil]], [[Peru]], and [[Mexico]].<ref>{{harvtxt|Grüter|2020|pp=25-27}}</ref><ref>{{harvtxt|Villas-Bôas|2018|pp=14-15}}</ref> == Geographical distribution == Stingless bees can be found in most tropical or subtropical regions of the world, such as the [[Africa|African continent]] ([[Afrotropical realm|Afrotropical]] region), [[Southeast Asia]] and [[Australia]] ([[Indomalayan realm|Indo-Malayan]] and [[Australasian realm|Australasian]] region), and tropical [[Americas|America]] ([[Neotropical realm|Neotropical]] region).<ref>{{harvtxt|Michener|2000|p=803}}</ref><ref name=":0">{{harvtxt|Grüter|2020|pp=47-49}}</ref><ref>{{harvtxt|Silveira|2002|p=31}}</ref><ref>{{harvtxt|Rasmussen|Thomas|Engel|2017}}</ref> The majority of [[Indigenous (ecology)|native]] [[eusocial]] bees of [[Central America|Central]] and [[South America]] are SB, although only a few of them produce honey on a scale such that they are farmed by humans.<ref>{{harvtxt|Cortopassi-Laurino|Imperatriz-Fonseca|Roubik|2006}}</ref><ref>{{harvtxt|Venturieri|Raiol|Pareira|2003}}</ref> The Neotropics, with approximately 426 species, boast the highest abundance and species richness, ranging from Cuba and Mexico in the north to Argentina in the south.<ref name=":0" /> They are also quite diverse in [[Africa]], including [[Madagascar]],<ref>{{harvtxt|Koch|2010}}</ref> and are farmed there also. Around 36 species exist on the continent. The equatorial regions harbor the greatest diversity, with the [[Sahara|Sahara Desert]] acting as a natural barrier to the north. The range extends southward to [[South Africa]] and southern Madagascar, with most African species inhabiting [[tropical forest]]s or both tropical forests and [[savanna]]hs.<ref name=":0" /> Meliponine honey is prized as a medicine in many African communities, as well as in South America. Some cultures use SB honey against [[Human digestive system|digestive]], [[Respiratory system|respiratory]], [[Eye|ocular]] and [[Reproductive system|reproductive]] problems, although more research is needed to disclose evidence that supports these practices.<ref name=":1" /><ref>{{harvtxt|Grüter|2020|p=|pp=28-29}}</ref><ref>{{harvtxt|Souza|Yuyama|Aguiar|Oliveira|2004}}</ref> In Asia and Australia, approximately 90 species of stingless bees span from [[India]] in the west to the [[Solomon Islands]] in the east, and from [[Nepal]], China ([[Yunnan]], [[Hainan]]), and [[Taiwan]] in the north to Australia in the south. [[File:Geographic distribution of stingless bees.png|center|thumb|399x399px|Geographic distribution of stingless bees.]] === Origin and dispersion === Phylogenetic analyses reveal three distinct groups in the evolutionary history of '''Meliponini''': the Afrotropical, the Indo-Malay/Australasia, and the Neotropical lineages. The evolutionary origin of the Meliponini is Neotropical. Studies observing contemporary [[species richness]] show that it remains highest in the Neotropics.<ref name=":2">{{harvtxt|Roubik|2023a}}</ref> The hypothesis proposes the potential dispersion of stingless bees from what is now [[North America]]. According to this scenario, these bees would have then traveled to Asia by crossing the [[Bering Strait]] ([[Beringia]] route) and reached [[Europe]] through [[Greenland]] ([[Thule Land Bridge|Thulean route]]).<ref>{{harvtxt|Grüter|2020|pp=54-56}}</ref><ref name=":2" /><ref>{{harvtxt|Rasmussen|Cameron|2009}}</ref><ref>{{harvtxt|Jalil|2014|p=126}}</ref> == Evolution and phylogeny == Meliponines form a [[clade]] within the corbiculate bees, characterized by unique pollen-carrying structures known as corbiculae ([[Pollen basket|pollen baskets]]) located on their hind legs. This group also includes other three tribes: honey bees ([[Apini]]), bumble bees ([[Bombini]]), and orchid bees ([[Euglossini]]). The concept of higher [[eusociality]], defined by the presence of distinct queen and worker castes and characterized by features such as perennial colony lifestyles and extensive food sharing among adults, is particularly relevant in understanding the social structure of these tribes. Both Meliponini and Apini tribes are considered higher eusocial, while Bombini is considered to be primitively eusocial.<ref name=":29">{{harvtxt|Grüter|2020|pp=43-44}}</ref> The phylogenetic relationships among the four tribes of corbiculate bees have been a topic of considerable debate within the scientific community. Two primary questions arise: the relationship of stingless bees to honey bees and bumble bees, and whether their eusocial behavior evolved independently or from a [[Common descent|common ancestor]]. Morphological and behavioral studies have suggested that Meliponini and Apini are sister groups, indicating a single origin of higher eusociality. In contrast, molecular studies often support a relationship between Meliponini and Bombini, proposing independent origins of higher eusociality in both Apini and Meliponini.<ref name=":29" /> A morphological, behavioral, and molecular data analysis provided strong support for the latter hypothesis of dual origins of higher eusociality. Subsequent research has reinforced the idea that stingless bees and honey bees evolved their eusocial lifestyles independently, resulting in distinct adaptive strategies for colony reproduction, brood rearing, foraging communication, and colony defense. This divergence helps explain the varied ecological and social solutions developed by these two groups of bees, such as [[foraging]] communication, colony defense/reproduction and brood rearing.<ref name=":29" /><ref>{{harvtxt|Payne|2014}}</ref> {{clade| {{clade |1={{clade |1=[[Melittidae]] |2={{clade |label1={{nowrap|long-tongued bees}} |1={{clade |label1=&nbsp;[[Apidae]]&nbsp; |1={{clade |label2={{nowrap|eusocial}} |1=[[Euglossini]] |2={{clade |label1={{nowrap|h. eusocial}} |1=[[Apini]] |2={{clade |1=[[Bombini]] |label1={{nowrap|p. eusocial}} |2=[[Meliponini]] |label2={{nowrap|h. eusocial}} |style2=background-color:#f9a6e8 }} }}}} |3=[[Megachilidae]] (mason, leafcutter bees) }} |label2={{nowrap|short-tongued bees}} |2={{clade |1=[[Andrenidae]] (mining bees) |2={{clade |1=[[Halictidae]] (sweat bees) |2={{clade |2=[[Colletidae]] (plasterer bees) |1=[[Stenotritidae]] (large Australian bees) }} }} }} }} }} }} |style=font-size:85%;line-height:85%|label1=[[Anthophila]] (bees)}} === Fossil history === The [[fossil]] record for stingless bees is notably robust compared to that of many other bee groups, with twelve extinct species currently identified. Fossils of these bees are primarily found in [[amber]] and [[copal]], where excellent preservation typically occurs. This favorable fossilization process may be attributed to the behaviors of stingless bees, which collect significant amounts of tree [[resin]] for building nests and defense, increasing the likelihood of entrapment.<ref name=":24">{{harvtxt|Grüter|2010|pp=49-51}}</ref> Despite this relatively good fossil record, the evolutionary history of stingless bees remains poorly understood, particularly regarding their widespread distribution across various ecological niches around the globe. The oldest known fossil stingless bee is ''[[Cretotrigona prisca]]'', a small worker bee approximately 5 mm in body length, discovered in New Jersey amber. This species is believed to have existed during the [[Late Cretaceous]] period, around 65–70 million years ago, marking it as the oldest confirmed fossil of an apid bee and the earliest fossil evidence of a eusocial bee. ''C. prisca'' exhibits striking similarities to extant stingless bees, indicating that the evolutionary lineage of meliponines may date back to this period.<ref name=":24" /> Some researchers suggest that stingless bees likely evolved in the Late Cretaceous, approximately 70–87 million years ago.<ref name=":24" /><ref>{{harvtxt|Rasmussen|Cameron|2009}}</ref><ref>{{harvtxt|Cardinal|Danforth|2011}}</ref> According to recent studies, corbiculate bees, which include stingless bees, are thought to have appeared around 84–87 million years ago, further supporting the notion of their evolution during this dynamic period in Earth's history.<ref name=":24" /><ref>{{harvtxt|Cardinal|Danforth|2013}}</ref><ref>{{harvtxt|Martins|Melo|Renner|2014}}</ref> == Behaviour, biology and ecology == === Overview === Meliponines, considered highly eusocial insects, exhibit a remarkable caste division. The colonies typically consist of a [[Queen bee|queen]], [[Worker bee|workers]], and sometimes male [[Drone (bee)|drones]].<ref name=":4">{{harvtxt|Nogueira-Neto|1997|p=78}}</ref> The queen is responsible for reproduction, while the workers perform various tasks such as foraging, nursing, and defending the colony. Individuals work together with a well-defined division of labor for the overall benefit.<ref name=":5">{{harvtxt|Nogueira-Neto|1997|p=82}}</ref> Stingless bees are valuable pollinators and contribute to ecosystem health by producing essential products. These insects collect and store honey, pollen, resin, propolis, and cerumen. Honey serves as their primary [[carbohydrate]] source, while pollen provides essential [[protein]]s. Resin, propolis, and cerumen are used in nest construction and maintenance.<ref>{{harvtxt|Nogueira-Neto|1997|pp=40-46}}</ref><ref>{{harvtxt|Grüter|2020|pp=12-15}}</ref> Nesting behavior varies among species and may involve hollow tree trunks, external hives, the soil, termite nest or even urban structures. This adaptability underscores their resilience and ability to coexist with human activities.<ref>{{harvtxt|Grüter|2020|pp=87-121}}</ref> === Castes === ==== Workers ==== {{Main|Worker bee}} In a SB colony, workers constitute the predominant segment of the population, serving as the colony's primary workforce. They undertake a multitude of responsibilities crucial for the colony's well-being, including defense, cleaning, handling building materials, and the collection and processing of food. Recognizable by the corbicula - a distinctive structure on their hind legs resembling a small basket - workers efficiently carry pollen, resin, clay, and other materials gathered from the environment. Given their abundance and unique physical feature, workers play a central role in sustaining the colony.<ref>{{harvtxt|Grüter|2020|pp=7-11}}</ref><ref name=":9">{{harvtxt|Villas-Bôas|2018|p=23}}</ref><ref>{{harvtxt|Nogueira-Neto|1997|pp=82-83}}</ref> ==== Queens ==== {{Main|Queen bee}} The principal egg layer in SB colonies is the queen, distinguished from the workers by differences in both size and shape. Stingless bee queens - except in the case of the ''[[Melipona]]'' genus, where queens and workers receive similar amounts of food and thus exhibit similar sizes - are generally larger and weigh more than workers (approximately 2–6 times). Post-mating, meliponine queens undergo [[Physogastrism|physogastry]], developing a distended abdomen. This physical transformation sets them apart from honey bee queens, and even ''Melipona'' queens can be easily identified by their enlarged abdomen after mating.<ref name=":4" /><ref name=":6">{{harvtxt|Villas-Bôas|2018|p=19}}</ref><ref name=":7">{{harvtxt|Grüter|2020|pp=7 & 5}}</ref> Stingless bee colonies typically follow a monogynous structure, featuring a single egg-laying queen. An exception is noted in ''[[Melipona bicolor]]'' colonies, which are often polygynous (large populations may have as many as 5 physogastric queens simultaneously involved in oviposition).<ref name=":7" /><ref name=":5" /><ref name=":8">{{harvtxt|Imperatriz-Fonseca|Zucchi|1995}}</ref> Depending on the species, queens can lay varying quantities of eggs daily, ranging from a dozen (e.g., ''[[Plebeia julianii]]'') to several hundred (e.g., ''[[Trigona recursa]]''). While information on queen lifespans is limited, available data suggest that queens generally outlive workers, with lifespans usually falling between 1 and 3 years, although some queens may live up to 7 years.<ref name=":7" /><ref name=":8" /> The laying queen assumes the crucial role of producing eggs that give rise to all castes within the colony. Additionally, she plays a pivotal role in organizing the colony, overseeing a complex communication system primarily reliant on the use of [[pheromone]]s.<ref name=":6" /> ==== Males (drones) ==== {{Main|Drone (bee)}} The primary function of males, or drones, is to mate with queens, performing limited tasks within the nest and leaving at around 2–3 weeks old, never to return. The production of males can vary, occurring continuously, sparsely or in large spurts when numerous drones emerge from brood combs for brief periods. Identifying a male can be challenging due to its similar body size to workers, but distinctive features such as the absence of a corbicula, larger eyes, slightly smaller [[Mandible (insect mouthpart)|mandibles]], slightly longer and v-shaped [[Antenna (biology)|antennae]], and often a lighter face color distinguish them. Clusters of males, numbering in the hundreds, can be observed outside colonies, awaiting the opportunity to mate with virgin queens.<ref name=":9" /><ref>{{harvtxt|Nogueira-Neto|1997|pp=83-85}}</ref><ref>{{harvtxt|Grüter|2020|p=11}}</ref> Males in a stingless bee colony, either produced mainly by the laying queen or primarily by the workers, play an important role in reproduction. Workers can produce males by laying unfertilized eggs, enabled by the [[haplodiploidy]] system, where males are [[Ploidy|haploid]], having only one set of [[chromosome]]s, while workers are diploid and incapable of producing female eggs due to their inability to mate. This sex determination system is common to all [[hymenoptera]]ns.<ref>{{harvtxt|Grüter|2020|p=11}}</ref> ==== Soldiers ==== While the existence of a soldier caste is well known in [[ant]]s and [[termite]]s, the phenomenon was unknown among bees until 2012, when some stingless bees were found to have a similar caste of defensive specialists that help guard the nest entrance against intruders.<ref>{{harvtxt|Grüter|Menezes|Imperatriz-Fonseca|Ratnieks|2012}}</ref> To date, at least 10 species have been documented to possess such "soldiers", including ''[[Tetragonisca angustula]]'', ''[[Tetragonisca fiebrigi|T. fiebrigi]]'', and ''[[Frieseomelitta longipes]]'', with the guards not only larger, but also sometimes a different color from ordinary workers.<ref>{{harvtxt|Grüter|Segers|Menezes|Vollet-Neto|Falcón|von Zuben|Bitondi|Nascimento|Almeida|2017}}</ref><ref>{{harvtxt|Grüter|2020|p=258}}</ref> ==== Division of labour ==== When the young worker bees emerge from their cells, they tend to initially remain inside the hive, performing different jobs. As workers age, they become guards or foragers. Unlike the [[larva]]e of [[honey bee]]s and many social wasps, meliponine larvae are not actively fed by adults ([[progressive provisioning]]). Pollen and nectar are placed in a cell, within which an [[egg]] is laid, and the cell is sealed until the adult bee emerges after [[pupa]]tion ([[mass provisioning]]). At any one time, hives can contain from 300 to more than 100,000 workers (with some authors claiming to calculate more than 150,000 workers, but with no methodology explanation), depending on species.<ref>{{harvtxt|Grüter|2020|pp=17-24}}</ref> === Products and materials === The industrious nature of stingless bees extends to their building activities. Unlike honey bees, they do not use pure wax for construction but combine it with resin to create cerumen, a material employed in constructing nest structures such as brood cells, food pots, and the protective involucrum. [[Beeswax|Wax]] is secreted by young bees through glands located on the top of their abdomen and this mixture not only provides structural strength but also offers antimicrobial properties, inhibiting the growth of fungi and bacteria. The creation of batumen involves combining cerumen with additional resin, mud, plant material, and sometimes even animal feces. Batumen, a stronger material, forms protective layers covering the walls of the nesting space, ensuring the safety of the colony.<ref name=":10">{{harvtxt|Nogueira-Neto|1997|pp=40-46}}</ref><ref name=":11">{{harvtxt|Grüter|2020|pp=12-15}}</ref><ref name=":12">{{harvtxt|Villas-Bôas|2018|pp=24-26}}</ref><ref name=":13">{{harvtxt|Venturieri|2004|pp=23-30}}</ref> On the other hand, [[clay]], sourced from the wild and exhibiting diverse colors based on its mineral origin, serves as another essential raw material for SB. While it can be used in its pure form, it is more common to combine clay with vegetable [[resin]]s to produce geopropolis. The inclusion of clay in this mixture enhances the durability and structural integrity of the resulting substance.<ref name=":10" /><ref name=":11" /><ref name=":12" /><ref name=":13" /> Vegetable resin, gathered from a variety of plant species in the wild, is an essential raw material brought back to the hive. Stored in small, sticky clumps in peripheral areas of the colony, it is often mistakenly treated as a synonym for [[propolis]]. However, in beekeeping terminology, propolis refers to a mixture of resin, wax, enzymes, and possibly other substances. Stingless bees go beyond the classic propolis by producing various derivatives from resins and wax, sometimes using pure resins for sealing or defense, a behavior not observed in ''Apis'' bees. Understanding these distinctions is vital for effective production and value addition to the meliponiculture activity.<ref name=":10" /><ref name=":11" /><ref name=":12" /><ref name=":13" /> [[Honey]], a prized product of bee colonies, is crafted through the processing of [[nectar]]s, [[Honeydew (secretion)|honeydews]], and fruit juices by worker bees. They store these collected substances in an extension of their gut called a [[crop (anatomy)|crop]]. Back at the hive, the bees ripen or dehydrate the nectar droplets by spinning them inside their mouthparts until honey is formed. Ripening concentrates the nectar and increases the sugar content, though it is not nearly as concentrated as the honey from ''Apis mellifera''. Stored in food pots, meliponines' honey is often referred to as pot-honey due to its distinctive storage method. Stingless bee honeys differ from ''A. mellifera'' honey in terms of color, texture, and flavor, being more liquid with a higher water content. Rich in [[mineral]]s, [[amino acid]]s, and [[flavonoid]] compounds, the composition of honey varies among colonies of the same species, influenced by factors such as season, habitat, and collected resources.<ref name=":10" /><ref name=":11" /><ref name=":12" /><ref name=":13" /> Special methods are being developed to harvest moderate amounts of honey from stingless bees in these areas without causing harm. For honey production, the bees need to be kept in a box specially designed to make the honey stores accessible without damaging the rest of the nest structure. Some recent box designs for honey production provide a separate compartment for the honey stores so the honey pots can be removed without spilling honey into other areas of the nest. Unlike a hive of commercial honeybees, which can produce 75 kg (165 lbs) of honey a year, a hive of Australian stingless bees produces less than 1 kg (2 lbs). Stingless bee honey has a distinctive "bush" taste—a mix of sweet and sour with a hint of fruit.<ref>{{harvtxt|Mduda|Hussein|Muruke|2023}}</ref><ref>{{harvtxt|Ferreira|Lencioni|Benassi|Barth|Bastos|2009}}</ref><ref>{{harvtxt|Sousa|Souza|Marques|Benassi|Gullón|Pintado|Magnani|2016}}</ref> The taste comes from plant resins—which the bees use to build their hives and honey pots—and varies at different times of year depending on the flowers and trees visited. In 2020 researchers at the [[University of Queensland]] found that some species of stingless bee in Australia, Malaysia, and Brazil produce honey that has [[trehalulose]]—a sugar with an unusually low [[Glycemic index|glycaemic index]] (GI) compared to that of glucose and fructose, the main sugars composing conventional honey.<ref>{{harvtxt|Mokaya|Nkoba|Ndunda|Vereecken|2002}}</ref><ref name=":25">{{harvtxt|Fletcher|Hungerford|Webber|Carpinelli de Jesus|Zhang|Stone|Blanchfield|Zawawi|2020a}}</ref> Such low glycaemic index honey is beneficial for humans because its consumption does not cause [[blood sugar level|blood sugar]] to spike, forcing the body to make more insulin in response. Honey with trehalulose is also beneficial as it this sugar cannot nourish the lactic acid-producing bacteria that cause [[tooth decay]]. The university's findings supported the long-standing claims of [[Indigenous Australian people]] that native honey is beneficial to human health.<ref name=":25" /><ref>{{Cite web |last=Layt |first=Stuart |date=2020-07-23 |title=Scientists say native stingless bee honey hits the sweet spot |url=https://www.brisbanetimes.com.au/national/queensland/scientists-say-native-stingless-bee-honey-hits-the-sweet-spot-20200723-p55eud.html |access-date=2020-07-27 |website=Brisbane Times |language=en}}</ref> This type of honey is scientifically supported as providing therapeutic value to humans as well.<ref name=":25" /><ref>{{harvtxt|Mduda|Hussein|Muruke|2023}}</ref><ref>{{harvtxt|Rodríguez-Malaver|Rasmussen|Gutiérrez|Gil|Nieves|Vit|2009}}</ref><ref>{{harvtxt|Nweze|Okafor|Nweze|Nweze|2017}}</ref><ref>{{harvtxt|Zulkhairi Amin|Sabri|Mohammad|Ismail|Chan|Ismail|Norhaizan|Norhaizan|2018}}</ref> === Nest === [[File:Natural cavity stingless bee nest.png|thumb|General main elements of stingless bees cavity nest colonies.]] Stingless bees, as a collective group, display remarkable adaptability to diverse nesting sites. They can be found in exposed nests in trees, from living in ant and termite nests above and below ground to cavities in trees, trunks, branches, rocks, or even human constructions.<ref name=":14">{{harvtxt|Grüter|2020|pp=87-97}}</ref> Many [[beekeeper]]s keep the bees in their original log [[beehive|hive]] or transfer them to a wooden box, as this makes controlling the hive easier. Some beekeepers put them in bamboos, flowerpots, coconut shells, and other recycling containers such as a water jug, a broken guitar, and other safe and closed containers.<ref>{{harvtxt|Venturieri|2004|pp=36-39}}</ref><ref>{{harvtxt|Contrera|Venturieri|2008}}</ref><ref name=":26" /> ==== Exposed nests ==== [[File:Arboreal stingless bee nest (Trigona sp.) Flores.jpg|thumb|''Trigona'' sp. exposed nest in a tree fork near [[Flores, Petén|Flores]], [[Guatemala]].]] Notably, certain species, such as the African ''Dactylurina'', construct hanging nests from the undersides of large branches for protection against adverse weather conditions. Additionally, some American ''[[Trigona]]'' species, including ''[[Trigona corvina|T. corvina]]'', ''[[Trigona spinipes|T. spinipes]]'', and T. nigerrima, as well as ''Tetragonisca weyrauchi'', build fully exposed nests.<ref name=":14" /> ==== Ground nests ==== [[File:Paratrigona ground nest.jpg|left|thumb|''Paratrigona'' sp. ground nest in [[Brasília]], [[Brazil]].]] A significant minority of meliponine species, belonging to genera like ''[[Camargoia]]'', ''[[Geotrigona]]'', ''[[Melipona]]'', ''[[Mourella]]'', ''[[Nogueirapis]]'', ''[[Paratrigona]]'', ''[[Partamona]]'', ''[[Schwarziana]]'', and others, opt for ground nests. These species take advantage of cavities in the ground, often utilizing abandoned nests of ants, termites, or rodents. Unlike some other cavity-nesting bees, stingless bees in this category do not excavate their own cavities but may enlarge existing ones.<ref name=":14" /> ==== Termite and ant shared nests ==== Numerous stingless bee species have evolved to coexist with termites. They inhabit parts of ant or termite nests, both above and below ground. These nests are often associated with various ant species, such as ''[[Azteca (ant)|Azteca]]'', ''[[Carpenter ant|Camponotus]]'', or ''[[Crematogaster]]'', and termite species like ''[[Nasutitermes]]'', ''[[Constrictotermes]]'', ''[[Macrotermes]]'', ''Microcerotermes'', ''[[Odontotermes]]'', or ''Pseudocanthotermes''. This strategy allows SB to utilize pre-existing cavities without the need for extensive excavation.<ref name=":14" /> ==== Cavity nests ==== The majority of stingless bees favor nesting in pre-existing cavities within tree trunks or branches. Nesting heights vary, with some colonies positioned close to the ground, typically below 5 meters, while others, like ''Trigona'' and ''[[Oxytrigona]]'', may nest at higher elevations, ranging from 10 to 25 meters. Some species, such as ''[[Melipona nigra]]'', exhibit unique nesting habits at the foot of a tree in root cavities or between roots. The choice of nesting height has implications for predation pressure and the microclimate experienced by the colony.<ref name=":14" /> The majority of stingless bee species exhibit a non-specific preference when it comes to selecting tree species for nesting. Instead, they opportunistically exploit whatever nesting sites are available This adaptability underscores the versatility of SB in adapting to various arboreal environments. Furthermore, cavity-nesting species can opportunistically utilize human constructions, nesting under roofs, in hollow spaces in walls, electricity boxes, or even metal tubes. In few cases, specific tree species, like ''[[Caryocar brasiliense]]'', may be preferred by certain stingless bee species (''[[Melipona quadrifasciata]]''), illustrating a degree of selectivity in nesting choices among different groups.<ref name=":14" /><ref>{{harvtxt|Antonini|Martins|2003}}</ref> ==== Entrances ==== Entrance tubes showcase a spectrum of characteristics, from being hard and brittle to soft and flexible. In many situations, the portion near the opening remains soft and flexible, aiding workers in sealing the entrance during the night. The tubes may also feature perforations and a coating of resin droplets, adding to the complexity of their design.<ref name=":15">{{harvtxt|Grüter|2020|pp=102-109}}</ref> The entrances serve as essential visual landmarks for returning bees, and they are often the first structures constructed at a new nest site. The diversity in entrance size influences foraging traffic, with larger entrances facilitating smoother traffic but potentially necessitating more entrance guards to ensure adequate defense.<ref name=":15" /> Some ''[[Partamona]]'' species exhibit a distinctive entrance architecture, where workers of ''[[Partamona helleri|P. helleri]]'' construct a large outer mud entrance leading to a smaller adjacent entrance. This unique design enables foragers to enter with high speed, bouncing off the ceiling of the outer entrance towards the smaller inner entrance. The peculiar appearance of this entrance has led to local names such as "toad mouth", highlighting the intriguing adaptations found in stingless bee nest entrances.<ref name=":15" /> <gallery perrow="4" widths="160" caption="Different species nest entrances in Brazilian stingless bees"> File:Abelha limão.jpg|''[[Lestrimelitta limao]]'' File:Frieseomelitta varia nest entrance.jpg|''[[Frieseomelitta varia]]'' File:Partamona helleri colmeia.jpg|''[[Partamona helleri]]'' File:Entrada de ninho de Scaptotrigona postica.jpg|''[[Scaptotrigona postica]]'' File:Parque Nacional da Serra do Itajaí - Liu Idárraga Orozco (86).jpg|''[[Oxytrigona tataira]]'' File:Melipona quadrifasciata nest guard.jpg|''[[Melipona quadrifasciata]]'' File:Tetragonisca angustula nest entrance.jpg|''[[Tetragonisca angustula]]'' File:Nannotrigona testaceicornis nest entrance.jpg|''[[Nannotrigona testaceicornis]]'' </gallery> [[File:Entrant tubes.png|thumb|Variation of entrant tube in some native stingless bee species found in Southeast Asia: (1) ''[[Geniotrigona thoracica]]'', (2) ''[[Homotrigona fimbriata]]'', (3) ''[[Lophotrigona canifrons]]'', (4) ''[[Tetragonilla collina]]'', (5) ''[[Pariotrigona klossi]]'', (6) ''[[Heterotrigona itama]]'', (7) ''[[Tetragonula fuscobalteata]]'', (8) ''[[Lepidotriogona terminata]]'', (9) ''[[Tetrigona melanoleuca]]'', (10) ''[[Tetrigona apicalis]]'', and (11) ''[[Tetragonula pagdeni]]''.]] ==== Brood cell arrangement ==== Stingless bee colonies exhibit a diversity of construction patterns of brood cells, primarily composed of soft cerumen, a mixture of wax and resin. Each crafted cell is designed to rear a single individual bee, emphasizing the precision and efficiency of their nest architecture.<ref name=":15" /> The quantity of brood cells within a nest displays significant variation across different stingless bee species. Nest size can range from a few brood cells, as observed in the Asian ''[[Lisotrigona carpenteri]]'', to remarkably expansive colonies with over 80,000 brood cells, particularly in some American ''[[Trigona]]'' species.<ref name=":15" /> Meliponine colonies exhibit diverse brood cell arrangements, primarily categorized into three main types: horizontal combs, vertical combs, and clustered cells. Despite these primary types, variations and intermediate forms are prevalent, contributing to the flexibility of nest structures.<ref name=":15" /><ref>{{harvtxt|Villas-Bôas|2018|pp=27-29}}</ref> The first type involves horizontal combs, often characterized by a spiral pattern or layers of cells. The presence of spirals may not be consistent within a species, varying among colonies or even within the same colony. Some species, such as ''[[Melipona]]'', ''[[Plebeia]]'', ''Plebeina'', ''[[Nannotrigona]]'', ''Trigona'', and ''Tetragona'', may occasionally build spirals alongside other comb structures, as observed in ''[[Oxytrigona mellicolor]]''. As space diminishes for upward construction, workers initiate the creation of a new comb at the bottom of the brood chamber. This innovative approach optimizes the available space when emerging bees vacate older, lower brood combs.<ref name=":15" />[[image:Sarang lebah kelulut.jpeg|thumb|250px|left|Hive box containing colony of ''[[Heterotrigona itama]]'']] The second prevalent brood cell arrangement involves clusters of cells held together with thin cerumen connections. This clustered style is observed in various distantly related genera, such as the American ''[[Trigonisca]]'', ''[[Frieseomelitta]]'', ''[[Leurotrigona]]'', the Australian ''[[Austroplebeia]]'', and the African ''[[Hypotrigona]]''. This arrangement is particularly useful for colonies in irregular cavities unsuitable for traditional comb building.<ref name=":15" /> The construction of vertical combs is a distinctive trait found in only two stingless bee species: the African ''[[Dactylurina]]'' and the American ''[[Scaura longula]]''. This vertical arrangement sets these species apart from the more commonly observed horizontal comb structures in other stingless bee genera.<ref name=":15" /> ==== Brood rearing ==== Stingless bee brood rearing is a sophisticated and intricately coordinated process involving various tasks performed by worker bees, closely synchronized with the queen's activities. The sequence begins with the completion of a new brood cell, marking the initiation of mass provisioning.<ref name=":16">{{harvtxt|Grüter|2020|p=161}}</ref> Upon finishing a brood cell, several workers engage in mass provisioning, regurgitating larval food into the cell. This collective effort is swiftly followed by the queen laying her egg on top of the provided larval food. The immediate sealing of the cell ensues shortly afterward, culminating this important phase of the brood rearing process.<ref name=":16" /> The practice of mass provisioning, oviposition, and cell sealing is considered an ancestral trait, shared with solitary wasps and bees. However, in the context of stingless bees, these actions represent distinct stages of a highly integrated social process. Notably, the queen plays a central role in orchestrating these activities, acting as a pacemaker for the entire colony.<ref name=":16" /> This process diverges significantly from brood rearing in ''Apis'' spp. In honeybee colonies, queens lay eggs into reusable empty cells, which are then progressively provisioned over several days before final sealing. The contrasting approaches in brood rearing highlight the unique social dynamics and adaptations within stingless bee colonies.<ref name=":16" /> === Swarming === {{See also|Swarming (honey bee)}} Stingless bees and honey bees, despite encountering a common challenge in establishing daughter colonies, employ contrasting strategies. There are three key differences: reproductive status and age of the queen that leaves the nest, temporal aspects of colony foundation, and communication processes for nest site selection.<ref name=":17">{{harvtxt|Grüter|2020|pp=131-152}}</ref> In HB (''Apis mellifera''), the mother queen, accompanied by a swarm of numerous workers, embarks on relocation to a new home once replacement queens have been reared. Conversely, in SB (meliponines), the departure is orchestrated by the unmated ("virgin") queen, leaving the mother queen in the original nest. Mated stingless bee cannot leave the hive due to damaged wings and increased abdominal size post-mating ([[physogastrism]]). The queen's weight in species like ''[[Scaptotrigona postica]]'' increases, for example, about 250%.<ref name=":17" /><ref>{{harvtxt|Engels|1987}}</ref> Unlike honey bees, stingless bee colonies are unable to perform absconding - a term denoting the abandonment of the nest and migration to a new location - making them reliant on alternative strategies to cope with challenges. Meliponines progressive found new colonies without abandonning their nest abruptly.<ref name=":17" /> These are the stages of stingless bees swarming:<ref name=":17" /> # Reconnaissance and preparation: Scouts inspect potential new nest sites for suitability, considering factors such as cavity size, entrance characteristics, and potential threats. The criteria for determining suitability remain largely unexplored. Some colonies engage in simultaneous preparation of multiple cavities before making a final decision and some others make the initial reconnaissance but do not move into the cavity; # Transport of building material and food: workers seal cracks in the chosen cavity using materials like resin, batumen, or mud. They construct an entrance tube, possibly serving as a visual beacon for nestmate workers. Early food pots are built and filled with honey, requiring a growing number of workers to transport cerumen and honey from the mother nest. # Progressive establishment and social link: the mother and daughter colony maintain a social link through workers traveling between the two nests. The duration of this link varies among species, ranging from a few days to several months. Stingless bee colonies display a preference for cavities previously used by other colonies, containing remnants of building material and nest structures. # Arrival of the queen: after initial preparations, an unmated queen, accompanied by additional workers, arrives at the new nest site. # Drone arrival: males (drones) aggregate outside the newly established nest. They often arrive shortly after swarming initiation, even before the completion of nest structures. Males can be observed near the entrance, awaiting further events. # Mating flight: males in aggregations do not enter the colony but await the queen's emergence for a mating flight. Although rarely observed, it is assumed that unmated stingless bee queens embark on a single mating flight, utilizing acquired sperm for the entirety of their reproductive life. === Natural enemies === In meliponiculture, beekeepers need to be aware of the presence of animals that can harm stingless bee colonies. There are several potential enemies, but the most damaging ones to meliponaries are listed below.<ref>{{harvtxt|Nogueira-Neto|1997|pp=367-390}}</ref> ==== Invertebrates ==== [[File:Forídeo e armadilha.png|thumb|Phorid fly and vinegar trap.]] [[Phoridae|Phorid]] flies in the genus ''[[Pseudohypocera]]'' pose a significant threat to stingless bee colonies, causing problems for beekeepers. These parasites lay eggs in open cells of pollen and honey, leading to potential extinction if not addressed. Early detection is crucial for manual removal or using [[vinegar]] traps. It's important never to leave an infested box unattended to prevent the cycle from restarting and avoid contaminating other colonies. Careful handling of food jars, especially during swarms transfers, is essential. Prompt removal of broken jars, sealing gaps with wax or tape, and maintaining vigilance during the rainy season for heightened phorid activity are recommended. Combatting these flies usually is a priority, particularly during increased reproductive periods.<ref name=":72">Embrapa. ''[https://www.embrapa.br/documents/1355163/39571288/Roteiro+Teórica+-+Cuidados+Especiais.pdf/4fd9e18e-34ea-638c-250b-73c2241e6f07 Inimigos Naturais & Cuidados Especiais].'' Curso Básico de Abelhas Sem Ferrão.</ref><ref name=":82">{{harvtxt|Villas-Bôas|2018|p=|pp=103-104}}</ref><ref>{{harvtxt|Grüter|2020|pp=235-236}}</ref> [[Termite]]s usually do not attack bees or their food pots. However, they can cause damage to the structure of hive boxes as there are many [[Xylophagy|xylophagous]] species. While termites do not usually pose major problems for beekeepers, they should still be monitored closely.<ref>{{harvtxt|Nogueira-Neto|1997|pp=368-370}}</ref><ref>{{harvtxt|Grüter|2020|p=236}}</ref>[[File:Hermetia illucens Black soldier fly edit1.jpg|thumb|[[Hermetia illucens|Black soldier fly.]]]] [[File:Caixa racional individual.png|thumb|166x166px|Rational box support with burnt oil to prevent ant attacks.]] [[Ant]]s are attracted to bee colonies by the smell of food. To prevent ant attacks, it's important to handle the hive boxes carefully and avoid exposing jars of pollen and honey. Although rare, when attacks do occur, there are intense conflicts between ants and bees. Stingless bees usually manage to defend themselves, but the damage to the bee population can be significant. To prevent ant infestations in meliponaries with individual supports, a useful strategy is to impregnate the box supports with burnt oil.<ref name=":92">{{harvtxt|Villas-Bôas|2018|p=106}}</ref><ref>{{harvtxt|Grüter|2020|p=235}}</ref> Another group of enemy flies are the [[Hermetia illucens|black soldier flies]] (''Hermetia illucens''). They lay their eggs in crevices of boxes and can extend the tip of their abdomen during laying, facilitating access to the inside of the hive. Larvae of this species feed on pollen, feces, and other materials found in colonies. In general, healthy bee colonies can coexist peacefully with soldier flies. However, in areas where these insects are prevalent, beekeepers must remain vigilant and protect the gaps in the colonies to prevent potential issues.<ref>{{harvtxt|Villas-Bôas|2018|p=105}}</ref> [[File:Entrada de ninho de Lestrimelitta tropica.jpg|left|thumb|''Lestrimelitta limao'' hive entrance.]] Cleptobiosis, also known as [[Kleptoparasitism|cleptoparasitism]], is a behaviour observed in various species of stingless bees, with over 30 identified species engaging in nest attacks, including honey bee nests. This behaviour serves the purpose of either resource theft or usurping the nest by swarming into an already occupied cavity and these bees are called robber bees. The Neotropical genus ''[[Lestrimelitta]]'' and the African genus ''[[Cleptotrigona]]'' represent bees with an obligate cleptobiotic lifestyle since they do not visit flowers for nectar or pollen. Furthermore, other species such as ''[[Melipona fuliginosa]]'', ''[[Oxytrigona tataira]],'' ''[[Trigona hyalinata]]'', ''[[Trigona spinipes|T. spinipes]]'', and ''[[Tetragona clavipes]]'' are reported to have comparable habits of pillaging and invading, which emphasises the variety of strategies employed by stingless bees in acquiring resources. Other enemies include: jumping spiders ([[Salticidae]]), moths, assassin bugs ([[Reduviidae]]), beetles, parasitoid wasps, predatory mites (''[[Amblyseius]]''), mantises ([[Mantodea]]), robber flies ([[Asilidae]]), etc.<ref>{{harvtxt|Grüter|2020|pp=236-238}}</ref> ==== Vertebrates ==== Human activities pose the most significant threat to stingless bees, whether through honey and nest removal, [[habitat destruction]], [[pesticide]] use or introduction of non-native competitors. Large-scale environmental alterations, particularly the conversion of natural habitats into urban or intensively [[Agriculture|farmed land]], are the most dramatic threats leading to habitat loss, reduced nest densities, and [[Extinction|species disappearance]].<ref name=":18">{{harvtxt|Grüter|2020|pp=238-239}}</ref> [[Primate]]s, including [[chimpanzee]]s, [[gorilla]]s, [[baboon]]s, and various monkey species, are known to threaten stingless bee colonies. [[Elephant]]s, [[honey badger]]s, [[sun bear]]s, [[spectacled bear]]s, [[anteater]]s, [[hog-nosed skunk]]s, [[armadillo]]s, [[tayra]]s, [[eyra cat]]s, [[kinkajou]]s, [[grisons]], and [[coyote]]s are among the mammals that consume or destroy stingless bee nests. Some, like the tayra and eyra cat, have specific preferences for stealing honey. [[Gecko]]s, [[lizard]]s, and [[toad]]s also pose threats by hunting adult bees or consuming workers at nest entrances. [[Woodpecker]]s and various bird species, including [[bee-eater]]s, [[woodcreeper]]s, [[drongo]]s, [[jacamar]]s, [[heron]]s, [[kingbird]]s, [[Flycatcher-shrike|flycatchers]], [[Swift (bird)|swifts]], and [[honeyeater]]s, occasionally prey on stingless bees. African [[honeyguide]]s have developed a mutualism with human honey-hunters, actively guiding them to bee nests for honey extraction and then consuming leftover wax and larvae.<ref name=":18" /> ==== Defense ==== Being [[tropics|tropical]], stingless bees are active all year round, although they are less active in cooler weather, with some species presenting [[diapause]].<ref name=":19">{{harvtxt|Grüter|2020|pp=248-260}}</ref><ref>{{harvtxt|Ribeiro|2002}}</ref><ref>{{harvtxt|Alves|Imperatriz-Fonseca|Santos-Filho|2009}}</ref> Unlike other eusocial bees, they do not sting, but will defend by biting if their nest is disturbed. In addition, a few (in the genus ''[[Oxytrigona]]'') have mandibular secretions, including [[formic acid]], that cause painful blisters.<ref name=":3" /><ref>{{harvtxt|Roubik|Smith|Carlson|1987}}</ref> Despite their lack of a [[sting (biology)|sting]], stingless bees, being eusocial, may have very large colonies made formidable by the number of defenders.<ref>{{harvtxt|Roubik|2006}}</ref><ref>{{harvtxt|Sarchet|2014}}</ref> Stingless bees use other sophisticated defence tactics to protect their colonies and ensure their survival. One important strategy is to choose nesting habitats with fewer natural enemies to reduce the risk of attacks. In addition, they use [[camouflage]] and [[mimicry]] to blend into their surroundings or imitate other animals to avoid detection. An effective strategy is to nest near colonies that provide protection, using collective strength to defend against potential invaders.<ref name=":19" /> Nest entrance guards play a vital role in colony defense by actively preventing unauthorized entry through attacking intruders and releasing alarm [[pheromone]]s to recruit additional defenders. It is worth noting that nest guards often carry sticky substances, such as resins and wax, in their corbiculae or mandibles. Stingless bees apply substances to attackers to immobilise them, thus thwarting potential threats to the colony. Some species (''Tetragonisca angustula'' and ''Nannotrigona testaceicornis'', for example) also close their nest entrances with a soft and porous layer of cerumen at night, further enhancing colony security during vulnerable periods. These intricate defence mechanisms demonstrate the adaptability and resilience of stingless bees in safeguarding their nests and resources.<ref name=":19" /> === Role differentiation === In a simplified sense, the sex of each bee depends on the number of [[chromosome]]s it receives. Female bees have two sets of chromosomes ([[diploid]])—one set from the [[queen (bee)|queen]] and another from one of the male bees or [[drone (bee)|drone]]s. Drones have only one set of chromosomes ([[haploid]]), and are the result of unfertilized eggs, though inbreeding can result in diploid drones. Unlike true honey bees, whose female bees may become workers or queens strictly depending on what kind of food they receive as larvae (queens are fed [[royal jelly]] and workers are fed pollen), the caste system in meliponines is variable, and commonly based simply on the amount of pollen consumed; larger amounts of pollen yield queens in the genus ''[[Melipona]]''. Also, a genetic component occurs, however, and as much as 25%<ref>{{harvtxt|Kerr|1950}}</ref> (typically 5–14%) of the female brood may be queens. Queen cells in the former case can be distinguished from others by their larger size, as they are stocked with more pollen, but in the latter case, the cells are identical to worker cells, and scattered among the worker brood. When the new queens emerge, they typically leave to mate, and most die.<ref name=":28">{{harvtxt |Wenseleers|Ratnieks|Ribeiro|Alves|Imperatriz-Fonseca|2005}}</ref> New nests are not established by swarms, but by a procession of workers that gradually construct a new nest at a secondary location. The nest is then joined by a newly mated queen, at which point many workers take up permanent residence and help the new queen raise her own workers. If a ruling queen is herself weak or dying, then a new queen can replace her. For ''[[Plebeia quadripunctata|Schwarziana quadripunctata]]'', although fewer than 1% of female worker cells produce dwarf queens, they comprise six of seven queen bees, and one of five proceed to head colonies of their own. They are reproductively active, but less fecund than large queens.<ref name=":28" /> == Interaction with humans == === Pollination === {{Main|Pollination}} [[File:Bee pollinating a rose.jpg|thumb|Solitary bee visiting and pollinating a flower.]] Bees play a critical role in the ecosystem, particularly in the pollination of natural vegetation. This activity is essential for the reproduction of various plant species, particularly in [[tropical forest]]s where most tree species rely on [[Entomophily|insect pollination]]. Even in temperate climates, where [[Anemophily|wind pollination]] is prevalent among forest trees, many bushes and herbaceous plants, rely on bees for pollination. The significance of bees extends to arid regions, such as [[Deserts and xeric shrublands|desertic and xeric shrublands]], where bee-pollinated plants are essential for preventing [[erosion]], supporting [[wildlife]], and ensuring [[Ecological stability|ecosystem stability]].<ref name=":23">{{harvtxt|Michener|2000|pp=4-5}}</ref> The impact of bee pollination on agriculture is substantial. In the late 1980s, certain plants were estimated to contribute between $4.6 to $18.9 billion to the U.S. economy, primarily through insect-pollinated crops. Although some bee-pollinated plants can self-pollinate in the absence of bees, the resulting crops often suffer from inbreeding depression. The quality and quantity of seeds or fruits are significantly enhanced when bees participate in the pollination process. Although estimates of crop pollination attributed to honey bees are uncertain, it is undeniable that bee pollination is a vital and economically valuable activity.<ref name=":23" /> Ramalho (2004) demonstrates that stingless bees amount to approximately 70% of all bees foraging on flowers in the Brazilian Tropical [[Atlantic Forest|Atlantic Rainforest]] even though they represented only 7% of all bee species.<ref>{{harvtxt|Ramalho|2004}}</ref> In a habitat in Costa Rica, stingless bees accounted for 50% of the observed foraging bees, despite representing only 16% of the recorded bee species.<ref>{{harvtxt|Brosi|Daily|Shih|Oviedo|Durán|2008}}</ref> Following this pattern, Cairns ''et al''. (2005) found that 52% of all bees visiting flowers in Mexican habitats were meliponines.<ref>{{harvtxt|Cairns|Villanueva-Gutiérrez|Koptur|Bray|2005}}</ref> Meliponine bees play a crucial role in tropical environments due to their high population rate, morphological diversity, diverse foraging strategies, generalist foraging habits ([[Polylectic|polylecty]]), and flower constancy during foraging trips. Nest densities and colony sizes can result in over a million individual stingless bees inhabiting a square kilometre of tropical habitat. Due to their diverse morphology and behaviour, bees are capable of collecting pollen and nectar from a wide range of flowering plants. Key plant families are reported as most visited by meliponines: [[Fabaceae]], [[Euphorbiaceae]], [[Asteraceae]] and [[Myrtaceae]].<ref name=":20">{{harvtxt|Grüter|2020|pp=323-330}}</ref> Grüter compiled some studies about twenty crops that substantially benefit from SB pollination (following table) and also lists seventy-four crops that are at least occasionally or potentially pollinated by stingless bees.<ref name=":20" /> {| class="wikitable mw-collapsible" |+ !Common name !Scientific name !Family !Pollinator genus !Reference |- |Annatto, achiote |''[[Bixa orellana]]'' |[[Bixaceae]] |''[[Melipona]]'' |<ref name=":21">{{harvtxt|Heard|1999}}</ref> |- |Aubergine |''[[Solanum melongena|Solanum melogena]]'' |[[Solanaceae]] |''Melipona'' |<ref>{{harvtxt|Nunes-Silva|Hrncir|da Silva|Roldão|Imperatriz-Fonseca|2013}}</ref> |- |Avocado |''[[Persea americana]]'' |[[Lauraceae]] |''[[Nannotrigona]], [[Trigona]]'' |<ref name=":21" /> |- |Camu-camu |''[[Myrciaria dubia]]'' |[[Myrtaceae]] |''Melipona, [[Scaptotrigona]]'' |<ref name=":21" /> |- |Carambola |''[[Averrhoa carambola]]'' |[[Oxalidaceae]] |''Trigona'' |<ref name=":21" /> |- |Chayote, choko |''[[Sechium edule]]'' |[[Cucurbitaceae|Cucubitaceae]] |''Trigona, [[Partamona]]'' |<ref name=":21" /> |- |Coconut |''[[Cocos nucifera]]'' |[[Arecaceae]] |various genera |<ref name=":21" /> |- |Coffee |''[[Coffea arabica]]'' |[[Rubiaceae]] |''[[Lepidotrigona]], Trigona'' |<ref name=":21" /><ref name=":22">{{harvtxt|Slaa|Chaves|Malagodi-Braga|Hofstede|2006}}</ref> |- |Coffee |''[[Coffea canephora]]'' |Rubiaceae |''Lepidotrigona, Trigona'' |<ref name=":22" /> |- |Cucumber |''[[Cucumis sativus]]'' |Curcubitaceae |''Nannotrigona, Scaptotrigona'' |<ref name=":21" /><ref name=":22" /> |- |Cupuaçu |''[[Theobroma grandiflorum]]'' |[[Malvaceae]] |''Trigona'' |<ref name=":21" /> |- |Macadamia |''[[Macadamia integrifolia]]'' |[[Proteaceae]] |''Trigona'' |<ref name=":21" /> |- |Mango |''[[Mangifera indica]]'' |[[Anacardiaceae]] |''Trigona'' |<ref name=":21" /> |- |Mapati, uvilla |''[[Pourouma cecropiifolia]]'' |[[Moraceae]] |''[[Oxytrigona]], Trigona'' |<ref name=":21" /> |- |Mealy sage |''[[Salvia farinacea]]'' |[[Lamiaceae]] |''Nannotrigona, [[Tetragonisca]]'' |<ref name=":22" /> |- |Rambutan |''[[Nephelium lappaceum]]'' |[[Sapindaceae]] |''Scaptotrigona'' |<ref name=":21" /><ref name=":22" /> |- |Rockmelon |''[[Cucumis melo]]'' |Curcubitaceae |''[[Heterotrigona]]'' |<ref>{{harvtxt|Azmi|Wan Sembok|Yusuf|Mohd Hatta|Salleh|Hamzah|Ramli|2019}}</ref> |- |Strawberry |''[[Fragaria]]'' sp. |[[Rosaceae]] |various genera |<ref name=":21" /><ref name=":22" /> |- |Sweet pepper |''[[Capsicum annuum]]'' |Solanaceae |''[[Austroplebeia]], Melipona, [[Tetragonula]]'' |<ref name=":21" /><ref name=":22" /><ref>{{harvtxt|Putra|Salmah|Swasti|2017}}</ref> |- |Tomato |''[[Lycopersicon esculentum|Solanum lycopersicum]]'' |Solanaceae |''Melipona, Nannotrigona'' |<ref name=":22" /> |} == Worldwide overview == === Africa === Stingless bees also play a vital ecological role across [[Sub-Saharan Africa]] and [[Madagascar]]. To understand these insects on the African continent, it's important to consider the prevailing socio-economic and cultural contexts. Despite their ecological significance, the diversity, conservation, and behavior of these bees remain underexplored, particularly compared to better-studied regions such as [[South America]] and [[Southeast Asia]]. Also, honeybees were extensively researched, in contrast to native meliponines.<ref name=":31">{{harvtxt|Kajobe|2008}}</ref><ref name=":32">{{harvtxt|Byarugaba|2004}}</ref> Africa is home to seven [[Biodiversity hotspot|biodiversity hotspots]], yet the recorded bee fauna is moderate relative to the continent's size. Madagascar stands out with exceptionally high levels of endemic species, though much of the bee diversity remains undocumented.<ref name=":30">{{harvtxt|Eardley|Gikungu|Schwarz|2009}}</ref>'''<ref name=":31" />''' Africa is home to aproximately 36 species of meliponines, including seven endemic to Madagascar. Most of these bees are found in equatorian regions (tropical forests and some savannahs).<ref>{{harvtxt|Grüter|2010|pp=47 & 49}}</ref> Factors such as habitat destruction, pesticide use, and invasive species pose significant threats to these pollinators. Furthermore, high rates of nest mortality, driven by predation and human activity, exacerbate conservation challenges. Research indicates that stingless bees in Africa face greater pressures than their counterparts in the American and Asian tropics, underlining the urgency for targeted conservation measures.<ref name=":30" />'''<ref name=":31" />''' Uganda's [[Bwindi Impenetrable National Park]] has shown the presence of at least five stingless bee species in, distributed across two genera: ''[[Meliponula]]'' and ''[[Hypotrigona]]''.<ref name=":31" />'''<ref name=":32" />''' In Madagascar, there is only one genus of stingless bees: ''[[Liotrigona]].''<ref name=":30" /> Meliponiculture, for example, is practised in Angola and Tanzania, and interest in managing stingless bees is growing in other African countries as well. <ref>{{harvtxt|Grüter|2010|p=27}}</ref> === Australia === Of the 1,600 species of wild bees native to Australia, about 14 are meliponines.<ref>{{cite news | url=http://www.abc.net.au/science/articles/2003/05/08/2045526.htm | title=Stingless bee rescue |author= Wendy Pyper |date=May 8, 2003 |work=ABC Science}}</ref> "''Coot-tha''", which derives from "''ku-ta''", is one of the Aboriginal names for "wild stingless bee honey".<ref>{{harvtxt|Vit|Pedro|Roubik|2018}}</ref> These species bear a variety of names, including Australian native honey bees, native bees, sugar-bag bees, and sweat bees (because they land on people's skin to collect sweat).<ref>{{Cite web |title=Australian Native Stingless Bees |url=https://www.aussiebee.com.au/australian-stingless-bees.html |access-date=2024-05-28 |website=www.aussiebee.com.au}}</ref> The various stingless species look quite similar, with the two most common species, ''[[Tetragonula carbonaria]]'' and ''[[Austroplebeia australis]]'', displaying the greatest variation, as the latter is smaller and less active. Both of these inhabit the area around [[Brisbane]].<ref>{{Cite web |last=Steve |date=2015-05-25 |title=Native Stingless Bees - Tetragonula carbonaria |url=https://www.nativebeehives.com/native-stingless-bees-tetragonula-carbonaria/ |access-date=2024-05-28 |website=www.nativebeehives.com |language=en-AU}}</ref> As stingless bees are usually harmless to humans, they have become an increasingly attractive addition to the suburban backyard. Most meliponine beekeepers do not keep the bees for [[honey]], but rather for the pleasure of conserving native species whose original [[habitat (ecology)|habitat]] is declining due to human development. In return, the bees [[pollinate]] crops, garden flowers, and bushland during their search for nectar and pollen. While a number of beekeepers fill a small niche market for bush honey, native meliponines only produce small amounts and the structure of their hives makes the honey difficult to extract. Only in warm areas of Australia such as [[Queensland]] and northern [[New South Wales]] are favorable for these bees to produce more honey than they need for their own survival. Most bees only come out of the hive when it is above about 18°C (64°F).<ref>{{Cite web|url=https://www.abc.net.au/news/2019-07-05/moves-underway-to-make-a-standard-for-native-bee-honey/11278346|title=Native bee honey set to have its own food standard|last=Thomas|first=Kerrin|date=2019-07-05|website=ABC News|language=en-AU|access-date=2019-07-10}}</ref> Harvesting honey from a nest in a cooler area could weaken or even kill the nest. ==== Pollination ==== Australian farmers rely almost exclusively on the introduced [[western honey bee]] to pollinate their crops. However, native bees may be better pollinators for certain agricultural crops. Stingless bees have been shown to be valuable pollinators of tropical plants such as [[macadamia]]s and [[mango]]s.<ref name=":21" /> Their foraging may also benefit strawberries, watermelons, citrus, avocados, [[lychee]]s, and many others.<ref name=":21" /><ref name=":22" /> Research into the use of stingless bees for crop pollination in Australia is still in its very early stages, but these bees show great potential. Studies at the [[University of Western Sydney]] have shown these bees are effective pollinators even in confined areas, such as glasshouses.<ref>{{cite journal |date=May 1999 |title=New Greenhouse Pollination Study With ''Trigona'' |url=http://www.aussiebee.com.au/ab10.html |journal=Aussie Bee Bulletin |issue=10 |quote=Pablo Occhiuzzi of the University of Western Sydney is studying the greenhouse pollination of capsicum with ''Trigona carbonaria''.}}</ref> === Brazil === {{see also|List of stingless bees of Brazil}} [[File:Abelhas sem ferrao brasil.png|thumb|Number of valid species of stingless bees in Brazil. State and region cut-outs. Study published in 2023.<ref name=":27" />]] [[File:Pollinating bee covered with pollen.jpg|thumb|''[[Trigona spinipes]],'' covered with [[pollen]], visiting a flower of the vegetable sponge gourd (''Luffa cylindrica'') in [[Campinas]], [[Brazil]].]] Brazil is home to several species bees belonging to Meliponini, with more than 300 species already identified and probably more yet to be discovered and described. They vary greatly in shape, size, and habits, and 20 to 30 of these species have good potential as honey producers. Although they are still quite unknown by most people, an increasing number of beekeepers (''meliponicultores'', in Portuguese) have been dedicated to these bees throughout the country.<ref name=":27" /><ref>{{harvtxt|Villas-Bôas|2018|p=17}}</ref> This activity has experienced significant growth since August 2004, when national laws were changed to allow native bee colonies to be freely marketed, which was previously forbidden in an unsuccessful attempt to protect these species. Nowadays the capture or destruction of existing colonies in nature is still forbidden, and only new colonies formed by the bees themselves in artificial traps can be collected from the wild.<ref>[https://conama.mma.gov.br/?option=com_sisconama&task=arquivo.download&id=440 CONAMA 2004 Resolution]</ref> Most marketed colonies are artificially produced by authorized beekeepers, through division of already existing captive colonies. Besides honey production, Brazilian stingless bees such as the jataí (''[[Tetragonisca angustula]]''), mandaguari (''[[Scaptotrigona postica]]''), and mandaçaia (''[[Melipona quadrifasciata]]'') serve as major pollinators of tropical plants and are considered the ecological equivalent of the honey bee.<ref name=":21" /><ref name=":22" /> Also, much practical and academic work is being done about the best ways of keeping such bees, multiplying their colonies, and exploring the honey they produce.<ref>{{Cite web |date=2024-07-26 |title=Arquivos Artigos - A.B.E.L.H.A. |url=https://abelha.org.br/artigos/ |access-date=2024-12-31 |website=abelha.org.br |language=pt-BR}}</ref> Among many others, species such as jandaíra (''[[Melipona subnitida]]'') and true ''uruçu'' (''[[Melipona scutellaris]]'') in the northeast of the country, mandaçaia (''[[Melipona quadrifasciata]]'') and yellow uruçu (''Melipona rufiventris'') in the south-southeast, tiúba or jupará (''[[Melipona interrupta]]'') and canudo (''[[Scaptotrigona polysticta]]'') in the north and jataí (''[[Tetragonisca angustula]]'') throughout the country are increasingly kept by small, medium, and large producers. Many other species as the mandaguari (''[[Scaptotrigona postica]]''), the guaraipo (''[[Melipona bicolor]]''), marmelada (''[[Frieseomelitta varia]]'') and the iraí (''[[Nannotrigona testaceicornis]]''), to mention a few, are also reared.<ref name=":33">{{Cite web |date=2022-02-21 |title=Fichas catalográficas das espécies relevantes para a meliponicultura - A.B.E.L.H.A. |url=https://abelha.org.br/fichas-catalograficas-das-especies-relevantes-para-a-meliponicultura-2/ |access-date=2024-12-31 |website=abelha.org.br |language=pt-BR}}</ref> According to [[Chico Mendes Institute for Biodiversity Conservation|ICMBio]] and the [[Ministry of Environment and Climate Change (Brazil)|Ministry of the Environment]] there are presently four species of Meliponini listed in the [[IUCN Red List|National Red List of Threatened Species]] in Brazil. ''[[Melipona capixaba]]'', ''[[Melipona rufiventris]]'', ''[[Melipona scutellaris]]'', and ''[[Partamona littoralis]]'' all listed as Endangered (EN).<ref>{{Cite web |title=IBAMA |url=https://www.ibama.gov.br/component/legislacao/?view=legislacao&force=1&legislacao=134521#:~:text=Ementa:,Extin%C3%A7%C3%A3o%20-%20Peixes%20e%20Invertebrados%20Aqu%C3%A1ticos. |access-date=2024-02-26 |website=www.ibama.gov.br}}</ref><ref>[[Instituto Chico Mendes de Conservação da Biodiversidade|ICMBio]]. 2018. [https://www.gov.br/icmbio/pt-br/centrais-de-conteudo/publicacoes/publicacoes-diversas/livro_vermelho_2018_vol7.pdf Livro Vermelho da Fauna Brasileira Ameaçada de Extinção: Volume VII - Invertebrados]. ''In'': ICMBio. (Org.). [https://www.gov.br/icmbio/pt-br/centrais-de-conteudo/publicacoes/publicacoes-diversas/livro_vermelho_2018_vol1.pdf Livro Vermelho da Fauna Brasileira Ameaçada de Extinção]. Brasília: ICMBio. 727p.</ref> ==== Honey production ==== Although the colony population of most of these bees is much smaller than that of European bees, the productivity per bee can be quite high. Interestingly, honey production is more connected to the body size, not the colony size. The manduri (''[[Melipona marginata]]''), jandaíra (''[[Melipona subnitida]]'') and the ''guaraipo'' (''[[Melipona bicolor|M. bicolor]]'') live in swarms of only around 300 individuals but can still produce up to 5 liters (1.3 US gallon) of honey a year under the right conditions.<ref>{{harvtxt|Grüter|2010|p=14}}</ref> In large bee farms, only the availability of flowers limits the honey production per colony. However, much larger numbers of beehives are required to produce amounts of honey comparable to that of European bees. Also, due to the fact of those bees storing honey in cerumen pots instead of standardized honeycombs as in the honeybee rearing makes extraction a lot more difficult and laborious.<ref>{{harvtxt|Fonseca|Sodré|Carvalho|Alves|Souza|Silva|Oliveira|Machado|Clarton|2007}}</ref> The honey from stingless bees has a higher water content, from 25% to 35%, compared to the honey from the genus ''Apis''. This contributes to its less cloying taste but also causes it to spoil more easily. Thus, for marketing, this honey needs to be processed through desiccation, fermentation or pasteurization. In its natural state, it should be kept under refrigeration.<ref>{{harvtxt|Grüter|2020|pp=12-14}}</ref> ==== Bees as pets ==== [[Image:Mandaçaias.jpg|left|thumb|A swarm of mandaçaias bees around an artificial [[beehive]] installed in a house's backyard in [[Brazil]]]] Due to the lack of a functional stinger and characteristic nonaggressive behavior of many Brazilian species of stingless bees, they can be reared without problems in densely populated environments (residential buildings, schools, urban parks), provided enough flowers are at their disposal nearby. Some breeders (''meliponicultores'') can produce honey even in apartments up to the 12th floor.<ref>{{cite web|url=http://meliponariocapixaba.blogspot.com.br/2009/11/e-possivel-criar-abelhas-em.html|title=Meliponario Capixaba: É POSSÍVEL CRIAR ABELHAS EM APARTAMENTOS?|author=João Luiz|access-date=12 March 2016}}</ref> The [[Melipona quadrifasciata|mandaçaias]] (''Melipona quadrifasciata'') are extremely tame, rarely attacking humans (only when their hives are opened for honey extraction or colony division). They form small, manageable colonies of only 400&ndash;600&nbsp;individuals. They are fairly large bees, up to 11&nbsp;mm (7/16") in length, and as a result have better body heat control, allowing them to live in regions where temperatures can drop a little lower than 0&nbsp;°C (32&nbsp;°F). However, they are somewhat selective about which flowers they will visit, preferring the flora that occurs in their natural environment. They are thus difficult to keep outside their region of origin (the eastern coast of Brazil). Once very common, the mandaçaia is now rather rare in nature, mainly due to the destruction of their native forests in the of Brazil.<ref>{{harvtxt|Brito|Faquinello|Paula-Leite|Carvalho|1997}}</ref><ref name=":33" /> Other groups of Brazilian stingless bees, genera ''[[Plebeia]]'' and ''[[Leurotrigona]]'', are also very tame and much smaller, with one of them (''[[Plebeia minima]]'') reaching no more than 2.5&nbsp;mm (3/32") in length, and the ''[[lambe-olhos]]'' ("lick-eyes" bee, ''[[Leurotrigona muelleri]]'') being even smaller, at no more than 1.5&nbsp;mm (3/32"). Many of these species are known as mirim (meaning 'small' in the [[Tupi-Guarani languages]]). As a result, they can be kept in very small artificial hives, thus being of interest for keepers who want them as pollinators in small glasshouses or just for the pleasure of having a 'toy' bee colony at home.<ref name=":33" /><ref>{{Cite web |title=Lambe-olhos – Leurotrigona muelleri – Laboratório de Sistemática de Plantas |url=https://sites.usp.br/jardimdabotanicausprp/lambe-olhos-leurotrigona-muelleri/ |access-date=2024-12-31 |website=sites.usp.br}}</ref><ref>{{Cite web |title=Mirim – Plebeia droryana – Laboratório de Sistemática de Plantas |url=https://sites.usp.br/jardimdabotanicausprp/mirim-plebeia-droryana/ |access-date=2024-12-31 |website=sites.usp.br}}</ref> Being so tiny, these species produce only a very small amount of honey, typically less than 500&nbsp;ml (1/2 US pint) a year, so are not interesting for commercial honey production. [[Image:Abelha-jatai.jpg|right|thumb|Entrance pipe of ''jataís´s'' colony: The ''jataí'' bees build a wax pipe in the entrance of their nest.]] Belonging to the same group, the jataí (''[[Tetragonisca angustula]]''), the marmelada (''[[Frieseomelitta varia]]''), and the moça-branca (''[[Frieseomelitta doederleini]]'') are intermediate in size between those very small species and the European bee. They are very adaptable species; the jataí, and can be reared in many different regions and environments, being quite common in most Brazilian cities. The jataí can bite when disturbed, but its jaws are weak, and in practice they are harmless, while the marmelada and moça-branca usually deposit propolis on their aggressors. Jataí is one of the first species to be kept by home beekeepers. Their nests can be easily identified in trees or wall cavities by the yellow wax pipe they build at the entrance, usually guarded by some soldier bees, which are stronger than regular worker bees. The marmelada and moça-branca make a little less honey, but it is denser and sweeter than most from other stingless bees and is considered very tasty.<ref name=":33" /><ref>{{harvtxt|Castanheira|Contel|2005|}}</ref> === Central America === [[Image:Colmena melipona.jpg|right|thumb|A Maya stingless bee hive: A piece of hollow log provides a home for meliponine bees in [[Belize]].]] The stingless bees ''[[Melipona beecheii]]'' and ''[[Melipona yucatanica|M. yucatanica]]'' are the primary native bees cultured in Central America, though a few other species are reported as being occasionally managed (e.g., ''[[Trigona fulviventris]]'' and ''[[Scaptotrigona mexicana]]'').<ref>{{harvtxt|Kent|1984}}</ref> They were extensively cultured by the [[Maya civilization]] for honey, and regarded as sacred. They continue to be cultivated by the modern [[Maya peoples]], although these bees are endangered due to massive deforestation, altered [[agriculture|agricultural]] practices (especially overuse of [[insecticide]]s), and changing beekeeping practices with the arrival of the [[Africanized honey bee]], which produces much greater honey crops.<ref name=":26">{{harvtxt|Villanueva|Roubik|Colli-Ucán|1998}}</ref> ==== History ==== Native meliponines (''M. beecheii'' being the most common) have been kept by the lowland Maya for thousands of years. The [[Yucatec Maya language]] name for this bee is ''xunan kab'', meaning "(royal, noble) lady bee".<ref>{{harvtxt|Grüter|2010|p=54}}</ref><ref>{{cite web|title=Diccionario Introductorio|url=http://www.uqroo.mx/libros/maya/diccionario.pdf|work=uqroo.mx|publisher=Universidad De Quintana Roo|access-date=27 March 2014|language=es}}</ref> The bees were once the subject of religious ceremonies and were a symbol of the bee-god [[Ah-Muzen-Cab]], known from the ''[[Madrid Codex (Maya)|Madrid Codex]]''.<ref name=":34">{{Cite web |date=2016-07-17 |title=People and bees. Mayan bees in the Madrid Codex |url=https://ecomercioagrario.com/en/people-and-bees-mayan-bees-in-the-madrid-codex/ |access-date=2024-12-31 |website=eComercio Agrario |language=en-US}}</ref> The bees were, and still are, treated as pets. Families would have one or many log-hives hanging in and around their houses. Although they are stingless, the bees do bite and can leave welts similar to a mosquito bite. The traditional way to gather bees, still favored among the locals, is find a wild hive, then the branch is cut around the hive to create a portable log, enclosing the colony. With proper maintenance, hives have been recorded as lasting over 80 years, being passed down through generations. In the archaeological record of Mesoamerica, stone discs have been found that are generally considered to be the caps of long-disintegrated logs that once housed the beehives.<ref name=":34" /><ref name=":35">{{Cite web |last=Kennedy |first=Jennifer |last2=Arghiris |first2=Richard |date=2019-01-31 |title=House of the Royal Lady Bee: Maya revive native bees and ancient beekeeping |url=https://news.mongabay.com/2019/01/house-of-the-royal-lady-bee-maya-revive-native-bees-and-ancient-beekeeping/ |access-date=2024-12-31 |website=Mongabay Environmental News |language=en-US}}</ref> ==== Tulum ==== [[Tulum]], the site of a [[pre-Columbian]] Maya city on the Caribbean coast 130&nbsp;km (81&nbsp;mi) south of Cancun, has a god depicted repeatedly all over the site. Upside down, he appears as a small figure over many doorways and entrances. One of the temples, the Temple of the Descending God (''Templo del Dios Descendente''), stands just left of the central plaza. Speculation is that he may be the "Bee God", Ah Muzen Cab, as seen in the ''Madrid Codex''. It is possible that this was a religious/trade center with emphasis on ''xunan kab'', the "royal lady".<ref name=":35" /> ==== Economic uses ==== [[Balché]], a traditional [[Mesoamerica]]n alcoholic beverage similar to [[mead]], was made from fermented honey and the bark of the leguminous'' balché'' tree (''[[Lonchocarpus violaceus]]''), hence its name. It was traditionally brewed in a canoe. The drink was known to have [[entheogen]]ic properties, that is, to produce mystical experiences, and was consumed in medicinal and ritual practices. Beekeepers would place the nests near the psychoactive plant ''[[Turbina corymbosa]]'' and possibly near ''balché'' trees, forcing the bees to use nectar from these plants to make their honey. Additionally, brewers would add extracts of the bark of the ''balché'' tree to the honey mixture before fermentation. The resulting beverage is responsible for psychotropic effects when consumed, due to the ergoline compounds in the pollen of the ''T. corymbosa'', the ''Melipona'' nectar gathered from the ''balché'' flowers, or the hallucinogenic compounds of the ''balché'' tree bark.<ref>{{harvtxt|Ott|1998}}</ref> [[Lost-wax casting]], a common metalworking method typically found where the inhabitants keep bees, was also used by the Maya. The wax from ''Melipona'' is soft and easy to work, especially in the humid Maya lowland. This allowed the Maya to create smaller works of art, jewelry, and other metalsmithing that would be difficult to forge. It also makes use of the leftovers from honey extraction. If the hive was damaged beyond repair, the whole of the comb could be used, thus using all of the hive. With experienced keepers, though, only the honey pot could be removed, the honey extracted, and the wax used for casting or other purposes.<ref>{{harvtxt|Pitses|2018|}}</ref><ref>{{harvtxt|Schwarz|1945|}}</ref> ==== Future ==== The outlook for meliponines in Mesoamerica is uncertain. The number of active Meliponini beekeepers is shy in comparison with the Africanized ''[[Apis mellifera]]'' breeders. The high honey yield, 100&nbsp;kg (220&nbsp;lbs) or more annually, along with the ease of hive care and ability to create new hives from existing stock, commonly outweighs the negative consequences of "killer bee" hive maintenance.<ref name=":36">{{harvtxt|Villanueva-G|Roubik|Colli-Ucán|2005|}}</ref> An additional blow to the art of meliponine beekeeping is that many of the ''meliponicultores'' are now elderly, and their hives may not be cared for once they die. The hives are considered similar to an old family collection, to be parted out once the collector dies or to be buried in whole or part along with the beekeeper upon death. In fact, a survey of a once-popular area of the Maya lowlands shows the rapid decline of beekeepers, down to around 70 in 2004 from thousands in the late 1980s. Conservation efforts are underway in several parts of Mesoamerica.<ref name=":36" /><ref>A comprehensive conservation guide can be found in the June 2005 issue of ''Bee World''.</ref> == References ==<!-- ActaAmazonica34 p333 --> {{Reflist|2}} ==Bibliography== === Articles and publications === * {{cite journal |last1=Alves |first1=D A |last2=Imperatriz-Fonseca |first2=V L |last3=Santos-Filho |first3=P S. |year=2009 |title=Production of workers, queens and males in ''Plebeia remota'' colonies (Hymenoptera, Apidae, Meliponini), a stingless bee with reproductive diapause |journal=Genetics and Molecular Research |volume=8 |issue=2 |pages=672–683 |doi=10.4238/vol8-2kerr030 |pmid=19554766 |doi-access=free}} * {{Cite journal |last1=Antonini |first1=Yasmine |last2=Martins |first2=Rogério P. |date=2003 |title=The value of a tree species (''Caryocar brasiliense'') for a stingless bee ''Melipona quadrifasciata quadrifasciata'' |url=http://link.springer.com/10.1023/A:1027378306119 |journal=Journal of Insect Conservation |volume=7 |issue=3 |pages=167–174 |doi=10.1023/A:1027378306119|bibcode=2003JICon...7..167A |s2cid=6080884 }} * {{Cite journal |last1=Azmi |first1=Wahizatul Afzan |last2=Wan Sembok |first2=W Z |last3=Yusuf |first3=N |last4=Mohd. 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B. |date=2002 |publisher=Silveira |isbn=978-85-903034-1-1 |edition=1st |location=Belo Horizonte}} * {{cite journal |last1=Venturieri |first1=G C. |year=2004 |title=Meliponicultura: Criação de Abelhas Indígenas Sem Ferrão |url=https://www.infoteca.cnptia.embrapa.br/infoteca/bitstream/doc/410121/1/livroabelha2014.pdf |journal=Comunicado Técnico Embrapa Amazônia Oriental |volume=118 |pages=1–4}} * Villas-Bôas, Jerônimo (2018). ''[https://ispn.org.br/site/wp-content/uploads/2018/10/ManualTecnologicoMel.pdf Manual Tecnológico de Aproveitamento Integral dos Produtos das Abelhas Nativas sem Ferrão]''. Brasília: Instituto Sociedade, População e Natureza (ISPN), v. 2. * {{Cite book |last1=Vit |first1=Patricia |url=https://books.google.com/books?id=NJxPDwAAQBAJ&dq=%22australia%22+%22native%22+%22stingless+bee%22&pg=PA401 |title=Pot-Pollen in Stingless Bee Melittology |last2=Pedro |first2=Silvia R. M. |last3=Roubik |first3=David W. |date=2018-03-05 |publisher=Springer |isbn=978-3-319-61839-5 |language=en}}< ==External links== {{Commons category|Meliponini}} {{Wikispecies|Meliponini|''Meliponini''}} *[http://www.ibra.org.uk/ International Bee Research Association] *[https://web.archive.org/web/20140215065345/http://www.ufv.br/dbg/bee/Versao2/stinglessbees.htm Stingless honey bees of Brazil] *[https://www.sciencedaily.com/releases/2005/06/050615062105.htm Mayan Stingless Bee Keeping: Going, Going, Gone?] *[http://green.blogs.nytimes.com/2012/02/17/a-different-kind-of-beekeeping-takes-flight/?_r=0 A Different Kind of Beekeeping Takes Flight] {{Taxonbar|from=Q127768}} {{Authority control}} [[Category:Meliponini| ]] [[Category:Entomology]] [[Category:Bees]]'
Unified diff of changes made by edit (edit_diff)
'@@ -185,5 +185,5 @@ ==== Soldiers ==== -While the existence of a soldier caste is well known in [[ant]]s and [[termite]]s, the phenomenon was unknown among bees until 2012, when some stingless bees were found to have a similar caste of defensive specialists that help guard the nest entrance against intruders.<ref>{{harvtxt|Grüter|Menezes|Imperatriz-Fonseca|Ratnieks|2012}}</ref> To date, at least 10 species have been documented to possess such "soldiers", including ''[[Tetragonisca angustula]]'', ''[[Tetragonisca fiebrigi|T. fiebrigi]]'', and ''[[Frieseomelitta longipes]]'', with the guards not only larger, but also sometimes a different color from ordinary workers.<ref>{{harvtxt|Grüter|Segers|Menezes|Vollet-Neto|Falcón|von Zuben|Bitondi|Nascimento|Almeida|2017}}</ref><ref>{{harvtxt|Grüter|2020|p=258}}</ref> +While the existence of a soldier caste is well known in [[ant]]s and [[termite]]s, the phenomenon was unknown among bees until 2012, when some stingless bees were found to have a similar caste of defensive specialists that help guard the nest entrance against intruders.<ref>{{harvtxt|Grüter|Menezes|Imperatriz-Fonseca|Ratnieks|2012}}</ref> To date, at least 10 species have been documented to possess such "soldiers", including ''[[Tetragonisca angustula]]'', ''[[Tetragonisca fiebrigi|T. fiebrigi]]'', and ''[[Frieseomelitta longipes]]'', with the guards not only larger, but also sometimes a different color from ordinary workers.<ref>{{harvtxt|Grüter|Segers|Menezes|Vollet-Neto|Falcón|von Zuben|Bitondi|Nascimento|Almeida|2017}}</ref><ref>{{harvtxt|Grüter|2020|p=258}}</ref> ==== Division of labour ==== @@ -199,5 +199,5 @@ Vegetable resin, gathered from a variety of plant species in the wild, is an essential raw material brought back to the hive. Stored in small, sticky clumps in peripheral areas of the colony, it is often mistakenly treated as a synonym for [[propolis]]. However, in beekeeping terminology, propolis refers to a mixture of resin, wax, enzymes, and possibly other substances. Stingless bees go beyond the classic propolis by producing various derivatives from resins and wax, sometimes using pure resins for sealing or defense, a behavior not observed in ''Apis'' bees. Understanding these distinctions is vital for effective production and value addition to the meliponiculture activity.<ref name=":10" /><ref name=":11" /><ref name=":12" /><ref name=":13" /> -[[Honey]], a prized product of bee colonies, is crafted through the processing of [[nectar]]s, [[Honeydew (secretion)|honeydews]], and fruit juices by worker bees. They store these collected substances in an extension of their gut called a [[crop (anatomy)|crop]]. Back at the hive, the bees ripen or dehydrate the nectar droplets by spinning them inside their mouthparts until honey is formed. Ripening concentrates the nectar and increases the sugar content, though it is not nearly as concentrated as the honey from ''Apis mellifera''. Stored in food pots, meliponines' honey is often referred to as pot-honey due to its distinctive storage method. Stingless bee honeys differ from ''A. mellifera'' honey in terms of color, texture, and flavor, being more liquid with a higher water content. Rich in [[mineral]]s, [[amino acid]]s, and [[flavonoid]] compounds, the composition of honey varies among colonies of the same species, influenced by factors such as season, habitat, and collected resources.<ref name=":10" /><ref name=":11" /><ref name=":12" /><ref name=":13" /> +[[Honey]], a prized product of bee colonies, is crafted through the processing of [[nectar]]s, [[Honeydew (secretion)|honeydews]], and fruit juices by worker bees. They store these collected substances in an extension of their gut called a [[crop (anatomy)|crop]]. Back at the hive, the bees ripen or dehydrate the nectar droplets by spinning them inside their mouthparts until honey is formed. Ripening concentrates the nectar and increases the sugar content, though it is not nearly as concentrated as the honey from ''Apis mellifera''. Stored in food pots, meliponines' honey is often referred to as pot-honey due to its distinctive storage method. Stingless bee honeys differ from ''A. mellifera'' honey in terms of color, texture, and flavor, being more liquid with a higher water content. Rich in [[mineral]]s, [[amino acid]]s, and [[flavonoid]] compounds, the composition of honey varies among colonies of the same species, influenced by factors such as season, habitat, and collected resources.<ref name=":10" /><ref name=":11" /><ref name=":12" /><ref name=":13" /> Special methods are being developed to harvest moderate amounts of honey from stingless bees in these areas without causing harm. For honey production, the bees need to be kept in a box specially designed to make the honey stores accessible without damaging the rest of the nest structure. Some recent box designs for honey production provide a separate compartment for the honey stores so the honey pots can be removed without spilling honey into other areas of the nest. Unlike a hive of commercial honeybees, which can produce 75 kg (165 lbs) of honey a year, a hive of Australian stingless bees produces less than 1 kg (2 lbs). Stingless bee honey has a distinctive "bush" taste—a mix of sweet and sour with a hint of fruit.<ref>{{harvtxt|Mduda|Hussein|Muruke|2023}}</ref><ref>{{harvtxt|Ferreira|Lencioni|Benassi|Barth|Bastos|2009}}</ref><ref>{{harvtxt|Sousa|Souza|Marques|Benassi|Gullón|Pintado|Magnani|2016}}</ref> The taste comes from plant resins—which the bees use to build their hives and honey pots—and varies at different times of year depending on the flowers and trees visited. @@ -209,5 +209,5 @@ Stingless bees, as a collective group, display remarkable adaptability to diverse nesting sites. They can be found in exposed nests in trees, from living in ant and termite nests above and below ground to cavities in trees, trunks, branches, rocks, or even human constructions.<ref name=":14">{{harvtxt|Grüter|2020|pp=87-97}}</ref> -Many [[beekeeper]]s keep the bees in their original log [[beehive|hive]] or transfer them to a wooden box, as this makes controlling the hive easier. Some beekeepers put them in bamboos, flowerpots, coconut shells, and other recycling containers such as a water jug, a broken guitar, and other safe and closed containers.<ref>{{harvtxt|Venturieri|2004|pp=36-39}}</ref><ref>{{harvtxt|Contrera|Venturieri|2008}}</ref><ref name=":26" /> +Many [[beekeeper]]s keep the bees in their original log [[beehive|hive]] or transfer them to a wooden box, as this makes controlling the hive easier. Some beekeepers put them in bamboos, flowerpots, coconut shells, and other recycling containers such as a water jug, a broken guitar, and other safe and closed containers.<ref>{{harvtxt|Venturieri|2004|pp=36-39}}</ref><ref>{{harvtxt|Contrera|Venturieri|2008}}</ref><ref name=":26" /> ==== Exposed nests ==== @@ -233,5 +233,5 @@ Some ''[[Partamona]]'' species exhibit a distinctive entrance architecture, where workers of ''[[Partamona helleri|P. helleri]]'' construct a large outer mud entrance leading to a smaller adjacent entrance. This unique design enables foragers to enter with high speed, bouncing off the ceiling of the outer entrance towards the smaller inner entrance. The peculiar appearance of this entrance has led to local names such as "toad mouth", highlighting the intriguing adaptations found in stingless bee nest entrances.<ref name=":15" /> -<gallery perrow="4" widths="160" caption="Different species nest entrances in Brazilian stingless bees"> +<gallery perrow="4" widths="160" caption="Different species nest entrances in Brazilian stingless bees"> File:Abelha limão.jpg|''[[Lestrimelitta limao]]'' File:Frieseomelitta varia nest entrance.jpg|''[[Frieseomelitta varia]]'' @@ -326,5 +326,5 @@ {{Main|Pollination}} [[File:Bee pollinating a rose.jpg|thumb|Solitary bee visiting and pollinating a flower.]] -Bees play a critical role in the ecosystem, particularly in the pollination of natural vegetation. This activity is essential for the reproduction of various plant species, particularly in [[tropical forest]]s where most tree species rely on [[Entomophily|insect pollination]]. Even in temperate climates, where [[Anemophily|wind pollination]] is prevalent among forest trees, many bushes and herbaceous plants, rely on bees for pollination. The significance of bees extends to arid regions, such as [[Deserts and xeric shrublands|desertic and xeric shrublands]], where bee-pollinated plants are essential for preventing [[erosion]], supporting [[wildlife]], and ensuring [[Ecological stability|ecosystem stability]].<ref name=":23">{{harvtxt|Michener|2000|pp=4-5}}</ref> +Bees play a critical role in the ecosystem, particularly in the pollination of natural vegetation. This activity is essential for the reproduction of various plant species, particularly in [[tropical forest]]s where most tree species rely on [[Entomophily|insect pollination]]. Even in temperate climates, where [[Anemophily|wind pollination]] is prevalent among forest trees, many bushes and herbaceous plants, rely on bees for pollination. The significance of bees extends to arid regions, such as [[Deserts and xeric shrublands|desertic and xeric shrublands]], where bee-pollinated plants are essential for preventing [[erosion]], supporting [[wildlife]], and ensuring [[Ecological stability|ecosystem stability]].<ref name=":23">{{harvtxt|Michener|2000|pp=4-5}}</ref> The impact of bee pollination on agriculture is substantial. In the late 1980s, certain plants were estimated to contribute between $4.6 to $18.9 billion to the U.S. economy, primarily through insect-pollinated crops. Although some bee-pollinated plants can self-pollinate in the absence of bees, the resulting crops often suffer from inbreeding depression. The quality and quantity of seeds or fruits are significantly enhanced when bees participate in the pollination process. Although estimates of crop pollination attributed to honey bees are uncertain, it is undeniable that bee pollination is a vital and economically valuable activity.<ref name=":23" /> @@ -463,8 +463,20 @@ |<ref name=":22" /> |} -== Relationship to humans == + +== Worldwide overview == + +=== Africa === +Stingless bees also play a vital ecological role across [[Sub-Saharan Africa]] and [[Madagascar]]. To understand these insects on the African continent, it's important to consider the prevailing socio-economic and cultural contexts. Despite their ecological significance, the diversity, conservation, and behavior of these bees remain underexplored, particularly compared to better-studied regions such as [[South America]] and [[Southeast Asia]]. Also, honeybees were extensively researched, in contrast to native meliponines.<ref name=":31">{{harvtxt|Kajobe|2008}}</ref><ref name=":32">{{harvtxt|Byarugaba|2004}}</ref> + +Africa is home to seven [[Biodiversity hotspot|biodiversity hotspots]], yet the recorded bee fauna is moderate relative to the continent's size. Madagascar stands out with exceptionally high levels of endemic species, though much of the bee diversity remains undocumented.<ref name=":30">{{harvtxt|Eardley|Gikungu|Schwarz|2009}}</ref>'''<ref name=":31" />''' Africa is home to aproximately 36 species of meliponines, including seven endemic to Madagascar. Most of these bees are found in equatorian regions (tropical forests and some savannahs).<ref>{{harvtxt|Grüter|2010|pp=47 & 49}}</ref> + +Factors such as habitat destruction, pesticide use, and invasive species pose significant threats to these pollinators. Furthermore, high rates of nest mortality, driven by predation and human activity, exacerbate conservation challenges. Research indicates that stingless bees in Africa face greater pressures than their counterparts in the American and Asian tropics, underlining the urgency for targeted conservation measures.<ref name=":30" />'''<ref name=":31" />''' + +Uganda's [[Bwindi Impenetrable National Park]] has shown the presence of at least five stingless bee species in, distributed across two genera: ''[[Meliponula]]'' and ''[[Hypotrigona]]''.<ref name=":31" />'''<ref name=":32" />''' In Madagascar, there is only one genus of stingless bees: ''[[Liotrigona]].''<ref name=":30" /> + +Meliponiculture, for example, is practised in Angola and Tanzania, and interest in managing stingless bees is growing in other African countries as well. <ref>{{harvtxt|Grüter|2010|p=27}}</ref> === Australia === -"''Coot-tha''", which derives from "''ku-ta''", is one of the Aboriginal names for "wild stingless bee honey".<ref>{{harvtxt |Vit|Pedro|Roubik|2018}}</ref> Of the 1,600 species of wild bees native to Australia, about 14 are meliponines.<ref>{{cite news | url=http://www.abc.net.au/science/articles/2003/05/08/2045526.htm | title=Stingless bee rescue |author= Wendy Pyper |date=May 8, 2003 |work=ABC Science}}</ref> These species bear a variety of names, including Australian native honey bees, native bees, sugar-bag bees, and sweat bees (because they land on people's skin to collect sweat).<ref>{{Cite web |title=Australian Native Stingless Bees |url=https://www.aussiebee.com.au/australian-stingless-bees.html |access-date=2024-05-28 |website=www.aussiebee.com.au}}</ref> All are small and usually black in colour, with hairy extended hind legs for carrying [[nectar]] and [[pollen]]; because of the latter, they are sometimes mistaken for [[bumblebee]]s. The various stingless species look quite similar, with the two most common species, ''[[Tetragonula carbonaria]]'' and ''[[Austroplebeia australis]]'', displaying the greatest variation, as the latter is smaller and less active. Both of these inhabit the area around [[Brisbane]].<ref>{{Cite web |last=Steve |date=2015-05-25 |title=Native Stingless Bees - Tetragonula carbonaria |url=https://www.nativebeehives.com/native-stingless-bees-tetragonula-carbonaria/ |access-date=2024-05-28 |website=www.nativebeehives.com |language=en-AU}}</ref> +Of the 1,600 species of wild bees native to Australia, about 14 are meliponines.<ref>{{cite news | url=http://www.abc.net.au/science/articles/2003/05/08/2045526.htm | title=Stingless bee rescue |author= Wendy Pyper |date=May 8, 2003 |work=ABC Science}}</ref> "''Coot-tha''", which derives from "''ku-ta''", is one of the Aboriginal names for "wild stingless bee honey".<ref>{{harvtxt|Vit|Pedro|Roubik|2018}}</ref> These species bear a variety of names, including Australian native honey bees, native bees, sugar-bag bees, and sweat bees (because they land on people's skin to collect sweat).<ref>{{Cite web |title=Australian Native Stingless Bees |url=https://www.aussiebee.com.au/australian-stingless-bees.html |access-date=2024-05-28 |website=www.aussiebee.com.au}}</ref> The various stingless species look quite similar, with the two most common species, ''[[Tetragonula carbonaria]]'' and ''[[Austroplebeia australis]]'', displaying the greatest variation, as the latter is smaller and less active. Both of these inhabit the area around [[Brisbane]].<ref>{{Cite web |last=Steve |date=2015-05-25 |title=Native Stingless Bees - Tetragonula carbonaria |url=https://www.nativebeehives.com/native-stingless-bees-tetragonula-carbonaria/ |access-date=2024-05-28 |website=www.nativebeehives.com |language=en-AU}}</ref> As stingless bees are usually harmless to humans, they have become an increasingly attractive addition to the suburban backyard. Most meliponine beekeepers do not keep the bees for [[honey]], but rather for the pleasure of conserving native species whose original [[habitat (ecology)|habitat]] is declining due to human development. In return, the bees [[pollinate]] crops, garden flowers, and bushland during their search for nectar and pollen. While a number of beekeepers fill a small niche market for bush honey, native meliponines only produce small amounts and the structure of their hives makes the honey difficult to extract. Only in warm areas of Australia such as [[Queensland]] and northern [[New South Wales]] are favorable for these bees to produce more honey than they need for their own survival. Most bees only come out of the hive when it is above about 18°C (64°F).<ref>{{Cite web|url=https://www.abc.net.au/news/2019-07-05/moves-underway-to-make-a-standard-for-native-bee-honey/11278346|title=Native bee honey set to have its own food standard|last=Thomas|first=Kerrin|date=2019-07-05|website=ABC News|language=en-AU|access-date=2019-07-10}}</ref> Harvesting honey from a nest in a cooler area could weaken or even kill the nest. @@ -478,55 +490,46 @@ [[File:Pollinating bee covered with pollen.jpg|thumb|''[[Trigona spinipes]],'' covered with [[pollen]], visiting a flower of the vegetable sponge gourd (''Luffa cylindrica'') in [[Campinas]], [[Brazil]].]] -Brazil is home to several species bees belonging to Meliponini, with more than 300 species already identified and probably more yet to be discovered and described. They vary greatly in shape, size, and habits, and 20 to 30 of these species have good potential as honey producers. Although they are still quite unknown by most people, an increasing number of beekeepers (''meliponicultores'', in Portuguese) have been dedicated to these bees throughout the country.<ref name=":27" /><ref>{{harvtxt|Villas-Bôas|2018|p=17}}</ref> This activity has experienced significant growth since August 2004, when national laws were changed to allow native bee colonies to be freely marketed, which was previously forbidden in an unsuccessful attempt to protect these species. Nowadays the capture or destruction of existing colonies in nature is still forbidden, and only new colonies formed by the bees themselves in artificial traps can be collected from the wild.<ref>[https://conama.mma.gov.br/?option=com_sisconama&task=arquivo.download&id=440 CONAMA 2004 Resolution]</ref> Most colonies marketed are artificially produced by authorized beekeepers, through division of already existing captive colonies. Besides honey production, Brazilian stingless bees such as the jataí (''[[Tetragonisca angustula]]''), mandaguari (''[[Scaptotrigona postica]]''), and mandaçaia (''[[Melipona quadrifasciata]]'') serve as major pollinators of tropical plants and are considered the ecological equivalent of the honey bee.<ref name=":21" /><ref name=":22" /> +Brazil is home to several species bees belonging to Meliponini, with more than 300 species already identified and probably more yet to be discovered and described. They vary greatly in shape, size, and habits, and 20 to 30 of these species have good potential as honey producers. Although they are still quite unknown by most people, an increasing number of beekeepers (''meliponicultores'', in Portuguese) have been dedicated to these bees throughout the country.<ref name=":27" /><ref>{{harvtxt|Villas-Bôas|2018|p=17}}</ref> This activity has experienced significant growth since August 2004, when national laws were changed to allow native bee colonies to be freely marketed, which was previously forbidden in an unsuccessful attempt to protect these species. Nowadays the capture or destruction of existing colonies in nature is still forbidden, and only new colonies formed by the bees themselves in artificial traps can be collected from the wild.<ref>[https://conama.mma.gov.br/?option=com_sisconama&task=arquivo.download&id=440 CONAMA 2004 Resolution]</ref> Most marketed colonies are artificially produced by authorized beekeepers, through division of already existing captive colonies. Besides honey production, Brazilian stingless bees such as the jataí (''[[Tetragonisca angustula]]''), mandaguari (''[[Scaptotrigona postica]]''), and mandaçaia (''[[Melipona quadrifasciata]]'') serve as major pollinators of tropical plants and are considered the ecological equivalent of the honey bee.<ref name=":21" /><ref name=":22" /> -Also, much practical and academic work is being done about the best ways of keeping such bees, multiplying their colonies, and exploring the honey they produce. Among many others, species such as jandaíra (''[[Melipona subnitida]]'') and true ''uruçu'' (''[[Melipona scutellaris]]'') in the northeast of the country, mandaçaia (''[[Melipona quadrifasciata]]'') and yellow uruçu (''Melipona rufiventris'') in the south-southeast, tiúba or jupará (''[[Melipona interrupta]]''<ref>{{Cite web|url=http://moure.cria.org.br/catalogue?id=34747|title=+Moure's Bee Catalogue}}</ref>) and straw-bee (''[[Scaptotrigona polysticta]]'') in the north and jataí (''[[Tetragonisca angustula]]'') throughout the country are increasingly kept by small, medium, and large producers. Many other species as the mandaguari (''[[Scaptotrigona postica]]''), the guaraipo (''[[Melipona bicolor]]''), marmelada (''[[Frieseomelitta varia]]'') and the iraí (''[[Nannotrigona testaceicornis]]''), to mention a few, are also reared. Through the cultivation of honey or selling of colonies, keeping stingless bees is an increasingly profitable activity. A single colony of species like mandaçaia and true "uruçu" can be divided up to four times a year, and each of the new colonies obtained this way can be sold for about US$100. +Also, much practical and academic work is being done about the best ways of keeping such bees, multiplying their colonies, and exploring the honey they produce.<ref>{{Cite web |date=2024-07-26 |title=Arquivos Artigos - A.B.E.L.H.A. |url=https://abelha.org.br/artigos/ |access-date=2024-12-31 |website=abelha.org.br |language=pt-BR}}</ref> Among many others, species such as jandaíra (''[[Melipona subnitida]]'') and true ''uruçu'' (''[[Melipona scutellaris]]'') in the northeast of the country, mandaçaia (''[[Melipona quadrifasciata]]'') and yellow uruçu (''Melipona rufiventris'') in the south-southeast, tiúba or jupará (''[[Melipona interrupta]]'') and canudo (''[[Scaptotrigona polysticta]]'') in the north and jataí (''[[Tetragonisca angustula]]'') throughout the country are increasingly kept by small, medium, and large producers. Many other species as the mandaguari (''[[Scaptotrigona postica]]''), the guaraipo (''[[Melipona bicolor]]''), marmelada (''[[Frieseomelitta varia]]'') and the iraí (''[[Nannotrigona testaceicornis]]''), to mention a few, are also reared.<ref name=":33">{{Cite web |date=2022-02-21 |title=Fichas catalográficas das espécies relevantes para a meliponicultura - A.B.E.L.H.A. |url=https://abelha.org.br/fichas-catalograficas-das-especies-relevantes-para-a-meliponicultura-2/ |access-date=2024-12-31 |website=abelha.org.br |language=pt-BR}}</ref> According to [[Chico Mendes Institute for Biodiversity Conservation|ICMBio]] and the [[Ministry of Environment and Climate Change (Brazil)|Ministry of the Environment]] there are presently four species of Meliponini listed in the [[IUCN Red List|National Red List of Threatened Species]] in Brazil. ''[[Melipona capixaba]]'', ''[[Melipona rufiventris]]'', ''[[Melipona scutellaris]]'', and ''[[Partamona littoralis]]'' all listed as Endangered (EN).<ref>{{Cite web |title=IBAMA |url=https://www.ibama.gov.br/component/legislacao/?view=legislacao&force=1&legislacao=134521#:~:text=Ementa:,Extin%C3%A7%C3%A3o%20-%20Peixes%20e%20Invertebrados%20Aqu%C3%A1ticos. |access-date=2024-02-26 |website=www.ibama.gov.br}}</ref><ref>[[Instituto Chico Mendes de Conservação da Biodiversidade|ICMBio]]. 2018. [https://www.gov.br/icmbio/pt-br/centrais-de-conteudo/publicacoes/publicacoes-diversas/livro_vermelho_2018_vol7.pdf Livro Vermelho da Fauna Brasileira Ameaçada de Extinção: Volume VII - Invertebrados]. ''In'': ICMBio. (Org.). [https://www.gov.br/icmbio/pt-br/centrais-de-conteudo/publicacoes/publicacoes-diversas/livro_vermelho_2018_vol1.pdf Livro Vermelho da Fauna Brasileira Ameaçada de Extinção]. Brasília: ICMBio. 727p.</ref> ==== Honey production ==== -Although the colony population of most of these bees is much smaller than that of European bees, the productivity per bee can be quite high. Probably the world champions in honey productivity, the manduri (''[[Melipona marginata]]'') and the ''guaraipo'' (''M. bicolor''), live in swarms of only around 300 individuals, but can still produce up to 5 liters (.79 US gallon) of honey a year under the right conditions. The manduri is one of the smallest bees in the genus ''[[Melipona]]'', varying in length from 6 to 7 mm (15/64" to 9/32").<ref>{{harvtxt|Witter|Blochtein|Santos|2007|p=27}}</ref> Species of the genus ''Scaptotrigona'' have very large colonies, with up to 20,000 individuals, and can produce from 3 to 8 liters (2 US gallons) of honey a year. - -Some large breeders have more than 3,000 hives of the tamer but still highly productive species in the genus ''Melipona'', such as the tiúba, the true ''uruçu'', and the jandaíra, each with 3,000 or more bees per colony. They can produce over 1.5 tons (3,000&nbsp;lbs) of honey every year.{{Citation needed|date=June 2024}} In large bee farms, only the availability of flowers limits the honey production per colony. Their honey is considered more palatable because it's not overly sweet, and it also is thought to have medicinal properties more pronounced than honey from bees of the genus ''Apis'' due to the higher level of antimicrobial substances. As a result, the honey from stingless bees returns very high profits, with prices greater than those for the more common honey produced by European bees. However, much larger numbers of beehives are required to produce amounts of honey comparable to that of European bees. Also, due to the fact of those bees storing honey in cerumen pots instead of standardized honeycombs as in the honeybee rearing makes extraction a lot more difficult and laborious.<ref>{{harvtxt|Fonseca|Sodré|Carvalho|Alves|Souza|Silva|Oliveira|Machado|Clarton|2007}}</ref> - -The honey from stingless bees has a lighter color and a higher water content, from 25% to 35%, compared to the honey from the genus ''Apis''. This contributes to its less cloying taste but also causes it to spoil more easily. Thus, for marketing, this honey needs to be processed through desiccation, fermentation or pasteurization. In its natural state, it should be kept under refrigeration.<ref>{{harvtxt|Grüter|2020|pp=12-14}}</ref> +Although the colony population of most of these bees is much smaller than that of European bees, the productivity per bee can be quite high. Interestingly, honey production is more connected to the body size, not the colony size. The manduri (''[[Melipona marginata]]''), jandaíra (''[[Melipona subnitida]]'') and the ''guaraipo'' (''[[Melipona bicolor|M. bicolor]]'') live in swarms of only around 300 individuals but can still produce up to 5 liters (1.3 US gallon) of honey a year under the right conditions.<ref>{{harvtxt|Grüter|2010|p=14}}</ref> In large bee farms, only the availability of flowers limits the honey production per colony. However, much larger numbers of beehives are required to produce amounts of honey comparable to that of European bees. Also, due to the fact of those bees storing honey in cerumen pots instead of standardized honeycombs as in the honeybee rearing makes extraction a lot more difficult and laborious.<ref>{{harvtxt|Fonseca|Sodré|Carvalho|Alves|Souza|Silva|Oliveira|Machado|Clarton|2007}}</ref> +The honey from stingless bees has a higher water content, from 25% to 35%, compared to the honey from the genus ''Apis''. This contributes to its less cloying taste but also causes it to spoil more easily. Thus, for marketing, this honey needs to be processed through desiccation, fermentation or pasteurization. In its natural state, it should be kept under refrigeration.<ref>{{harvtxt|Grüter|2020|pp=12-14}}</ref> ==== Bees as pets ==== [[Image:Mandaçaias.jpg|left|thumb|A swarm of mandaçaias bees around an artificial [[beehive]] installed in a house's backyard in [[Brazil]]]] -Due to the lack of a functional stinger and characteristic nonaggressive behavior of many Brazilian species of stingless bees, they can be reared without problems in densely populated environments such as cities, provided enough flowers are at their disposal nearby. Some breeders (''meliponicultores'') can produce honey even in apartments up to the 12th floor.<ref>{{cite web|url=http://meliponariocapixaba.blogspot.com.br/2009/11/e-possivel-criar-abelhas-em.html|title=Meliponario Capixaba: É POSSÍVEL CRIAR ABELHAS EM APARTAMENTOS?|author=João Luiz|access-date=12 March 2016}}</ref> +Due to the lack of a functional stinger and characteristic nonaggressive behavior of many Brazilian species of stingless bees, they can be reared without problems in densely populated environments (residential buildings, schools, urban parks), provided enough flowers are at their disposal nearby. Some breeders (''meliponicultores'') can produce honey even in apartments up to the 12th floor.<ref>{{cite web|url=http://meliponariocapixaba.blogspot.com.br/2009/11/e-possivel-criar-abelhas-em.html|title=Meliponario Capixaba: É POSSÍVEL CRIAR ABELHAS EM APARTAMENTOS?|author=João Luiz|access-date=12 March 2016}}</ref> -The [[Melipona quadrifasciata|mandaçaias]] (''Melipona quadrifasciata'') are extremely tame, rarely attacking humans (only when their hives are opened for honey extraction or colony division). They form small, manageable colonies of only 400&ndash;600&nbsp;individuals. At the same time, a single rational beehive of ''mandaçaia'' can produce up to 4&nbsp;liters (1&nbsp;US gallon) of honey a year, making the species very attractive for home keepers. They are fairly large bees, up to 11&nbsp;mm (7/16") in length, and as a result have better body heat control, allowing them to live in regions where temperatures can drop a little lower than 0&nbsp;°C (32&nbsp;°F). However, they are somewhat selective about which flowers they will visit, preferring the flora that occurs in their natural environment. They are thus difficult to keep outside their region of origin (the northeast coast of Brazil).<ref>{{harvtxt|Brito|Faquinello|Paula-Leite|Carvalho|1997}}</ref> +The [[Melipona quadrifasciata|mandaçaias]] (''Melipona quadrifasciata'') are extremely tame, rarely attacking humans (only when their hives are opened for honey extraction or colony division). They form small, manageable colonies of only 400&ndash;600&nbsp;individuals. They are fairly large bees, up to 11&nbsp;mm (7/16") in length, and as a result have better body heat control, allowing them to live in regions where temperatures can drop a little lower than 0&nbsp;°C (32&nbsp;°F). However, they are somewhat selective about which flowers they will visit, preferring the flora that occurs in their natural environment. They are thus difficult to keep outside their region of origin (the eastern coast of Brazil). Once very common, the mandaçaia is now rather rare in nature, mainly due to the destruction of their native forests in the of Brazil.<ref>{{harvtxt|Brito|Faquinello|Paula-Leite|Carvalho|1997}}</ref><ref name=":33" /> -Other species, like the ''tiúba'' and the true ''uruçu'', are also very tame and highly productive. Their colonies have from 3,000&ndash;5,000 individuals (for comparison, the population of honeybee swarms can peak at 80,000&nbsp;individuals) and can produce up to 10 liters (2.6 US gallons) of honey a year.{{Citation needed|date=June 2024}} They can be easily kept at home but will survive only in regions with a warm climate, their larvae dying at temperatures lower than 12&nbsp;°C (54&nbsp;°F). The yellow ''uruçu'', however, can survive at temperatures lower than 0&nbsp;°C (32&nbsp;°F), and their colonies, bearing about 3,500&nbsp;individuals, can produce up to 6&nbsp;liters (1.5&nbsp;US gallons) of honey a year.{{Citation needed|date=June 2024}} But this species will react with powerful nipping if its nests are disturbed, and usually they are only kept by professional meliponicultors. +Other groups of Brazilian stingless bees, genera ''[[Plebeia]]'' and ''[[Leurotrigona]]'', are also very tame and much smaller, with one of them (''[[Plebeia minima]]'') reaching no more than 2.5&nbsp;mm (3/32") in length, and the ''[[lambe-olhos]]'' ("lick-eyes" bee, ''[[Leurotrigona muelleri]]'') being even smaller, at no more than 1.5&nbsp;mm (3/32"). Many of these species are known as mirim (meaning 'small' in the [[Tupi-Guarani languages]]). As a result, they can be kept in very small artificial hives, thus being of interest for keepers who want them as pollinators in small glasshouses or just for the pleasure of having a 'toy' bee colony at home.<ref name=":33" /><ref>{{Cite web |title=Lambe-olhos – Leurotrigona muelleri – Laboratório de Sistemática de Plantas |url=https://sites.usp.br/jardimdabotanicausprp/lambe-olhos-leurotrigona-muelleri/ |access-date=2024-12-31 |website=sites.usp.br}}</ref><ref>{{Cite web |title=Mirim – Plebeia droryana – Laboratório de Sistemática de Plantas |url=https://sites.usp.br/jardimdabotanicausprp/mirim-plebeia-droryana/ |access-date=2024-12-31 |website=sites.usp.br}}</ref> Being so tiny, these species produce only a very small amount of honey, typically less than 500&nbsp;ml (1/2 US pint) a year, so are not interesting for commercial honey production. -Another suitable species for keeping at home is the ''guaraipo'' (''[[Melipona bicolor]]''). It is also quite tame, never attacking the beekeeper, and their colonies have fewer than 600 individuals. They can withstand temperatures as low as &minus;10&nbsp;°C (14&nbsp;°F), and each colony can produce over 3&nbsp;liters ({{frac|3|4}}&nbsp;US gallons) of honey a year.{{Citation needed|date=June 2024}} Their colonies usually have more than one single queen at a time (usually two or three, but sometimes up to five), a phenomenon called [[Gyne|polygyne]], and thus are less sensitive to the death of one queen, which can cause the loss of a whole colony in other species. But the ''guaraipo'' is very sensitive to low levels of humidity, and their hives must be equipped with means to keep a high moisture content. Once very common, the ''guaraipo'' is now rather rare in nature, mainly due to the destruction of their native forests in the south-southeast of Brazil. +[[Image:Abelha-jatai.jpg|right|thumb|Entrance pipe of ''jataís´s'' colony: The ''jataí'' bees build a wax pipe in the entrance of their nest.]] +Belonging to the same group, the jataí (''[[Tetragonisca angustula]]''), the marmelada (''[[Frieseomelitta varia]]''), and the moça-branca (''[[Frieseomelitta doederleini]]'') are intermediate in size between those very small species and the European bee. They are very adaptable species; the jataí, and can be reared in many different regions and environments, being quite common in most Brazilian cities. The jataí can bite when disturbed, but its jaws are weak, and in practice they are harmless, while the marmelada and moça-branca usually deposit propolis on their aggressors. Jataí is one of the first species to be kept by home beekeepers. Their nests can be easily identified in trees or wall cavities by the yellow wax pipe they build at the entrance, usually guarded by some soldier bees, which are stronger than regular worker bees. The marmelada and moça-branca make a little less honey, but it is denser and sweeter than most from other stingless bees and is considered very tasty.<ref name=":33" /><ref>{{harvtxt|Castanheira|Contel|2005|}}</ref> -Other groups of Brazilian stingless bees, genera ''[[Plebeia]]'' and ''[[Leurotrigona]]'', are also very tame and much smaller, with one of them (''[[Plebeia minima]]'') reaching no more than 2.5&nbsp;mm (3/32") in length, and the ''[[lambe-olhos]]'' ("lick-eyes" bee, ''[[Leurotrigona muelleri]]'') being even smaller, at no more than 1.5&nbsp;mm (3/32"). Many of these species are known as ''mirim'' (meaning 'small' in the [[Tupi-Guarani languages]]). As a result, they can be kept in very small artificial hives, thus being of interest for keepers who want them as pollinators in small glasshouses or just for the pleasure of having a 'toy' bee colony at home. Being so tiny, these species produce only a very small amount of honey, typically less than 500&nbsp;ml (1/2 US pint) a year, so are not interesting for commercial honey production. - -[[Image:Abelha-jatai.jpg|right|thumb|Entrance pipe of ''jataís´s'' colony: The ''jataí'' bees build a wax pipe in the entrance of their nest.]] -Belonging to the same group, the ''jataí'' (''[[Tetragonisca angustula]]''), the ''marmelada ''(''[[Frieseomelitta varia]]''), and the ''moça-branca'' (''[[Frieseomelitta doederleini]]'') are intermediate in size between those very small species and the European bee. They are very adaptable species; the ''jataí'', and can be reared in many different regions and environments, being quite common in most Brazilian cities. The ''jataí'' can bite when disturbed, but its jaws are weak, and in practice they are harmless, while the ''marmelada'' and ''moça-branca'' usually deposit propolis on their aggressors. Producing up to 1.5&nbsp;liters (0.4&nbsp;US gallons) of honey a year, their honey is considered among the best from stingless bees. In fact, the ''jataí'' was one of the first species to be kept by home beekeepers. Their nests can be easily identified in trees or wall cavities by the yellow wax pipe they build at the entrance, usually guarded by some soldier bees, which are stronger than regular worker bees. The'' marmelada'' and ''moça-branca'' make a little less honey, but it is denser and sweeter than most from other stingless bees and is considered very tasty. - -=== Maya stingless bees of Central America === +=== Central America === [[Image:Colmena melipona.jpg|right|thumb|A Maya stingless bee hive: A piece of hollow log provides a home for meliponine bees in [[Belize]].]] The stingless bees ''[[Melipona beecheii]]'' and ''[[Melipona yucatanica|M. yucatanica]]'' are the primary native bees cultured in Central America, though a few other species are reported as being occasionally managed (e.g., ''[[Trigona fulviventris]]'' and ''[[Scaptotrigona mexicana]]'').<ref>{{harvtxt|Kent|1984}}</ref> They were extensively cultured by the [[Maya civilization]] for honey, and regarded as sacred. They continue to be cultivated by the modern [[Maya peoples]], although these bees are endangered due to massive deforestation, altered [[agriculture|agricultural]] practices (especially overuse of [[insecticide]]s), and changing beekeeping practices with the arrival of the [[Africanized honey bee]], which produces much greater honey crops.<ref name=":26">{{harvtxt|Villanueva|Roubik|Colli-Ucán|1998}}</ref> ==== History ==== -Native meliponines (''M. beecheii'' being the favorite) have been kept by the lowland Maya for thousands of years. The [[Yucatec Maya language]] name for this bee is ''xunan kab'', meaning "(royal, noble) lady bee".<ref>{{cite web|title=Diccionario Introductorio|url=http://www.uqroo.mx/libros/maya/diccionario.pdf|work=uqroo.mx|publisher=Universidad De Quintana Roo|access-date=27 March 2014|language=es}}</ref> The bees were once the subject of religious ceremonies and were a symbol of the bee-god [[Ah-Muzen-Cab]], known from the ''[[Madrid Codex (Maya)|Madrid Codex]]''. +Native meliponines (''M. beecheii'' being the most common) have been kept by the lowland Maya for thousands of years. The [[Yucatec Maya language]] name for this bee is ''xunan kab'', meaning "(royal, noble) lady bee".<ref>{{harvtxt|Grüter|2010|p=54}}</ref><ref>{{cite web|title=Diccionario Introductorio|url=http://www.uqroo.mx/libros/maya/diccionario.pdf|work=uqroo.mx|publisher=Universidad De Quintana Roo|access-date=27 March 2014|language=es}}</ref> The bees were once the subject of religious ceremonies and were a symbol of the bee-god [[Ah-Muzen-Cab]], known from the ''[[Madrid Codex (Maya)|Madrid Codex]]''.<ref name=":34">{{Cite web |date=2016-07-17 |title=People and bees. Mayan bees in the Madrid Codex |url=https://ecomercioagrario.com/en/people-and-bees-mayan-bees-in-the-madrid-codex/ |access-date=2024-12-31 |website=eComercio Agrario |language=en-US}}</ref> -The bees were, and still are, treated as pets. Families would have one or many log-hives hanging in and around their houses. Although they are stingless, the bees do bite and can leave welts similar to a mosquito bite. The traditional way to gather bees, still favored among the locals, is find a wild hive, then the branch is cut around the hive to create a portable log, enclosing the colony. This log is then capped on both ends with another piece of wood or pottery and sealed with mud. This clever method keeps the melipine bees from mixing their brood, pollen, and honey in the same comb as do the European bees. The brood is kept in the middle of the hive, and the honey is stored in vertical "pots" on the outer edges of the hive. A temporary, replaceable cap at the end of the log allows for easy access to the honey while doing minimal damage to the hive. However, inexperienced handlers can still do irreversible damage to a hive, causing the hive to swarm and abscond from the log. With proper maintenance, though, hives have been recorded as lasting over 80 years, being passed down through generations. In the archaeological record of Mesoamerica, stone discs have been found that are generally considered to be the caps of long-disintegrated logs that once housed the beehives. +The bees were, and still are, treated as pets. Families would have one or many log-hives hanging in and around their houses. Although they are stingless, the bees do bite and can leave welts similar to a mosquito bite. The traditional way to gather bees, still favored among the locals, is find a wild hive, then the branch is cut around the hive to create a portable log, enclosing the colony. With proper maintenance, hives have been recorded as lasting over 80 years, being passed down through generations. In the archaeological record of Mesoamerica, stone discs have been found that are generally considered to be the caps of long-disintegrated logs that once housed the beehives.<ref name=":34" /><ref name=":35">{{Cite web |last=Kennedy |first=Jennifer |last2=Arghiris |first2=Richard |date=2019-01-31 |title=House of the Royal Lady Bee: Maya revive native bees and ancient beekeeping |url=https://news.mongabay.com/2019/01/house-of-the-royal-lady-bee-maya-revive-native-bees-and-ancient-beekeeping/ |access-date=2024-12-31 |website=Mongabay Environmental News |language=en-US}}</ref> ==== Tulum ==== -[[Tulum]], the site of a [[pre-Columbian]] Maya city on the Caribbean coast 130&nbsp;km (81&nbsp;mi) south of Cancun, has a god depicted repeatedly all over the site. Upside down, he appears as a small figure over many doorways and entrances. One of the temples, the "Templo del Dios Descendente" or the Temple of the Descending God, stands just left of the central plaza. Speculation is that he may be the "Bee God", Ah Muzen Cab, as seen in the Madrid Codex. It is possible that this was a religious/trade center with emphasis on ''xunan kab'', the "royal lady". +[[Tulum]], the site of a [[pre-Columbian]] Maya city on the Caribbean coast 130&nbsp;km (81&nbsp;mi) south of Cancun, has a god depicted repeatedly all over the site. Upside down, he appears as a small figure over many doorways and entrances. One of the temples, the Temple of the Descending God (''Templo del Dios Descendente''), stands just left of the central plaza. Speculation is that he may be the "Bee God", Ah Muzen Cab, as seen in the ''Madrid Codex''. It is possible that this was a religious/trade center with emphasis on ''xunan kab'', the "royal lady".<ref name=":35" /> ==== Economic uses ==== -[[Balché]], a traditional [[Mesoamerica]]n alcoholic beverage similar to [[mead]], was made from fermented honey and the bark of the leguminous'' balché'' tree (''[[Lonchocarpus violaceus]]''), hence its name.<ref>{{harvtxt|Ott|1998}}</ref> It was traditionally brewed in a canoe. The drink was known to have [[entheogen]]ic properties, that is, to produce mystical experiences, and was consumed in medicinal and ritual practices. Beekeepers would place the nests near the psychoactive plant ''[[Turbina corymbosa]]'' and possibly near ''balché'' trees, forcing the bees to use nectar from these plants to make their honey.<ref>{{harvtxt|Ott|1998}}</ref> Additionally, brewers would add extracts of the bark of the ''balché'' tree to the honey mixture before fermentation. The resulting beverage is responsible for psychotropic effects when consumed, due to the ergoline compounds in the pollen of the ''T. corymbosa'', the ''Melipona'' nectar gathered from the ''balché'' flowers, or the hallucinogenic compounds of the ''balché'' tree bark. +[[Balché]], a traditional [[Mesoamerica]]n alcoholic beverage similar to [[mead]], was made from fermented honey and the bark of the leguminous'' balché'' tree (''[[Lonchocarpus violaceus]]''), hence its name. It was traditionally brewed in a canoe. The drink was known to have [[entheogen]]ic properties, that is, to produce mystical experiences, and was consumed in medicinal and ritual practices. Beekeepers would place the nests near the psychoactive plant ''[[Turbina corymbosa]]'' and possibly near ''balché'' trees, forcing the bees to use nectar from these plants to make their honey. Additionally, brewers would add extracts of the bark of the ''balché'' tree to the honey mixture before fermentation. The resulting beverage is responsible for psychotropic effects when consumed, due to the ergoline compounds in the pollen of the ''T. corymbosa'', the ''Melipona'' nectar gathered from the ''balché'' flowers, or the hallucinogenic compounds of the ''balché'' tree bark.<ref>{{harvtxt|Ott|1998}}</ref> -[[Lost-wax casting]], a common metalworking method typically found where the inhabitants keep bees, was also used by the Maya. The wax from ''Melipona'' is soft and easy to work, especially in the humid Maya lowland. This allowed the Maya to create smaller works of art, jewelry, and other metalsmithing that would be difficult to forge. It also makes use of the leftovers from honey extraction. If the hive was damaged beyond repair, the whole of the comb could be used, thus using all of the hive. With experienced keepers, though, only the honey pot could be removed, the honey extracted, and the wax used for casting or other purposes. +[[Lost-wax casting]], a common metalworking method typically found where the inhabitants keep bees, was also used by the Maya. The wax from ''Melipona'' is soft and easy to work, especially in the humid Maya lowland. This allowed the Maya to create smaller works of art, jewelry, and other metalsmithing that would be difficult to forge. It also makes use of the leftovers from honey extraction. If the hive was damaged beyond repair, the whole of the comb could be used, thus using all of the hive. With experienced keepers, though, only the honey pot could be removed, the honey extracted, and the wax used for casting or other purposes.<ref>{{harvtxt|Pitses|2018|}}</ref><ref>{{harvtxt|Schwarz|1945|}}</ref> ==== Future ==== -The outlook for meliponines in Mesoamerica is uncertain. The number of active Meliponini beekeepers is shy in comparison with the Africanized ''[[Apis mellifera]]'' breeders. The high honey yield, 100&nbsp;kg (220&nbsp;lbs) or more annually, along with the ease of hive care and ability to create new hives from existing stock, commonly outweighs the negative consequences of "killer bee" hive maintenance. - -Further complicating the issue, [[Africanized honey bee]]s do not visit some flora, such as those in the [[Solanaceae|tomato family]], and several forest trees and shrubs, which rely on the native stingless bees for pollination. A decline in populations of native flora has already occurred in areas where stingless bees have been displaced by Africanized honey bees. +The outlook for meliponines in Mesoamerica is uncertain. The number of active Meliponini beekeepers is shy in comparison with the Africanized ''[[Apis mellifera]]'' breeders. The high honey yield, 100&nbsp;kg (220&nbsp;lbs) or more annually, along with the ease of hive care and ability to create new hives from existing stock, commonly outweighs the negative consequences of "killer bee" hive maintenance.<ref name=":36">{{harvtxt|Villanueva-G|Roubik|Colli-Ucán|2005|}}</ref> -An additional blow to the art of meliponine beekeeping is that many of the meliponine beekeepers are now elderly, and their hives may not be cared for once they die. The hives are considered similar to an old family collection, to be parted out once the collector dies or to be buried in whole or part along with the beekeeper upon death. In fact, a survey of a once-popular area of the Maya lowlands shows the rapid decline of beekeepers, down to around 70 in 2004 from thousands in the late 1980s. It is traditional in the Maya lowlands that the hive itself or parts of the hive be buried along with the beekeeper to ''volar al cielo'', "to fly to heaven".{{citation needed|date=March 2007}} Conservation efforts are underway in several parts of Mesoamerica.<ref>A comprehensive conservation guide can be found in the June 2005 issue of ''Bee World''.</ref> +An additional blow to the art of meliponine beekeeping is that many of the ''meliponicultores'' are now elderly, and their hives may not be cared for once they die. The hives are considered similar to an old family collection, to be parted out once the collector dies or to be buried in whole or part along with the beekeeper upon death. In fact, a survey of a once-popular area of the Maya lowlands shows the rapid decline of beekeepers, down to around 70 in 2004 from thousands in the late 1980s. Conservation efforts are underway in several parts of Mesoamerica.<ref name=":36" /><ref>A comprehensive conservation guide can be found in the June 2005 issue of ''Bee World''.</ref> == References ==<!-- ActaAmazonica34 p333 --> @@ -540,4 +543,5 @@ * {{Cite journal |last1=Antonini |first1=Yasmine |last2=Martins |first2=Rogério P. |date=2003 |title=The value of a tree species (''Caryocar brasiliense'') for a stingless bee ''Melipona quadrifasciata quadrifasciata'' |url=http://link.springer.com/10.1023/A:1027378306119 |journal=Journal of Insect Conservation |volume=7 |issue=3 |pages=167–174 |doi=10.1023/A:1027378306119|bibcode=2003JICon...7..167A |s2cid=6080884 }} * {{Cite journal |last1=Azmi |first1=Wahizatul Afzan |last2=Wan Sembok |first2=W Z |last3=Yusuf |first3=N |last4=Mohd. 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|last9=Nogueira-Neto |first9=P. |display-authors=3 |year=2006 |title=Global meliponiculture: challenges and opportunities |journal=Apidologie |volume=37 |issue=2 |pages=275–292 |doi=10.1051/apido:2006027 |doi-access=free}} +* {{Cite journal |last=Eardley |first=Connal D. |last2=Gikungu |first2=Mary |last3=Schwarz |first3=Michael P. |date=2009 |title=Bee conservation in Sub-Saharan Africa and Madagascar: diversity, status and threats |url=http://link.springer.com/10.1051/apido/2009016 |journal=Apidologie |volume=40 |issue=3 |pages=355–366 |doi=10.1051/apido/2009016 |issn=0044-8435}} * {{Cite journal |last=Engels |first=Wolf |date=1987 |title=Pheromones and reproduction in brazilian stingless bees |journal=Memórias do Instituto Oswaldo Cruz |language=en |volume=82 |pages=35–45 |doi=10.1590/S0074-02761987000700009 |issn=0074-0276|doi-access=free }} * {{Cite journal |last1=Ferreira |first1=E.L. |last2=Lencioni |first2=C. |last3=Benassi |first3=M.T. |last4=Barth |first4=M.O. |last5=Bastos |first5=D.H.M. |date=2009-07-30 |title=Descriptive Sensory Analysis and Acceptance of Stingless Bee Honey |url=http://journals.sagepub.com/doi/10.1177/1082013209341136 |journal=Food Science and Technology International |language=en |volume=15 |issue=3 |pages=251–258 |doi=10.1177/1082013209341136 |issn=1082-0132 |s2cid=84700846}} @@ -559,6 +565,7 @@ * {{Cite journal |last1=Imperatriz-Fonseca |first1=V. L. |last2=Zucchi |first2=R. |date=1995 |title=Virgin queens in stingless bee (Apidae, Meliponinae) colonies: a review |url=http://dx.doi.org/10.1051/apido:19950305 |journal=Apidologie |language=en |volume=26 |issue=3 |pages=231–244 |doi=10.1051/apido:19950305 |issn=0044-8435}} * {{Cite journal |last=Kajobe |first=Robert |date=2006-12-12 |title=Nesting biology of equatorial Afrotropical stingless bees (Apidae; Meliponini) in Bwindi Impenetrable National Park, Uganda |url=https://www.tandfonline.com/doi/full/10.1080/00218839.2007.11101403 |journal=Journal of Apicultural Research |language=en |volume=46 |issue=4 |pages=245–255 |doi=10.1080/00218839.2007.11101403 |issn=0021-8839 |s2cid=84923320}} -* <{{Cite journal |last=Kent |first=Robert B. |date=1984 |title=Mesoamerican Stingless Beekeeping |url=http://www.tandfonline.com/doi/abs/10.1080/08873638409478571 |journal=Journal of Cultural Geography |language=en |volume=4 |issue=2 |pages=14–28 |doi=10.1080/08873638409478571 |issn=0887-3631}} +* {{Cite journal |last=Kent |first=Robert B. |date=1984 |title=Mesoamerican Stingless Beekeeping |url=http://www.tandfonline.com/doi/abs/10.1080/08873638409478571 |journal=Journal of Cultural Geography |language=en |volume=4 |issue=2 |pages=14–28 |doi=10.1080/08873638409478571 |issn=0887-3631}} * {{cite journal |last1=Kerr |first1=W E. |year=1950 |title=Genetic determination of castes in the genus ''Melipona'' |journal=Genetics |volume=35 |issue=2 |pages=143–152 |doi=10.1093/genetics/35.2.143 |pmc=1209477 |pmid=17247339}} +* {{Cite journal |last=Kerr |first=Warwick Estevam |last2=Petrere Jr. |first2=Miguel |last3=Diniz Filho |first3=José Alexandre Felizola |date=2001-03 |title=Informações biológicas e estimativa do tamanho ideal da colmeia para a abelha tiúba do Maranhão (Melipona compressipes fasciculata Smith - Hymenoptera, Apidae) |url=https://www.scielo.br/j/rbzool/a/r74hTMg94tR9PTJGr8cvJwH/?lang=pt |journal=Revista Brasileira de Zoologia |language=pt |volume=18 |pages=45–52 |doi=10.1590/S0101-81752001000100003 |issn=0101-8175}} * {{cite journal |last1=Koch |first1=H. |year=2010 |title=Combining morphology and DNA barcoding resolves the taxonomy of Western Malagasy ''Liotrigona'' Moure", 1961 |url=http://www.africaninvertebrates.org.za/Koch_2010_51_2_474.aspx][http://www.tb1.ethz.ch/PublicationsEO/PDFpapers/Koch_AFRICAN_INVERTEBRATES_2010_51_413-421.pdf |url-status=dead |journal=African Invertebrates |volume=51 |issue=2 |pages=413–421 |doi=10.5733/afin.051.0210 |s2cid=49266406 |archive-url=https://web.archive.org/web/20110912162337/http://www.africaninvertebrates.org.za/Koch_2010_51_2_474.aspx |archive-date=2011-09-12}} * {{Cite journal |last=Martins |first=Aline C. |last2=Melo |first2=Gabriel A. R. |last3=Renner |first3=Susanne S. |date=2014-11-01 |title=The corbiculate bees arose from New World oil-collecting bees: Implications for the origin of pollen baskets |url=https://linkinghub.elsevier.com/retrieve/pii/S1055790314002450 |journal=Molecular Phylogenetics and Evolution |volume=80 |pages=88–94 |doi=10.1016/j.ympev.2014.07.003 |issn=1055-7903}} @@ -582,4 +589,5 @@ * {{cite journal |last1=Roubik |first1=D W. |year=2006 |title=Stingless bee nesting biology |journal=Apidologie |volume=37 |issue=2 |pages=124–143 |doi=10.1051/apido:2006026 |doi-access=free}} * {{Cite web |last=Sarchet |first=Penny |date=14 November 2014 |title=Zoologger: Stingless suicidal bees bite until they die |url=https://www.newscientist.com/article/dn26562-zoologger-stingless-suicidal-bees-bite-until-they-die/ |access-date=2020-01-24 |website=New Scientist |language=en-US}} +* {{Cite journal |last=Schwarz |first=Herbert F. |date=1945 |title=The Wax of Stingless Bees (Meliponidæ) and the Uses to Which It Has Been Put |url=https://www.jstor.org/stable/25005104?utm_source=chatgpt.com |journal=Journal of the New York Entomological Society |volume=53 |issue=2 |pages=137–144 |issn=0028-7199}} * {{Cite journal |last1=Slaa |first1=Ester Judith |last2=Chaves |first2=Luis Alejandro Sánchez |last3=Malagodi-Braga |first3=Katia Sampaio |last4=Hofstede |first4=Frouke Elisabeth |date=2006-03-01 |title=Stingless bees in applied pollination: practice and perspectives |url=http://dx.doi.org/10.1051/apido:2006022 |journal=Apidologie |language=en |volume=37 |issue=2 |pages=293–315 |doi=10.1051/apido:2006022 |s2cid=55280074 |issn=0044-8435}} * {{Cite journal |last1=Sousa |first1=Janaína Maria Batista de |last2=Souza |first2=Evandro Leite de |last3=Marques |first3=Gilmardes |last4=Benassi |first4=Marta de Toledo |last5=Gullón |first5=Beatriz |last6=Pintado |first6=Maria Manuela |last7=Magnani |first7=Marciane |date=2016-01-01 |title=Sugar profile, physicochemical and sensory aspects of monofloral honeys produced by different stingless bee species in Brazilian semi-arid region |journal=LWT - Food Science and Technology |volume=65 |pages=645–651 |doi=10.1016/j.lwt.2015.08.058 |issn=0023-6438 |doi-access=free}} @@ -588,4 +596,5 @@ * {{Cite journal |last1=Viana |first1=Blandina Felipe |last2=Coutinho |first2=Jeferson Gabriel da Encarnação |last3=Garibaldi |first3=Lucas Alejandro |last4=Castagnino |first4=Guido Laercio Bragança |last5=Gramacho |first5=Kátia Peres |last6=Silva |first6=Fabiana Oliveira |date=2014-10-09 |title=Stingless bees further improve apple pollination and production |url=https://pollinationecology.org/index.php/jpe/article/view/309 |journal=Journal of Pollination Ecology |language=en |volume=14 |pages=261–269 |doi=10.26786/1920-7603(2014)26 |issn=1920-7603|doi-access=free }} * {{cite journal |author=Villanueva, Rogel |display-authors=et al. |year=2005 |title=Extinction of ''Melipona beecheii'' and traditional beekeeping in the Yucatán peninsula |journal=Bee World |volume=86 |issue=2 |pages=35–41 |doi=10.1080/0005772X.2005.11099651 |s2cid=31943555}} +* {{Cite journal |last=Villanueva-G |first=Rogel |last2=Roubik |first2=David W |last3=Colli-Ucán |first3=Wilberto |date=2005-01 |title=Extinction of Melipona beecheii and traditional beekeeping in the Yucatán peninsula |url=http://www.tandfonline.com/doi/full/10.1080/0005772X.2005.11099651 |journal=Bee World |language=en |volume=86 |issue=2 |pages=35–41 |doi=10.1080/0005772X.2005.11099651 |issn=0005-772X}} * {{Cite journal |last1=Wenseleers |first1=T. |last2=Ratnieks |first2=F. L. W. |last3=Ribeiro |first3=M. D. F. |last4=Alves |first4=D. D. A. |last5=Imperatriz-Fonseca |first5=V. L. |date=June 2005 |title=Working-class royalty: Bees beat the caste system |journal=[[Biology Letters]] |volume=1 |issue=2 |pages=125–128 |doi=10.1098/rsbl.2004.0281 |pmc=1626201 |pmid=17148145}}{{open access}} * {{Cite journal |last1=Witter |first1=Sidia |last2=Blochtein |first2=Betina |last3=Santos |first3=Camila dos |date=August 2007 |title=Abelhas sem ferrão do Rio Grande do Sul: manejo e conservação |url=https://www.agricultura.rs.gov.br/upload/arquivos/202105/11142006-boletim-15.pdf |journal=Boletim FEPAGRO |issue=15}} @@ -595,4 +604,5 @@ * {{Cite book |last=Grüter |first=Christoph |url=http://link.springer.com/10.1007/978-3-030-60090-7 |title=Stingless Bees: Their Behaviour, Ecology and Evolution |date=2020 |publisher=Springer International Publishing |isbn=978-3-030-60089-1 |series=Fascinating Life Sciences |location=Cham |language=en |doi=10.1007/978-3-030-60090-7|s2cid=227250633 }} * {{Cite book |last=Jalil |first=Abu Hassan |url=https://books.google.com/books?id=fUh9BAAAQBAJ |title=Beescape for Meliponines: Conservation of Indo-Malayan Stingless Bees |date=2014 |publisher=Partridge Publishing Singapore |isbn=978-1-4828-2362-2 |language=ar}} +* {{Cite book |last=Kajobe |first=R. |url=https://books.google.com.br/books/about/Foraging_Behaviour_of_Equatorial_Afrotro.html?id=bMuezQEACAAJ&redir_esc=y |title=Foraging Behaviour of Equatorial Afrotropical Stingless Bees: Habitat Selection and Competition for Resources |date=2008 |publisher=Utrecht University |language=en}} * {{Cite book |last=Michener |first=Charles D. |url=https://books.google.com/books?id=bu_1gmY13FIC |title=The Bees of the World |date=2000 |publisher=Johns Hopkins Univ. Press |isbn=978-0-8018-6133-8 |location=Baltimore, Md.}} * {{Cite book |last=Roubik |first=David W. |url=https://www.cambridge.org/core/books/ecology-and-natural-history-of-tropical-bees/01B9D0798E5CBA8A68865ED0CC166A4A |title=Ecology and natural history of tropical bees |date=1989 |publisher=Cambridge Univ. Pr |isbn=978-0-521-26236-1 |series=Cambridge tropical biology series |location=Cambridge}} '
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[ 0 => 'While the existence of a soldier caste is well known in [[ant]]s and [[termite]]s, the phenomenon was unknown among bees until 2012, when some stingless bees were found to have a similar caste of defensive specialists that help guard the nest entrance against intruders.<ref>{{harvtxt|Grüter|Menezes|Imperatriz-Fonseca|Ratnieks|2012}}</ref> To date, at least 10 species have been documented to possess such "soldiers", including ''[[Tetragonisca angustula]]'', ''[[Tetragonisca fiebrigi|T. fiebrigi]]'', and ''[[Frieseomelitta longipes]]'', with the guards not only larger, but also sometimes a different color from ordinary workers.<ref>{{harvtxt|Grüter|Segers|Menezes|Vollet-Neto|Falcón|von Zuben|Bitondi|Nascimento|Almeida|2017}}</ref><ref>{{harvtxt|Grüter|2020|p=258}}</ref>', 1 => '[[Honey]], a prized product of bee colonies, is crafted through the processing of [[nectar]]s, [[Honeydew (secretion)|honeydews]], and fruit juices by worker bees. They store these collected substances in an extension of their gut called a [[crop (anatomy)|crop]]. Back at the hive, the bees ripen or dehydrate the nectar droplets by spinning them inside their mouthparts until honey is formed. Ripening concentrates the nectar and increases the sugar content, though it is not nearly as concentrated as the honey from ''Apis mellifera''. Stored in food pots, meliponines' honey is often referred to as pot-honey due to its distinctive storage method. Stingless bee honeys differ from ''A. mellifera'' honey in terms of color, texture, and flavor, being more liquid with a higher water content. Rich in [[mineral]]s, [[amino acid]]s, and [[flavonoid]] compounds, the composition of honey varies among colonies of the same species, influenced by factors such as season, habitat, and collected resources.<ref name=":10" /><ref name=":11" /><ref name=":12" /><ref name=":13" />', 2 => 'Many [[beekeeper]]s keep the bees in their original log [[beehive|hive]] or transfer them to a wooden box, as this makes controlling the hive easier. Some beekeepers put them in bamboos, flowerpots, coconut shells, and other recycling containers such as a water jug, a broken guitar, and other safe and closed containers.<ref>{{harvtxt|Venturieri|2004|pp=36-39}}</ref><ref>{{harvtxt|Contrera|Venturieri|2008}}</ref><ref name=":26" />', 3 => '<gallery perrow="4" widths="160" caption="Different species nest entrances in Brazilian stingless bees">', 4 => 'Bees play a critical role in the ecosystem, particularly in the pollination of natural vegetation. This activity is essential for the reproduction of various plant species, particularly in [[tropical forest]]s where most tree species rely on [[Entomophily|insect pollination]]. Even in temperate climates, where [[Anemophily|wind pollination]] is prevalent among forest trees, many bushes and herbaceous plants, rely on bees for pollination. The significance of bees extends to arid regions, such as [[Deserts and xeric shrublands|desertic and xeric shrublands]], where bee-pollinated plants are essential for preventing [[erosion]], supporting [[wildlife]], and ensuring [[Ecological stability|ecosystem stability]].<ref name=":23">{{harvtxt|Michener|2000|pp=4-5}}</ref>', 5 => '', 6 => '== Worldwide overview ==', 7 => '', 8 => '=== Africa ===', 9 => 'Stingless bees also play a vital ecological role across [[Sub-Saharan Africa]] and [[Madagascar]]. To understand these insects on the African continent, it's important to consider the prevailing socio-economic and cultural contexts. Despite their ecological significance, the diversity, conservation, and behavior of these bees remain underexplored, particularly compared to better-studied regions such as [[South America]] and [[Southeast Asia]]. Also, honeybees were extensively researched, in contrast to native meliponines.<ref name=":31">{{harvtxt|Kajobe|2008}}</ref><ref name=":32">{{harvtxt|Byarugaba|2004}}</ref> ', 10 => '', 11 => 'Africa is home to seven [[Biodiversity hotspot|biodiversity hotspots]], yet the recorded bee fauna is moderate relative to the continent's size. Madagascar stands out with exceptionally high levels of endemic species, though much of the bee diversity remains undocumented.<ref name=":30">{{harvtxt|Eardley|Gikungu|Schwarz|2009}}</ref>'''<ref name=":31" />''' Africa is home to aproximately 36 species of meliponines, including seven endemic to Madagascar. Most of these bees are found in equatorian regions (tropical forests and some savannahs).<ref>{{harvtxt|Grüter|2010|pp=47 & 49}}</ref> ', 12 => '', 13 => 'Factors such as habitat destruction, pesticide use, and invasive species pose significant threats to these pollinators. Furthermore, high rates of nest mortality, driven by predation and human activity, exacerbate conservation challenges. Research indicates that stingless bees in Africa face greater pressures than their counterparts in the American and Asian tropics, underlining the urgency for targeted conservation measures.<ref name=":30" />'''<ref name=":31" />'''', 14 => '', 15 => 'Uganda's [[Bwindi Impenetrable National Park]] has shown the presence of at least five stingless bee species in, distributed across two genera: ''[[Meliponula]]'' and ''[[Hypotrigona]]''.<ref name=":31" />'''<ref name=":32" />''' In Madagascar, there is only one genus of stingless bees: ''[[Liotrigona]].''<ref name=":30" />', 16 => '', 17 => 'Meliponiculture, for example, is practised in Angola and Tanzania, and interest in managing stingless bees is growing in other African countries as well. <ref>{{harvtxt|Grüter|2010|p=27}}</ref>', 18 => 'Of the 1,600 species of wild bees native to Australia, about 14 are meliponines.<ref>{{cite news | url=http://www.abc.net.au/science/articles/2003/05/08/2045526.htm | title=Stingless bee rescue |author= Wendy Pyper |date=May 8, 2003 |work=ABC Science}}</ref> "''Coot-tha''", which derives from "''ku-ta''", is one of the Aboriginal names for "wild stingless bee honey".<ref>{{harvtxt|Vit|Pedro|Roubik|2018}}</ref> These species bear a variety of names, including Australian native honey bees, native bees, sugar-bag bees, and sweat bees (because they land on people's skin to collect sweat).<ref>{{Cite web |title=Australian Native Stingless Bees |url=https://www.aussiebee.com.au/australian-stingless-bees.html |access-date=2024-05-28 |website=www.aussiebee.com.au}}</ref> The various stingless species look quite similar, with the two most common species, ''[[Tetragonula carbonaria]]'' and ''[[Austroplebeia australis]]'', displaying the greatest variation, as the latter is smaller and less active. Both of these inhabit the area around [[Brisbane]].<ref>{{Cite web |last=Steve |date=2015-05-25 |title=Native Stingless Bees - Tetragonula carbonaria |url=https://www.nativebeehives.com/native-stingless-bees-tetragonula-carbonaria/ |access-date=2024-05-28 |website=www.nativebeehives.com |language=en-AU}}</ref>', 19 => 'Brazil is home to several species bees belonging to Meliponini, with more than 300 species already identified and probably more yet to be discovered and described. They vary greatly in shape, size, and habits, and 20 to 30 of these species have good potential as honey producers. Although they are still quite unknown by most people, an increasing number of beekeepers (''meliponicultores'', in Portuguese) have been dedicated to these bees throughout the country.<ref name=":27" /><ref>{{harvtxt|Villas-Bôas|2018|p=17}}</ref> This activity has experienced significant growth since August 2004, when national laws were changed to allow native bee colonies to be freely marketed, which was previously forbidden in an unsuccessful attempt to protect these species. Nowadays the capture or destruction of existing colonies in nature is still forbidden, and only new colonies formed by the bees themselves in artificial traps can be collected from the wild.<ref>[https://conama.mma.gov.br/?option=com_sisconama&task=arquivo.download&id=440 CONAMA 2004 Resolution]</ref> Most marketed colonies are artificially produced by authorized beekeepers, through division of already existing captive colonies. Besides honey production, Brazilian stingless bees such as the jataí (''[[Tetragonisca angustula]]''), mandaguari (''[[Scaptotrigona postica]]''), and mandaçaia (''[[Melipona quadrifasciata]]'') serve as major pollinators of tropical plants and are considered the ecological equivalent of the honey bee.<ref name=":21" /><ref name=":22" />', 20 => 'Also, much practical and academic work is being done about the best ways of keeping such bees, multiplying their colonies, and exploring the honey they produce.<ref>{{Cite web |date=2024-07-26 |title=Arquivos Artigos - A.B.E.L.H.A. |url=https://abelha.org.br/artigos/ |access-date=2024-12-31 |website=abelha.org.br |language=pt-BR}}</ref> Among many others, species such as jandaíra (''[[Melipona subnitida]]'') and true ''uruçu'' (''[[Melipona scutellaris]]'') in the northeast of the country, mandaçaia (''[[Melipona quadrifasciata]]'') and yellow uruçu (''Melipona rufiventris'') in the south-southeast, tiúba or jupará (''[[Melipona interrupta]]'') and canudo (''[[Scaptotrigona polysticta]]'') in the north and jataí (''[[Tetragonisca angustula]]'') throughout the country are increasingly kept by small, medium, and large producers. Many other species as the mandaguari (''[[Scaptotrigona postica]]''), the guaraipo (''[[Melipona bicolor]]''), marmelada (''[[Frieseomelitta varia]]'') and the iraí (''[[Nannotrigona testaceicornis]]''), to mention a few, are also reared.<ref name=":33">{{Cite web |date=2022-02-21 |title=Fichas catalográficas das espécies relevantes para a meliponicultura - A.B.E.L.H.A. |url=https://abelha.org.br/fichas-catalograficas-das-especies-relevantes-para-a-meliponicultura-2/ |access-date=2024-12-31 |website=abelha.org.br |language=pt-BR}}</ref>', 21 => 'Although the colony population of most of these bees is much smaller than that of European bees, the productivity per bee can be quite high. Interestingly, honey production is more connected to the body size, not the colony size. The manduri (''[[Melipona marginata]]''), jandaíra (''[[Melipona subnitida]]'') and the ''guaraipo'' (''[[Melipona bicolor|M. bicolor]]'') live in swarms of only around 300 individuals but can still produce up to 5 liters (1.3 US gallon) of honey a year under the right conditions.<ref>{{harvtxt|Grüter|2010|p=14}}</ref> In large bee farms, only the availability of flowers limits the honey production per colony. However, much larger numbers of beehives are required to produce amounts of honey comparable to that of European bees. Also, due to the fact of those bees storing honey in cerumen pots instead of standardized honeycombs as in the honeybee rearing makes extraction a lot more difficult and laborious.<ref>{{harvtxt|Fonseca|Sodré|Carvalho|Alves|Souza|Silva|Oliveira|Machado|Clarton|2007}}</ref>', 22 => 'The honey from stingless bees has a higher water content, from 25% to 35%, compared to the honey from the genus ''Apis''. This contributes to its less cloying taste but also causes it to spoil more easily. Thus, for marketing, this honey needs to be processed through desiccation, fermentation or pasteurization. In its natural state, it should be kept under refrigeration.<ref>{{harvtxt|Grüter|2020|pp=12-14}}</ref>', 23 => 'Due to the lack of a functional stinger and characteristic nonaggressive behavior of many Brazilian species of stingless bees, they can be reared without problems in densely populated environments (residential buildings, schools, urban parks), provided enough flowers are at their disposal nearby. Some breeders (''meliponicultores'') can produce honey even in apartments up to the 12th floor.<ref>{{cite web|url=http://meliponariocapixaba.blogspot.com.br/2009/11/e-possivel-criar-abelhas-em.html|title=Meliponario Capixaba: É POSSÍVEL CRIAR ABELHAS EM APARTAMENTOS?|author=João Luiz|access-date=12 March 2016}}</ref>', 24 => 'The [[Melipona quadrifasciata|mandaçaias]] (''Melipona quadrifasciata'') are extremely tame, rarely attacking humans (only when their hives are opened for honey extraction or colony division). They form small, manageable colonies of only 400&ndash;600&nbsp;individuals. They are fairly large bees, up to 11&nbsp;mm (7/16") in length, and as a result have better body heat control, allowing them to live in regions where temperatures can drop a little lower than 0&nbsp;°C (32&nbsp;°F). However, they are somewhat selective about which flowers they will visit, preferring the flora that occurs in their natural environment. They are thus difficult to keep outside their region of origin (the eastern coast of Brazil). Once very common, the mandaçaia is now rather rare in nature, mainly due to the destruction of their native forests in the of Brazil.<ref>{{harvtxt|Brito|Faquinello|Paula-Leite|Carvalho|1997}}</ref><ref name=":33" />', 25 => 'Other groups of Brazilian stingless bees, genera ''[[Plebeia]]'' and ''[[Leurotrigona]]'', are also very tame and much smaller, with one of them (''[[Plebeia minima]]'') reaching no more than 2.5&nbsp;mm (3/32") in length, and the ''[[lambe-olhos]]'' ("lick-eyes" bee, ''[[Leurotrigona muelleri]]'') being even smaller, at no more than 1.5&nbsp;mm (3/32"). Many of these species are known as mirim (meaning 'small' in the [[Tupi-Guarani languages]]). As a result, they can be kept in very small artificial hives, thus being of interest for keepers who want them as pollinators in small glasshouses or just for the pleasure of having a 'toy' bee colony at home.<ref name=":33" /><ref>{{Cite web |title=Lambe-olhos – Leurotrigona muelleri – Laboratório de Sistemática de Plantas |url=https://sites.usp.br/jardimdabotanicausprp/lambe-olhos-leurotrigona-muelleri/ |access-date=2024-12-31 |website=sites.usp.br}}</ref><ref>{{Cite web |title=Mirim – Plebeia droryana – Laboratório de Sistemática de Plantas |url=https://sites.usp.br/jardimdabotanicausprp/mirim-plebeia-droryana/ |access-date=2024-12-31 |website=sites.usp.br}}</ref> Being so tiny, these species produce only a very small amount of honey, typically less than 500&nbsp;ml (1/2 US pint) a year, so are not interesting for commercial honey production.', 26 => '[[Image:Abelha-jatai.jpg|right|thumb|Entrance pipe of ''jataís´s'' colony: The ''jataí'' bees build a wax pipe in the entrance of their nest.]]', 27 => 'Belonging to the same group, the jataí (''[[Tetragonisca angustula]]''), the marmelada (''[[Frieseomelitta varia]]''), and the moça-branca (''[[Frieseomelitta doederleini]]'') are intermediate in size between those very small species and the European bee. They are very adaptable species; the jataí, and can be reared in many different regions and environments, being quite common in most Brazilian cities. The jataí can bite when disturbed, but its jaws are weak, and in practice they are harmless, while the marmelada and moça-branca usually deposit propolis on their aggressors. Jataí is one of the first species to be kept by home beekeepers. Their nests can be easily identified in trees or wall cavities by the yellow wax pipe they build at the entrance, usually guarded by some soldier bees, which are stronger than regular worker bees. The marmelada and moça-branca make a little less honey, but it is denser and sweeter than most from other stingless bees and is considered very tasty.<ref name=":33" /><ref>{{harvtxt|Castanheira|Contel|2005|}}</ref>', 28 => '=== Central America ===', 29 => 'Native meliponines (''M. beecheii'' being the most common) have been kept by the lowland Maya for thousands of years. The [[Yucatec Maya language]] name for this bee is ''xunan kab'', meaning "(royal, noble) lady bee".<ref>{{harvtxt|Grüter|2010|p=54}}</ref><ref>{{cite web|title=Diccionario Introductorio|url=http://www.uqroo.mx/libros/maya/diccionario.pdf|work=uqroo.mx|publisher=Universidad De Quintana Roo|access-date=27 March 2014|language=es}}</ref> The bees were once the subject of religious ceremonies and were a symbol of the bee-god [[Ah-Muzen-Cab]], known from the ''[[Madrid Codex (Maya)|Madrid Codex]]''.<ref name=":34">{{Cite web |date=2016-07-17 |title=People and bees. Mayan bees in the Madrid Codex |url=https://ecomercioagrario.com/en/people-and-bees-mayan-bees-in-the-madrid-codex/ |access-date=2024-12-31 |website=eComercio Agrario |language=en-US}}</ref>', 30 => 'The bees were, and still are, treated as pets. Families would have one or many log-hives hanging in and around their houses. Although they are stingless, the bees do bite and can leave welts similar to a mosquito bite. The traditional way to gather bees, still favored among the locals, is find a wild hive, then the branch is cut around the hive to create a portable log, enclosing the colony. With proper maintenance, hives have been recorded as lasting over 80 years, being passed down through generations. In the archaeological record of Mesoamerica, stone discs have been found that are generally considered to be the caps of long-disintegrated logs that once housed the beehives.<ref name=":34" /><ref name=":35">{{Cite web |last=Kennedy |first=Jennifer |last2=Arghiris |first2=Richard |date=2019-01-31 |title=House of the Royal Lady Bee: Maya revive native bees and ancient beekeeping |url=https://news.mongabay.com/2019/01/house-of-the-royal-lady-bee-maya-revive-native-bees-and-ancient-beekeeping/ |access-date=2024-12-31 |website=Mongabay Environmental News |language=en-US}}</ref>', 31 => '[[Tulum]], the site of a [[pre-Columbian]] Maya city on the Caribbean coast 130&nbsp;km (81&nbsp;mi) south of Cancun, has a god depicted repeatedly all over the site. Upside down, he appears as a small figure over many doorways and entrances. One of the temples, the Temple of the Descending God (''Templo del Dios Descendente''), stands just left of the central plaza. Speculation is that he may be the "Bee God", Ah Muzen Cab, as seen in the ''Madrid Codex''. It is possible that this was a religious/trade center with emphasis on ''xunan kab'', the "royal lady".<ref name=":35" />', 32 => '[[Balché]], a traditional [[Mesoamerica]]n alcoholic beverage similar to [[mead]], was made from fermented honey and the bark of the leguminous'' balché'' tree (''[[Lonchocarpus violaceus]]''), hence its name. It was traditionally brewed in a canoe. The drink was known to have [[entheogen]]ic properties, that is, to produce mystical experiences, and was consumed in medicinal and ritual practices. Beekeepers would place the nests near the psychoactive plant ''[[Turbina corymbosa]]'' and possibly near ''balché'' trees, forcing the bees to use nectar from these plants to make their honey. Additionally, brewers would add extracts of the bark of the ''balché'' tree to the honey mixture before fermentation. The resulting beverage is responsible for psychotropic effects when consumed, due to the ergoline compounds in the pollen of the ''T. corymbosa'', the ''Melipona'' nectar gathered from the ''balché'' flowers, or the hallucinogenic compounds of the ''balché'' tree bark.<ref>{{harvtxt|Ott|1998}}</ref>', 33 => '[[Lost-wax casting]], a common metalworking method typically found where the inhabitants keep bees, was also used by the Maya. The wax from ''Melipona'' is soft and easy to work, especially in the humid Maya lowland. This allowed the Maya to create smaller works of art, jewelry, and other metalsmithing that would be difficult to forge. It also makes use of the leftovers from honey extraction. If the hive was damaged beyond repair, the whole of the comb could be used, thus using all of the hive. With experienced keepers, though, only the honey pot could be removed, the honey extracted, and the wax used for casting or other purposes.<ref>{{harvtxt|Pitses|2018|}}</ref><ref>{{harvtxt|Schwarz|1945|}}</ref>', 34 => 'The outlook for meliponines in Mesoamerica is uncertain. The number of active Meliponini beekeepers is shy in comparison with the Africanized ''[[Apis mellifera]]'' breeders. The high honey yield, 100&nbsp;kg (220&nbsp;lbs) or more annually, along with the ease of hive care and ability to create new hives from existing stock, commonly outweighs the negative consequences of "killer bee" hive maintenance.<ref name=":36">{{harvtxt|Villanueva-G|Roubik|Colli-Ucán|2005|}}</ref>', 35 => 'An additional blow to the art of meliponine beekeeping is that many of the ''meliponicultores'' are now elderly, and their hives may not be cared for once they die. The hives are considered similar to an old family collection, to be parted out once the collector dies or to be buried in whole or part along with the beekeeper upon death. In fact, a survey of a once-popular area of the Maya lowlands shows the rapid decline of beekeepers, down to around 70 in 2004 from thousands in the late 1980s. Conservation efforts are underway in several parts of Mesoamerica.<ref name=":36" /><ref>A comprehensive conservation guide can be found in the June 2005 issue of ''Bee World''.</ref>', 36 => '* {{Cite journal |last=Byarugaba |first=Dominic |date=2004 |title=Stingless bees (Hymenoptera: Apidae) of Bwindi impenetrable forest, Uganda and Abayanda indigenous knowledge |url=http://www.journals.cambridge.org/abstract_S1742758404000128 |journal=International Journal of Tropical Insect Science |language=en |volume=24 |issue=01 |doi=10.1079/IJT20048 |issn=1742-7584}}', 37 => '* {{Cite journal |last=Castanheira |first=Eliana Barroso |last2=Contel |first2=Eucleia Primo Betioli |date=2005-01-01 |url=https://www.tandfonline.com/doi/abs/10.1080/00218839.2005.11101157 |doi=10.1080/00218839.2005.11101157 |issn=0021-8839}}', 38 => '* {{Cite journal |last=Eardley |first=Connal D. |last2=Gikungu |first2=Mary |last3=Schwarz |first3=Michael P. |date=2009 |title=Bee conservation in Sub-Saharan Africa and Madagascar: diversity, status and threats |url=http://link.springer.com/10.1051/apido/2009016 |journal=Apidologie |volume=40 |issue=3 |pages=355–366 |doi=10.1051/apido/2009016 |issn=0044-8435}}', 39 => '* {{Cite journal |last=Kent |first=Robert B. |date=1984 |title=Mesoamerican Stingless Beekeeping |url=http://www.tandfonline.com/doi/abs/10.1080/08873638409478571 |journal=Journal of Cultural Geography |language=en |volume=4 |issue=2 |pages=14–28 |doi=10.1080/08873638409478571 |issn=0887-3631}}', 40 => '* {{Cite journal |last=Kerr |first=Warwick Estevam |last2=Petrere Jr. |first2=Miguel |last3=Diniz Filho |first3=José Alexandre Felizola |date=2001-03 |title=Informações biológicas e estimativa do tamanho ideal da colmeia para a abelha tiúba do Maranhão (Melipona compressipes fasciculata Smith - Hymenoptera, Apidae) |url=https://www.scielo.br/j/rbzool/a/r74hTMg94tR9PTJGr8cvJwH/?lang=pt |journal=Revista Brasileira de Zoologia |language=pt |volume=18 |pages=45–52 |doi=10.1590/S0101-81752001000100003 |issn=0101-8175}}', 41 => '* {{Cite journal |last=Schwarz |first=Herbert F. |date=1945 |title=The Wax of Stingless Bees (Meliponidæ) and the Uses to Which It Has Been Put |url=https://www.jstor.org/stable/25005104?utm_source=chatgpt.com |journal=Journal of the New York Entomological Society |volume=53 |issue=2 |pages=137–144 |issn=0028-7199}}', 42 => '* {{Cite journal |last=Villanueva-G |first=Rogel |last2=Roubik |first2=David W |last3=Colli-Ucán |first3=Wilberto |date=2005-01 |title=Extinction of Melipona beecheii and traditional beekeeping in the Yucatán peninsula |url=http://www.tandfonline.com/doi/full/10.1080/0005772X.2005.11099651 |journal=Bee World |language=en |volume=86 |issue=2 |pages=35–41 |doi=10.1080/0005772X.2005.11099651 |issn=0005-772X}}', 43 => '* {{Cite book |last=Kajobe |first=R. |url=https://books.google.com.br/books/about/Foraging_Behaviour_of_Equatorial_Afrotro.html?id=bMuezQEACAAJ&redir_esc=y |title=Foraging Behaviour of Equatorial Afrotropical Stingless Bees: Habitat Selection and Competition for Resources |date=2008 |publisher=Utrecht University |language=en}}' ]
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[ 0 => 'While the existence of a soldier caste is well known in [[ant]]s and [[termite]]s, the phenomenon was unknown among bees until 2012, when some stingless bees were found to have a similar caste of defensive specialists that help guard the nest entrance against intruders.<ref>{{harvtxt|Grüter|Menezes|Imperatriz-Fonseca|Ratnieks|2012}}</ref> To date, at least 10 species have been documented to possess such "soldiers", including ''[[Tetragonisca angustula]]'', ''[[Tetragonisca fiebrigi|T. fiebrigi]]'', and ''[[Frieseomelitta longipes]]'', with the guards not only larger, but also sometimes a different color from ordinary workers.<ref>{{harvtxt|Grüter|Segers|Menezes|Vollet-Neto|Falcón|von Zuben|Bitondi|Nascimento|Almeida|2017}}</ref><ref>{{harvtxt|Grüter|2020|p=258}}</ref>', 1 => '[[Honey]], a prized product of bee colonies, is crafted through the processing of [[nectar]]s, [[Honeydew (secretion)|honeydews]], and fruit juices by worker bees. They store these collected substances in an extension of their gut called a [[crop (anatomy)|crop]]. Back at the hive, the bees ripen or dehydrate the nectar droplets by spinning them inside their mouthparts until honey is formed. Ripening concentrates the nectar and increases the sugar content, though it is not nearly as concentrated as the honey from ''Apis mellifera''. Stored in food pots, meliponines' honey is often referred to as pot-honey due to its distinctive storage method. Stingless bee honeys differ from ''A. mellifera'' honey in terms of color, texture, and flavor, being more liquid with a higher water content. Rich in [[mineral]]s, [[amino acid]]s, and [[flavonoid]] compounds, the composition of honey varies among colonies of the same species, influenced by factors such as season, habitat, and collected resources.<ref name=":10" /><ref name=":11" /><ref name=":12" /><ref name=":13" />', 2 => 'Many [[beekeeper]]s keep the bees in their original log [[beehive|hive]] or transfer them to a wooden box, as this makes controlling the hive easier. Some beekeepers put them in bamboos, flowerpots, coconut shells, and other recycling containers such as a water jug, a broken guitar, and other safe and closed containers.<ref>{{harvtxt|Venturieri|2004|pp=36-39}}</ref><ref>{{harvtxt|Contrera|Venturieri|2008}}</ref><ref name=":26" />', 3 => '<gallery perrow="4" widths="160" caption="Different species nest entrances in Brazilian stingless bees">', 4 => 'Bees play a critical role in the ecosystem, particularly in the pollination of natural vegetation. This activity is essential for the reproduction of various plant species, particularly in [[tropical forest]]s where most tree species rely on [[Entomophily|insect pollination]]. Even in temperate climates, where [[Anemophily|wind pollination]] is prevalent among forest trees, many bushes and herbaceous plants, rely on bees for pollination. The significance of bees extends to arid regions, such as [[Deserts and xeric shrublands|desertic and xeric shrublands]], where bee-pollinated plants are essential for preventing [[erosion]], supporting [[wildlife]], and ensuring [[Ecological stability|ecosystem stability]].<ref name=":23">{{harvtxt|Michener|2000|pp=4-5}}</ref>', 5 => '== Relationship to humans ==', 6 => '"''Coot-tha''", which derives from "''ku-ta''", is one of the Aboriginal names for "wild stingless bee honey".<ref>{{harvtxt |Vit|Pedro|Roubik|2018}}</ref> Of the 1,600 species of wild bees native to Australia, about 14 are meliponines.<ref>{{cite news | url=http://www.abc.net.au/science/articles/2003/05/08/2045526.htm | title=Stingless bee rescue |author= Wendy Pyper |date=May 8, 2003 |work=ABC Science}}</ref> These species bear a variety of names, including Australian native honey bees, native bees, sugar-bag bees, and sweat bees (because they land on people's skin to collect sweat).<ref>{{Cite web |title=Australian Native Stingless Bees |url=https://www.aussiebee.com.au/australian-stingless-bees.html |access-date=2024-05-28 |website=www.aussiebee.com.au}}</ref> All are small and usually black in colour, with hairy extended hind legs for carrying [[nectar]] and [[pollen]]; because of the latter, they are sometimes mistaken for [[bumblebee]]s. The various stingless species look quite similar, with the two most common species, ''[[Tetragonula carbonaria]]'' and ''[[Austroplebeia australis]]'', displaying the greatest variation, as the latter is smaller and less active. Both of these inhabit the area around [[Brisbane]].<ref>{{Cite web |last=Steve |date=2015-05-25 |title=Native Stingless Bees - Tetragonula carbonaria |url=https://www.nativebeehives.com/native-stingless-bees-tetragonula-carbonaria/ |access-date=2024-05-28 |website=www.nativebeehives.com |language=en-AU}}</ref>', 7 => 'Brazil is home to several species bees belonging to Meliponini, with more than 300 species already identified and probably more yet to be discovered and described. They vary greatly in shape, size, and habits, and 20 to 30 of these species have good potential as honey producers. Although they are still quite unknown by most people, an increasing number of beekeepers (''meliponicultores'', in Portuguese) have been dedicated to these bees throughout the country.<ref name=":27" /><ref>{{harvtxt|Villas-Bôas|2018|p=17}}</ref> This activity has experienced significant growth since August 2004, when national laws were changed to allow native bee colonies to be freely marketed, which was previously forbidden in an unsuccessful attempt to protect these species. Nowadays the capture or destruction of existing colonies in nature is still forbidden, and only new colonies formed by the bees themselves in artificial traps can be collected from the wild.<ref>[https://conama.mma.gov.br/?option=com_sisconama&task=arquivo.download&id=440 CONAMA 2004 Resolution]</ref> Most colonies marketed are artificially produced by authorized beekeepers, through division of already existing captive colonies. Besides honey production, Brazilian stingless bees such as the jataí (''[[Tetragonisca angustula]]''), mandaguari (''[[Scaptotrigona postica]]''), and mandaçaia (''[[Melipona quadrifasciata]]'') serve as major pollinators of tropical plants and are considered the ecological equivalent of the honey bee.<ref name=":21" /><ref name=":22" />', 8 => 'Also, much practical and academic work is being done about the best ways of keeping such bees, multiplying their colonies, and exploring the honey they produce. Among many others, species such as jandaíra (''[[Melipona subnitida]]'') and true ''uruçu'' (''[[Melipona scutellaris]]'') in the northeast of the country, mandaçaia (''[[Melipona quadrifasciata]]'') and yellow uruçu (''Melipona rufiventris'') in the south-southeast, tiúba or jupará (''[[Melipona interrupta]]''<ref>{{Cite web|url=http://moure.cria.org.br/catalogue?id=34747|title=+Moure's Bee Catalogue}}</ref>) and straw-bee (''[[Scaptotrigona polysticta]]'') in the north and jataí (''[[Tetragonisca angustula]]'') throughout the country are increasingly kept by small, medium, and large producers. Many other species as the mandaguari (''[[Scaptotrigona postica]]''), the guaraipo (''[[Melipona bicolor]]''), marmelada (''[[Frieseomelitta varia]]'') and the iraí (''[[Nannotrigona testaceicornis]]''), to mention a few, are also reared. Through the cultivation of honey or selling of colonies, keeping stingless bees is an increasingly profitable activity. A single colony of species like mandaçaia and true "uruçu" can be divided up to four times a year, and each of the new colonies obtained this way can be sold for about US$100.', 9 => 'Although the colony population of most of these bees is much smaller than that of European bees, the productivity per bee can be quite high. Probably the world champions in honey productivity, the manduri (''[[Melipona marginata]]'') and the ''guaraipo'' (''M. bicolor''), live in swarms of only around 300 individuals, but can still produce up to 5 liters (.79 US gallon) of honey a year under the right conditions. The manduri is one of the smallest bees in the genus ''[[Melipona]]'', varying in length from 6 to 7 mm (15/64" to 9/32").<ref>{{harvtxt|Witter|Blochtein|Santos|2007|p=27}}</ref> Species of the genus ''Scaptotrigona'' have very large colonies, with up to 20,000 individuals, and can produce from 3 to 8 liters (2 US gallons) of honey a year.', 10 => '', 11 => 'Some large breeders have more than 3,000 hives of the tamer but still highly productive species in the genus ''Melipona'', such as the tiúba, the true ''uruçu'', and the jandaíra, each with 3,000 or more bees per colony. They can produce over 1.5 tons (3,000&nbsp;lbs) of honey every year.{{Citation needed|date=June 2024}} In large bee farms, only the availability of flowers limits the honey production per colony. Their honey is considered more palatable because it's not overly sweet, and it also is thought to have medicinal properties more pronounced than honey from bees of the genus ''Apis'' due to the higher level of antimicrobial substances. As a result, the honey from stingless bees returns very high profits, with prices greater than those for the more common honey produced by European bees. However, much larger numbers of beehives are required to produce amounts of honey comparable to that of European bees. Also, due to the fact of those bees storing honey in cerumen pots instead of standardized honeycombs as in the honeybee rearing makes extraction a lot more difficult and laborious.<ref>{{harvtxt|Fonseca|Sodré|Carvalho|Alves|Souza|Silva|Oliveira|Machado|Clarton|2007}}</ref>', 12 => '', 13 => 'The honey from stingless bees has a lighter color and a higher water content, from 25% to 35%, compared to the honey from the genus ''Apis''. This contributes to its less cloying taste but also causes it to spoil more easily. Thus, for marketing, this honey needs to be processed through desiccation, fermentation or pasteurization. In its natural state, it should be kept under refrigeration.<ref>{{harvtxt|Grüter|2020|pp=12-14}}</ref>', 14 => 'Due to the lack of a functional stinger and characteristic nonaggressive behavior of many Brazilian species of stingless bees, they can be reared without problems in densely populated environments such as cities, provided enough flowers are at their disposal nearby. Some breeders (''meliponicultores'') can produce honey even in apartments up to the 12th floor.<ref>{{cite web|url=http://meliponariocapixaba.blogspot.com.br/2009/11/e-possivel-criar-abelhas-em.html|title=Meliponario Capixaba: É POSSÍVEL CRIAR ABELHAS EM APARTAMENTOS?|author=João Luiz|access-date=12 March 2016}}</ref>', 15 => 'The [[Melipona quadrifasciata|mandaçaias]] (''Melipona quadrifasciata'') are extremely tame, rarely attacking humans (only when their hives are opened for honey extraction or colony division). They form small, manageable colonies of only 400&ndash;600&nbsp;individuals. At the same time, a single rational beehive of ''mandaçaia'' can produce up to 4&nbsp;liters (1&nbsp;US gallon) of honey a year, making the species very attractive for home keepers. They are fairly large bees, up to 11&nbsp;mm (7/16") in length, and as a result have better body heat control, allowing them to live in regions where temperatures can drop a little lower than 0&nbsp;°C (32&nbsp;°F). However, they are somewhat selective about which flowers they will visit, preferring the flora that occurs in their natural environment. They are thus difficult to keep outside their region of origin (the northeast coast of Brazil).<ref>{{harvtxt|Brito|Faquinello|Paula-Leite|Carvalho|1997}}</ref>', 16 => 'Other species, like the ''tiúba'' and the true ''uruçu'', are also very tame and highly productive. Their colonies have from 3,000&ndash;5,000 individuals (for comparison, the population of honeybee swarms can peak at 80,000&nbsp;individuals) and can produce up to 10 liters (2.6 US gallons) of honey a year.{{Citation needed|date=June 2024}} They can be easily kept at home but will survive only in regions with a warm climate, their larvae dying at temperatures lower than 12&nbsp;°C (54&nbsp;°F). The yellow ''uruçu'', however, can survive at temperatures lower than 0&nbsp;°C (32&nbsp;°F), and their colonies, bearing about 3,500&nbsp;individuals, can produce up to 6&nbsp;liters (1.5&nbsp;US gallons) of honey a year.{{Citation needed|date=June 2024}} But this species will react with powerful nipping if its nests are disturbed, and usually they are only kept by professional meliponicultors.', 17 => 'Another suitable species for keeping at home is the ''guaraipo'' (''[[Melipona bicolor]]''). It is also quite tame, never attacking the beekeeper, and their colonies have fewer than 600 individuals. They can withstand temperatures as low as &minus;10&nbsp;°C (14&nbsp;°F), and each colony can produce over 3&nbsp;liters ({{frac|3|4}}&nbsp;US gallons) of honey a year.{{Citation needed|date=June 2024}} Their colonies usually have more than one single queen at a time (usually two or three, but sometimes up to five), a phenomenon called [[Gyne|polygyne]], and thus are less sensitive to the death of one queen, which can cause the loss of a whole colony in other species. But the ''guaraipo'' is very sensitive to low levels of humidity, and their hives must be equipped with means to keep a high moisture content. Once very common, the ''guaraipo'' is now rather rare in nature, mainly due to the destruction of their native forests in the south-southeast of Brazil.', 18 => 'Other groups of Brazilian stingless bees, genera ''[[Plebeia]]'' and ''[[Leurotrigona]]'', are also very tame and much smaller, with one of them (''[[Plebeia minima]]'') reaching no more than 2.5&nbsp;mm (3/32") in length, and the ''[[lambe-olhos]]'' ("lick-eyes" bee, ''[[Leurotrigona muelleri]]'') being even smaller, at no more than 1.5&nbsp;mm (3/32"). Many of these species are known as ''mirim'' (meaning 'small' in the [[Tupi-Guarani languages]]). As a result, they can be kept in very small artificial hives, thus being of interest for keepers who want them as pollinators in small glasshouses or just for the pleasure of having a 'toy' bee colony at home. Being so tiny, these species produce only a very small amount of honey, typically less than 500&nbsp;ml (1/2 US pint) a year, so are not interesting for commercial honey production.', 19 => '', 20 => '[[Image:Abelha-jatai.jpg|right|thumb|Entrance pipe of ''jataís´s'' colony: The ''jataí'' bees build a wax pipe in the entrance of their nest.]]', 21 => 'Belonging to the same group, the ''jataí'' (''[[Tetragonisca angustula]]''), the ''marmelada ''(''[[Frieseomelitta varia]]''), and the ''moça-branca'' (''[[Frieseomelitta doederleini]]'') are intermediate in size between those very small species and the European bee. They are very adaptable species; the ''jataí'', and can be reared in many different regions and environments, being quite common in most Brazilian cities. The ''jataí'' can bite when disturbed, but its jaws are weak, and in practice they are harmless, while the ''marmelada'' and ''moça-branca'' usually deposit propolis on their aggressors. Producing up to 1.5&nbsp;liters (0.4&nbsp;US gallons) of honey a year, their honey is considered among the best from stingless bees. In fact, the ''jataí'' was one of the first species to be kept by home beekeepers. Their nests can be easily identified in trees or wall cavities by the yellow wax pipe they build at the entrance, usually guarded by some soldier bees, which are stronger than regular worker bees. The'' marmelada'' and ''moça-branca'' make a little less honey, but it is denser and sweeter than most from other stingless bees and is considered very tasty.', 22 => '', 23 => '=== Maya stingless bees of Central America ===', 24 => 'Native meliponines (''M. beecheii'' being the favorite) have been kept by the lowland Maya for thousands of years. The [[Yucatec Maya language]] name for this bee is ''xunan kab'', meaning "(royal, noble) lady bee".<ref>{{cite web|title=Diccionario Introductorio|url=http://www.uqroo.mx/libros/maya/diccionario.pdf|work=uqroo.mx|publisher=Universidad De Quintana Roo|access-date=27 March 2014|language=es}}</ref> The bees were once the subject of religious ceremonies and were a symbol of the bee-god [[Ah-Muzen-Cab]], known from the ''[[Madrid Codex (Maya)|Madrid Codex]]''.', 25 => 'The bees were, and still are, treated as pets. Families would have one or many log-hives hanging in and around their houses. Although they are stingless, the bees do bite and can leave welts similar to a mosquito bite. The traditional way to gather bees, still favored among the locals, is find a wild hive, then the branch is cut around the hive to create a portable log, enclosing the colony. This log is then capped on both ends with another piece of wood or pottery and sealed with mud. This clever method keeps the melipine bees from mixing their brood, pollen, and honey in the same comb as do the European bees. The brood is kept in the middle of the hive, and the honey is stored in vertical "pots" on the outer edges of the hive. A temporary, replaceable cap at the end of the log allows for easy access to the honey while doing minimal damage to the hive. However, inexperienced handlers can still do irreversible damage to a hive, causing the hive to swarm and abscond from the log. With proper maintenance, though, hives have been recorded as lasting over 80 years, being passed down through generations. In the archaeological record of Mesoamerica, stone discs have been found that are generally considered to be the caps of long-disintegrated logs that once housed the beehives.', 26 => '[[Tulum]], the site of a [[pre-Columbian]] Maya city on the Caribbean coast 130&nbsp;km (81&nbsp;mi) south of Cancun, has a god depicted repeatedly all over the site. Upside down, he appears as a small figure over many doorways and entrances. One of the temples, the "Templo del Dios Descendente" or the Temple of the Descending God, stands just left of the central plaza. Speculation is that he may be the "Bee God", Ah Muzen Cab, as seen in the Madrid Codex. It is possible that this was a religious/trade center with emphasis on ''xunan kab'', the "royal lady".', 27 => '[[Balché]], a traditional [[Mesoamerica]]n alcoholic beverage similar to [[mead]], was made from fermented honey and the bark of the leguminous'' balché'' tree (''[[Lonchocarpus violaceus]]''), hence its name.<ref>{{harvtxt|Ott|1998}}</ref> It was traditionally brewed in a canoe. The drink was known to have [[entheogen]]ic properties, that is, to produce mystical experiences, and was consumed in medicinal and ritual practices. Beekeepers would place the nests near the psychoactive plant ''[[Turbina corymbosa]]'' and possibly near ''balché'' trees, forcing the bees to use nectar from these plants to make their honey.<ref>{{harvtxt|Ott|1998}}</ref> Additionally, brewers would add extracts of the bark of the ''balché'' tree to the honey mixture before fermentation. The resulting beverage is responsible for psychotropic effects when consumed, due to the ergoline compounds in the pollen of the ''T. corymbosa'', the ''Melipona'' nectar gathered from the ''balché'' flowers, or the hallucinogenic compounds of the ''balché'' tree bark.', 28 => '[[Lost-wax casting]], a common metalworking method typically found where the inhabitants keep bees, was also used by the Maya. The wax from ''Melipona'' is soft and easy to work, especially in the humid Maya lowland. This allowed the Maya to create smaller works of art, jewelry, and other metalsmithing that would be difficult to forge. It also makes use of the leftovers from honey extraction. If the hive was damaged beyond repair, the whole of the comb could be used, thus using all of the hive. With experienced keepers, though, only the honey pot could be removed, the honey extracted, and the wax used for casting or other purposes.', 29 => 'The outlook for meliponines in Mesoamerica is uncertain. The number of active Meliponini beekeepers is shy in comparison with the Africanized ''[[Apis mellifera]]'' breeders. The high honey yield, 100&nbsp;kg (220&nbsp;lbs) or more annually, along with the ease of hive care and ability to create new hives from existing stock, commonly outweighs the negative consequences of "killer bee" hive maintenance.', 30 => '', 31 => 'Further complicating the issue, [[Africanized honey bee]]s do not visit some flora, such as those in the [[Solanaceae|tomato family]], and several forest trees and shrubs, which rely on the native stingless bees for pollination. A decline in populations of native flora has already occurred in areas where stingless bees have been displaced by Africanized honey bees.', 32 => 'An additional blow to the art of meliponine beekeeping is that many of the meliponine beekeepers are now elderly, and their hives may not be cared for once they die. The hives are considered similar to an old family collection, to be parted out once the collector dies or to be buried in whole or part along with the beekeeper upon death. In fact, a survey of a once-popular area of the Maya lowlands shows the rapid decline of beekeepers, down to around 70 in 2004 from thousands in the late 1980s. It is traditional in the Maya lowlands that the hive itself or parts of the hive be buried along with the beekeeper to ''volar al cielo'', "to fly to heaven".{{citation needed|date=March 2007}} Conservation efforts are underway in several parts of Mesoamerica.<ref>A comprehensive conservation guide can be found in the June 2005 issue of ''Bee World''.</ref>', 33 => '* <{{Cite journal |last=Kent |first=Robert B. |date=1984 |title=Mesoamerican Stingless Beekeeping |url=http://www.tandfonline.com/doi/abs/10.1080/08873638409478571 |journal=Journal of Cultural Geography |language=en |volume=4 |issue=2 |pages=14–28 |doi=10.1080/08873638409478571 |issn=0887-3631}}' ]
All external links added in the edit (added_links)
[ 0 => 'https://abelha.org.br/artigos/', 1 => 'https://abelha.org.br/fichas-catalograficas-das-especies-relevantes-para-a-meliponicultura-2/', 2 => 'https://sites.usp.br/jardimdabotanicausprp/lambe-olhos-leurotrigona-muelleri/', 3 => 'https://sites.usp.br/jardimdabotanicausprp/mirim-plebeia-droryana/', 4 => 'https://ecomercioagrario.com/en/people-and-bees-mayan-bees-in-the-madrid-codex/', 5 => 'https://news.mongabay.com/2019/01/house-of-the-royal-lady-bee-maya-revive-native-bees-and-ancient-beekeeping/', 6 => 'http://www.journals.cambridge.org/abstract_S1742758404000128', 7 => 'https://doi.org/10.1079%2FIJT20048', 8 => 'https://search.worldcat.org/issn/1742-7584', 9 => 'https://doi.org/10.1080%2F00218839.2005.11101157', 10 => 'https://www.tandfonline.com/doi/abs/10.1080/00218839.2005.11101157', 11 => 'http://link.springer.com/10.1051/apido/2009016', 12 => 'https://doi.org/10.1051%2Fapido%2F2009016', 13 => 'https://www.scielo.br/j/rbzool/a/r74hTMg94tR9PTJGr8cvJwH/?lang=pt', 14 => 'https://doi.org/10.1590%2FS0101-81752001000100003', 15 => 'https://search.worldcat.org/issn/0101-8175', 16 => 'https://www.jstor.org/stable/25005104?utm_source=chatgpt.com', 17 => 'https://search.worldcat.org/issn/0028-7199', 18 => 'http://www.tandfonline.com/doi/full/10.1080/0005772X.2005.11099651', 19 => 'https://search.worldcat.org/issn/0005-772X', 20 => 'https://books.google.com.br/books/about/Foraging_Behaviour_of_Equatorial_Afrotro.html?id=bMuezQEACAAJ&redir_esc=y' ]
All external links in the new text (all_links)
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Parsed HTML source of the new revision (new_html)
'<div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Bee tribe, reduced stingers, strong bites</div> <p class="mw-empty-elt"> </p> <table class="infobox biota" style="text-align: left; width: 200px; font-size: 100%"> <tbody><tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(235,235,210)">Stingless bees<br /><div style="font-size: 85%;">Temporal range: <span class="noprint"><span style="display:inline-block;"></span><span style="display:inline-block;">Maastrichtian-Present</span> <span style="display:inline-block;"></span><div id="Timeline-row" style="margin: 4px auto 0; clear:both; width:220px; padding:0px; height:18px; overflow:visible; white-space:nowrap; border:1px #666; border-style:solid none; position:relative; z-index:0; font-size:97%;"> <div style="position:absolute; height:100%; left:0px; width:207.23076923077px; padding-left:5px; text-align:left; background-color:rgb(254,217,106); background-image: linear-gradient(to right, rgba(255,255,255,1), rgba(254,217,106,1) 15%, rgba(254,217,106,1));"><a href="/enwiki/wiki/Precambrian" title="Precambrian">PreꞒ</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(127,160,86); left:37.636923076923px; width:18.073846153846px;"><a href="/enwiki/wiki/Cambrian" title="Cambrian">Ꞓ</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(0,146,112); left:55.710769230769px; width:14.08px;"><a href="/enwiki/wiki/Ordovician" title="Ordovician">O</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(179,225,182); left:69.790769230769px; width:8.3261538461539px;"><a href="/enwiki/wiki/Silurian" title="Silurian">S</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(203,140,55); left:78.116923076923px; width:20.409230769231px;"><a href="/enwiki/wiki/Devonian" title="Devonian">D</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(103,165,153); left:98.526153846154px; width:20.307692307692px;"><a href="/enwiki/wiki/Carboniferous" title="Carboniferous">C</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(240,64,40); left:118.83384615385px; width:15.907015384615px;"><a href="/enwiki/wiki/Permian" title="Permian">P</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(129,43,146); left:134.74086153846px; width:17.092984615385px;"><a href="/enwiki/wiki/Triassic" title="Triassic">T</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(52,178,201); left:151.83384615385px; width:19.089230769231px;"><a href="/enwiki/wiki/Jurassic" title="Jurassic">J</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(127,198,78); left:170.92307692308px; width:26.738461538462px;"><a href="/enwiki/wiki/Cretaceous" title="Cretaceous">K</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(253,154,82); left:197.66153846154px; width:14.543692307692px;"><a href="/enwiki/wiki/Paleogene" title="Paleogene">Pg</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(255,230,25); left:212.20523076923px; width:6.9215384615385px;"><a href="/enwiki/wiki/Neogene" title="Neogene">N</a></div> <div id="end-border" style="position:absolute; height:100%; background-color:#666; width:1px; left:219px"></div><div style="margin:0 auto; line-height:0; clear:both; width:220px; padding:0px; height:8px; overflow:visible; background-color:transparent; position:relative; top:-4px; z-index:100;"><div style="position:absolute; height:8px; left:196.10461538462px; width:23.895384615385px; background-color:#360; opacity:0.42;"></div> <div style="position:absolute; height:8px; left:196.10461538462px; width:23.895384615385px; background-color:#360; opacity:1;"></div> <div style="position:absolute; height:6px; top:1px; left:197.10461538462px; width:21.895384615385px; background-color:#6c3;"></div> </div> </div></span></div> </th></tr> <tr> <td colspan="2" style="text-align: center"><span class="mw-default-size" typeof="mw:File/Frameless"><a href="/enwiki/wiki/File:Meliponula_ferruginea.jpg" class="mw-file-description"><img src="/upwiki/wikipedia/commons/thumb/1/12/Meliponula_ferruginea.jpg/220px-Meliponula_ferruginea.jpg" decoding="async" width="220" height="330" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/1/12/Meliponula_ferruginea.jpg/330px-Meliponula_ferruginea.jpg 1.5x, /upwiki/wikipedia/commons/thumb/1/12/Meliponula_ferruginea.jpg/440px-Meliponula_ferruginea.jpg 2x" data-file-width="1026" data-file-height="1540" /></a></span> </td></tr> <tr> <td colspan="2" style="text-align: center; font-size: 88%"><i>Meliponula ferruginea</i> </td></tr> <tr> <th colspan="2" style="color:inherit; min-width:15em; text-align: center; background-color: rgb(235,235,210)"><a href="/enwiki/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">Scientific classification</a> <span class="plainlinks taxobox-edit-taxonomy skin-invert" style="font-size:smaller; float:right; padding-right:0.4em; margin-left:-3em;"><span typeof="mw:File"><a href="/enwiki/wiki/Template:Taxonomy/Meliponini" title="Edit this classification"><img alt="Edit this classification" src="/upwiki/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/15px-OOjs_UI_icon_edit-ltr.svg.png" decoding="async" width="15" height="15" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/23px-OOjs_UI_icon_edit-ltr.svg.png 1.5x, /upwiki/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/30px-OOjs_UI_icon_edit-ltr.svg.png 2x" data-file-width="20" data-file-height="20" /></a></span></span> </th></tr> <tr> <td>Domain: </td> <td><a href="/enwiki/wiki/Eukaryote" title="Eukaryote">Eukaryota</a> </td></tr> <tr> <td>Kingdom: </td> <td><a href="/enwiki/wiki/Animal" title="Animal">Animalia</a> </td></tr> <tr> <td>Phylum: </td> <td><a href="/enwiki/wiki/Arthropod" title="Arthropod">Arthropoda</a> </td></tr> <tr> <td>Class: </td> <td><a href="/enwiki/wiki/Insect" title="Insect">Insecta</a> </td></tr> <tr> <td>Order: </td> <td><a href="/enwiki/wiki/Hymenoptera" title="Hymenoptera">Hymenoptera</a> </td></tr> <tr> <td>Family: </td> <td><a href="/enwiki/wiki/Apidae" title="Apidae">Apidae</a> </td></tr> <tr> <td><i>Clade</i>: </td> <td><a href="/enwiki/wiki/Corbiculata" class="mw-redirect" title="Corbiculata">Corbiculata</a> </td></tr> <tr> <td>Tribe: </td> <td><a class="mw-selflink selflink">Meliponini</a><br /><small><a href="/enwiki/wiki/Am%C3%A9d%C3%A9e_Louis_Michel_le_Peletier,_comte_de_Saint-Fargeau" title="Amédée Louis Michel le Peletier, comte de Saint-Fargeau">Lepeletier</a>, 1836</small> </td></tr> <tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(235,235,210)">Genera </th></tr> <tr> <td colspan="2" style="text-align: left"> <style data-mw-deduplicate="TemplateStyles:r1126788409">.mw-parser-output .plainlist ol,.mw-parser-output .plainlist ul{line-height:inherit;list-style:none;margin:0;padding:0}.mw-parser-output .plainlist ol li,.mw-parser-output .plainlist ul li{margin-bottom:0}</style><div class="plainlist"><ul><li><i><a href="/enwiki/w/index.php?title=Aparatrigona&amp;action=edit&amp;redlink=1" class="new" title="Aparatrigona (page does not exist)">Aparatrigona</a></i></li><li><i><a href="/enwiki/w/index.php?title=Apotrigona&amp;action=edit&amp;redlink=1" class="new" title="Apotrigona (page does not exist)">Apotrigona</a></i></li><li><i><a href="/enwiki/w/index.php?title=Asperplebeia&amp;action=edit&amp;redlink=1" class="new" title="Asperplebeia (page does not exist)">Asperplebeia</a></i></li><li><i><a href="/enwiki/wiki/Austroplebeia" title="Austroplebeia">Austroplebeia</a></i></li><li><i><a href="/enwiki/w/index.php?title=Axestotrigona&amp;action=edit&amp;redlink=1" class="new" title="Axestotrigona (page does not exist)">Axestotrigona</a></i></li><li><i><a href="/enwiki/wiki/Camargoia" title="Camargoia">Camargoia</a></i></li><li><i><a href="/enwiki/wiki/Celetrigona" title="Celetrigona">Celetrigona</a></i></li><li><i><a href="/enwiki/wiki/Cephalotrigona" title="Cephalotrigona">Cephalotrigona</a></i></li><li><i><a href="/enwiki/w/index.php?title=Cleptotrigona&amp;action=edit&amp;redlink=1" class="new" title="Cleptotrigona (page does not exist)">Cleptotrigona</a></i></li><li><a href="/enwiki/wiki/Extinction" title="Extinction">†</a><i><a href="/enwiki/w/index.php?title=Cretotrigona&amp;action=edit&amp;redlink=1" class="new" title="Cretotrigona (page does not exist)">Cretotrigona</a></i></li><li><i><a href="/enwiki/wiki/Dactylurina" title="Dactylurina">Dactylurina</a></i></li><li><i><a href="/enwiki/wiki/Dolichotrigona" title="Dolichotrigona">Dolichotrigona</a></i></li><li><i><a href="/enwiki/wiki/Duckeola" title="Duckeola">Duckeola</a></i></li><li><i><a href="/enwiki/w/index.php?title=Ebaiotrigona&amp;action=edit&amp;redlink=1" class="new" title="Ebaiotrigona (page does not exist)">Ebaiotrigona</a></i></li><li>†<i><a href="/enwiki/w/index.php?title=Exebotrigona&amp;action=edit&amp;redlink=1" class="new" title="Exebotrigona (page does not exist)">Exebotrigona</a></i></li><li><i><a href="/enwiki/wiki/Friesella" title="Friesella">Friesella</a></i></li><li><i><a href="/enwiki/wiki/Frieseomelitta" title="Frieseomelitta">Frieseomelitta</a></i></li><li><i><a href="/enwiki/w/index.php?title=Geniotrigona&amp;action=edit&amp;redlink=1" class="new" title="Geniotrigona (page does not exist)">Geniotrigona</a></i></li><li><i><a href="/enwiki/wiki/Geotrigona" title="Geotrigona">Geotrigona</a></i></li><li><i><a href="/enwiki/w/index.php?title=Heterotrigona&amp;action=edit&amp;redlink=1" class="new" title="Heterotrigona (page does not exist)">Heterotrigona</a></i></li><li><i><a href="/enwiki/w/index.php?title=Homotrigona&amp;action=edit&amp;redlink=1" class="new" title="Homotrigona (page does not exist)">Homotrigona</a></i></li><li><i><a href="/enwiki/w/index.php?title=Hypotrigona&amp;action=edit&amp;redlink=1" class="new" title="Hypotrigona (page does not exist)">Hypotrigona</a></i></li><li>†<i><a href="/enwiki/w/index.php?title=Kelneriapis&amp;action=edit&amp;redlink=1" class="new" title="Kelneriapis (page does not exist)">Kelneriapis</a></i></li><li><i><a href="/enwiki/w/index.php?title=Lepidotrigona&amp;action=edit&amp;redlink=1" class="new" title="Lepidotrigona (page does not exist)">Lepidotrigona</a></i></li><li><i><a href="/enwiki/wiki/Lestrimelitta" title="Lestrimelitta">Lestrimelitta</a></i></li><li><i><a href="/enwiki/wiki/Leurotrigona" title="Leurotrigona">Leurotrigona</a></i></li><li><i><a href="/enwiki/w/index.php?title=Liotrigona&amp;action=edit&amp;redlink=1" class="new" title="Liotrigona (page does not exist)">Liotrigona</a></i></li><li>†<i><a href="/enwiki/w/index.php?title=Liotrigonopsis&amp;action=edit&amp;redlink=1" class="new" title="Liotrigonopsis (page does not exist)">Liotrigonopsis</a></i></li><li><i><a href="/enwiki/w/index.php?title=Lisotrigona&amp;action=edit&amp;redlink=1" class="new" title="Lisotrigona (page does not exist)">Lisotrigona</a></i></li><li><i><a href="/enwiki/w/index.php?title=Lophotrigona&amp;action=edit&amp;redlink=1" class="new" title="Lophotrigona (page does not exist)">Lophotrigona</a></i></li><li><i><a href="/enwiki/w/index.php?title=Meliplebeia&amp;action=edit&amp;redlink=1" class="new" title="Meliplebeia (page does not exist)">Meliplebeia</a></i></li><li><i><a href="/enwiki/wiki/Melipona" title="Melipona">Melipona</a></i></li><li>†<i><a href="/enwiki/w/index.php?title=Meliponorytes&amp;action=edit&amp;redlink=1" class="new" title="Meliponorytes (page does not exist)">Meliponorytes</a></i></li><li><i><a href="/enwiki/wiki/Meliponula" title="Meliponula">Meliponula</a></i></li><li><i><a href="/enwiki/w/index.php?title=Meliwillea&amp;action=edit&amp;redlink=1" class="new" title="Meliwillea (page does not exist)">Meliwillea</a></i></li><li><i><a href="/enwiki/w/index.php?title=Mourella&amp;action=edit&amp;redlink=1" class="new" title="Mourella (page does not exist)">Mourella</a></i></li><li><i><a href="/enwiki/wiki/Nannotrigona" title="Nannotrigona">Nannotrigona</a></i></li><li><i><a href="/enwiki/w/index.php?title=Nanoplebeia&amp;action=edit&amp;redlink=1" class="new" title="Nanoplebeia (page does not exist)">Nanoplebeia</a></i></li><li><i><a href="/enwiki/wiki/Nogueirapis" title="Nogueirapis">Nogueirapis</a></i></li><li><i><a href="/enwiki/w/index.php?title=Odontotrigona&amp;action=edit&amp;redlink=1" class="new" title="Odontotrigona (page does not exist)">Odontotrigona</a></i></li><li><i><a href="/enwiki/wiki/Oxytrigona" title="Oxytrigona">Oxytrigona</a></i></li><li><i><a href="/enwiki/w/index.php?title=Papuatrigona&amp;action=edit&amp;redlink=1" class="new" title="Papuatrigona (page does not exist)">Papuatrigona</a></i></li><li><i><a href="/enwiki/w/index.php?title=Parapartamona&amp;action=edit&amp;redlink=1" class="new" title="Parapartamona (page does not exist)">Parapartamona</a></i></li><li><i><a href="/enwiki/wiki/Paratrigona" title="Paratrigona">Paratrigona</a></i></li><li><i><a href="/enwiki/w/index.php?title=Pariotrigona&amp;action=edit&amp;redlink=1" class="new" title="Pariotrigona (page does not exist)">Pariotrigona</a></i></li><li><i><a href="/enwiki/wiki/Paratrigonoides" title="Paratrigonoides">Paratrigonoides</a></i></li><li><i><a href="/enwiki/wiki/Partamona" title="Partamona">Partamona</a></i></li><li><i><a href="/enwiki/w/index.php?title=Platytrigona&amp;action=edit&amp;redlink=1" class="new" title="Platytrigona (page does not exist)">Platytrigona</a></i></li><li><i><a href="/enwiki/wiki/Plebeia" title="Plebeia">Plebeia</a></i></li><li><i><a href="/enwiki/w/index.php?title=Plebeiella&amp;action=edit&amp;redlink=1" class="new" title="Plebeiella (page does not exist)">Plebeiella</a></i></li><li><i><a href="/enwiki/wiki/Plebeina" class="mw-redirect" title="Plebeina">Plebeina</a></i></li><li><i><a href="/enwiki/w/index.php?title=Plectoplebeia&amp;action=edit&amp;redlink=1" class="new" title="Plectoplebeia (page does not exist)">Plectoplebeia</a></i></li><li>†<i><a href="/enwiki/w/index.php?title=Proplebeia&amp;action=edit&amp;redlink=1" class="new" title="Proplebeia (page does not exist)">Proplebeia</a></i></li><li><i><a href="/enwiki/wiki/Ptilotrigona" title="Ptilotrigona">Ptilotrigona</a></i></li><li><i><a href="/enwiki/w/index.php?title=Sahulotrigona&amp;action=edit&amp;redlink=1" class="new" title="Sahulotrigona (page does not exist)">Sahulotrigona</a></i></li><li><i><a href="/enwiki/wiki/Scaptotrigona" title="Scaptotrigona">Scaptotrigona</a></i></li><li><i><a href="/enwiki/wiki/Scaura" title="Scaura">Scaura</a></i></li><li><i><a href="/enwiki/wiki/Schwarziana" title="Schwarziana">Schwarziana</a></i></li><li><i><a href="/enwiki/w/index.php?title=Sundatrigona&amp;action=edit&amp;redlink=1" class="new" title="Sundatrigona (page does not exist)">Sundatrigona</a></i></li><li><i><a href="/enwiki/wiki/Tetragona" title="Tetragona">Tetragona</a></i></li><li><i><a href="/enwiki/w/index.php?title=Tetragonilla&amp;action=edit&amp;redlink=1" class="new" title="Tetragonilla (page does not exist)">Tetragonilla</a></i></li><li><i><a href="/enwiki/wiki/Tetragonisca" title="Tetragonisca">Tetragonisca</a></i></li><li><i><a href="/enwiki/wiki/Tetragonula" title="Tetragonula">Tetragonula</a></i></li><li><i><a href="/enwiki/w/index.php?title=Tetrigona&amp;action=edit&amp;redlink=1" class="new" title="Tetrigona (page does not exist)">Tetrigona</a></i></li><li><i><a href="/enwiki/wiki/Trichotrigona" title="Trichotrigona">Trichotrigona</a></i></li><li><i><a href="/enwiki/wiki/Trigona_(genus)" class="mw-redirect" title="Trigona (genus)">Trigona</a></i></li><li><i><a href="/enwiki/wiki/Trigonisca" title="Trigonisca">Trigonisca</a></i></li><li><i><a href="/enwiki/w/index.php?title=Wallacetrigona&amp;action=edit&amp;redlink=1" class="new" title="Wallacetrigona (page does not exist)">Wallacetrigona</a></i></li></ul></div> </td></tr> </tbody></table><style data-mw-deduplicate="TemplateStyles:r1238732961">@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox.biota tr{background:transparent!important}html.skin-theme-clientpref-night .mw-parser-output .infobox.biota img{background:transparent}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox.biota tr{background:transparent!important}html.skin-theme-clientpref-os .mw-parser-output .infobox.biota img{background:white}}.mw-parser-output .infobox.biota .taxobox-edit-taxonomy img{background:transparent!important}body.skin-vector .mw-parser-output table.biota.infobox{margin-top:0.5em}body.skin--responsive .mw-parser-output table.biota.infobox tr.taxonrow td{padding:2px 10px}</style> <p><b>Stingless bees (SB)</b>, sometimes called <b>stingless honey bees</b> or simply <b>meliponines</b>, are a large group of bees (from about 462 to 552 described species),<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">&#91;</span>1<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:27_2-0" class="reference"><a href="#cite_note-:27-2"><span class="cite-bracket">&#91;</span>2<span class="cite-bracket">&#93;</span></a></sup> comprising the <a href="/enwiki/wiki/Tribe_(biology)" title="Tribe (biology)">tribe</a> <b>Meliponini</b><sup id="cite_ref-3" class="reference"><a href="#cite_note-3"><span class="cite-bracket">&#91;</span>3<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">&#91;</span>4<span class="cite-bracket">&#93;</span></a></sup> (or <a href="/enwiki/wiki/Subtribe" title="Subtribe">subtribe</a> Meliponina according to other authors).<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">&#91;</span>5<span class="cite-bracket">&#93;</span></a></sup> They belong in the <a href="/enwiki/wiki/Family_(biology)" title="Family (biology)">family</a> <a href="/enwiki/wiki/Apidae" title="Apidae">Apidae</a> (<a href="/enwiki/wiki/Subfamily" title="Subfamily">subfamily</a> <a href="/enwiki/wiki/Apinae" title="Apinae">Apinae</a>), and are closely related to common <a href="/enwiki/wiki/Honey_bee" title="Honey bee">honey bees</a> (HB, tribe <a href="/enwiki/wiki/Apini" class="mw-redirect" title="Apini">Apini</a>), orchid bees (tribe <a href="/enwiki/wiki/Euglossini" title="Euglossini">Euglossini</a>), and <a href="/enwiki/wiki/Bumblebee" title="Bumblebee">bumblebees</a> (tribe <a href="/enwiki/wiki/Bombini" title="Bombini">Bombini</a>). These four bee tribes belong to the <a href="/enwiki/wiki/Pollen_basket" title="Pollen basket">corbiculate</a> bees <a href="/enwiki/wiki/Monophyly" title="Monophyly">monophyletic</a> group.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">&#91;</span>6<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">&#91;</span>7<span class="cite-bracket">&#93;</span></a></sup> Meliponines have stingers, but they are highly reduced and cannot be used for defense, though these bees exhibit other defensive behaviors and mechanisms.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">&#91;</span>8<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:1_9-0" class="reference"><a href="#cite_note-:1-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup> Meliponines are not the only type of bee incapable of stinging: all male bees and many female bees of several other families, such as <a href="/enwiki/wiki/Andrenidae" title="Andrenidae">Andrenidae</a> and <a href="/enwiki/wiki/Megachilidae" title="Megachilidae">Megachilidae</a> (tribe <a href="/enwiki/wiki/Dioxyini" title="Dioxyini">Dioxyini</a>), also cannot sting.<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">&#91;</span>10<span class="cite-bracket">&#93;</span></a></sup> </p><p>Some stingless bees have powerful mandibles and can inflict painful bites.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">&#91;</span>11<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">&#91;</span>12<span class="cite-bracket">&#93;</span></a></sup> Some species can present large mandibular glands for the secretion of caustic defense substances, secrete unpleasant smells or use sticky materials to immobilise enemies.<sup id="cite_ref-:3_13-0" class="reference"><a href="#cite_note-:3-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">&#91;</span>14<span class="cite-bracket">&#93;</span></a></sup> </p><p>The main honey-producing bees of this group generally belong to the genera <i><a href="/enwiki/wiki/Scaptotrigona" title="Scaptotrigona">Scaptotrigona</a></i>, <i><a href="/enwiki/wiki/Tetragonisca" title="Tetragonisca">Tetragonisca</a></i>, <i><a href="/enwiki/wiki/Melipona" title="Melipona">Melipona</a></i> and <i><a href="/enwiki/wiki/Austroplebeia" title="Austroplebeia">Austroplebeia</a></i>, although there are other genera containing species that produce some usable honey. They are farmed in <a href="/enwiki/wiki/Meliponiculture" title="Meliponiculture">meliponiculture</a> in the same way that European honey bees (genus <i><a href="/enwiki/wiki/Apis_(genus)" class="mw-redirect" title="Apis (genus)">Apis</a></i>) are cultivated in <a href="/enwiki/wiki/Apiculture" class="mw-redirect" title="Apiculture">apiculture</a>. </p><p>Throughout <a href="/enwiki/wiki/Mesoamerica" title="Mesoamerica">Mesoamerica</a>, the <a href="/enwiki/wiki/Maya_peoples" title="Maya peoples">Mayans</a> have engaged in extensive meliponiculture on a large scale since before the arrival of Columbus. Meliponiculture played a significant role in <a href="/enwiki/wiki/Maya_society" title="Maya society">Maya society</a>, influencing their social, economic, and religious activities. The practice of maintaining <b>stingless bees</b> in man-made structures is prevalent across the Americas, with notable instances in countries such as <a href="/enwiki/wiki/Brazil" title="Brazil">Brazil</a>, <a href="/enwiki/wiki/Peru" title="Peru">Peru</a>, and <a href="/enwiki/wiki/Mexico" title="Mexico">Mexico</a>.<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">&#91;</span>15<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">&#91;</span>16<span class="cite-bracket">&#93;</span></a></sup> </p> <div id="toc" class="toc" role="navigation" aria-labelledby="mw-toc-heading"><input type="checkbox" role="button" id="toctogglecheckbox" class="toctogglecheckbox" style="display:none" /><div class="toctitle" lang="en" dir="ltr"><h2 id="mw-toc-heading">Contents</h2><span class="toctogglespan"><label class="toctogglelabel" for="toctogglecheckbox"></label></span></div> <ul> <li class="toclevel-1 tocsection-1"><a href="#Geographical_distribution"><span class="tocnumber">1</span> <span class="toctext">Geographical distribution</span></a> <ul> <li class="toclevel-2 tocsection-2"><a href="#Origin_and_dispersion"><span class="tocnumber">1.1</span> <span class="toctext">Origin and dispersion</span></a></li> </ul> </li> <li class="toclevel-1 tocsection-3"><a href="#Evolution_and_phylogeny"><span class="tocnumber">2</span> <span class="toctext">Evolution and phylogeny</span></a> <ul> <li class="toclevel-2 tocsection-4"><a href="#Fossil_history"><span class="tocnumber">2.1</span> <span class="toctext">Fossil history</span></a></li> </ul> </li> <li class="toclevel-1 tocsection-5"><a href="#Behaviour,_biology_and_ecology"><span class="tocnumber">3</span> <span class="toctext">Behaviour, biology and ecology</span></a> <ul> <li class="toclevel-2 tocsection-6"><a href="#Overview"><span class="tocnumber">3.1</span> <span class="toctext">Overview</span></a></li> <li class="toclevel-2 tocsection-7"><a href="#Castes"><span class="tocnumber">3.2</span> <span class="toctext">Castes</span></a> <ul> <li class="toclevel-3 tocsection-8"><a href="#Workers"><span class="tocnumber">3.2.1</span> <span class="toctext">Workers</span></a></li> <li class="toclevel-3 tocsection-9"><a href="#Queens"><span class="tocnumber">3.2.2</span> <span class="toctext">Queens</span></a></li> <li class="toclevel-3 tocsection-10"><a href="#Males_(drones)"><span class="tocnumber">3.2.3</span> <span class="toctext">Males (drones)</span></a></li> <li class="toclevel-3 tocsection-11"><a href="#Soldiers"><span class="tocnumber">3.2.4</span> <span class="toctext">Soldiers</span></a></li> <li class="toclevel-3 tocsection-12"><a href="#Division_of_labour"><span class="tocnumber">3.2.5</span> <span class="toctext">Division of labour</span></a></li> </ul> </li> <li class="toclevel-2 tocsection-13"><a href="#Products_and_materials"><span class="tocnumber">3.3</span> <span class="toctext">Products and materials</span></a></li> <li class="toclevel-2 tocsection-14"><a href="#Nest"><span class="tocnumber">3.4</span> <span class="toctext">Nest</span></a> <ul> <li class="toclevel-3 tocsection-15"><a href="#Exposed_nests"><span class="tocnumber">3.4.1</span> <span class="toctext">Exposed nests</span></a></li> <li class="toclevel-3 tocsection-16"><a href="#Ground_nests"><span class="tocnumber">3.4.2</span> <span class="toctext">Ground nests</span></a></li> <li class="toclevel-3 tocsection-17"><a href="#Termite_and_ant_shared_nests"><span class="tocnumber">3.4.3</span> <span class="toctext">Termite and ant shared nests</span></a></li> <li class="toclevel-3 tocsection-18"><a href="#Cavity_nests"><span class="tocnumber">3.4.4</span> <span class="toctext">Cavity nests</span></a></li> <li class="toclevel-3 tocsection-19"><a href="#Entrances"><span class="tocnumber">3.4.5</span> <span class="toctext">Entrances</span></a></li> <li class="toclevel-3 tocsection-20"><a href="#Brood_cell_arrangement"><span class="tocnumber">3.4.6</span> <span class="toctext">Brood cell arrangement</span></a></li> <li class="toclevel-3 tocsection-21"><a href="#Brood_rearing"><span class="tocnumber">3.4.7</span> <span class="toctext">Brood rearing</span></a></li> </ul> </li> <li class="toclevel-2 tocsection-22"><a href="#Swarming"><span class="tocnumber">3.5</span> <span class="toctext">Swarming</span></a></li> <li class="toclevel-2 tocsection-23"><a href="#Natural_enemies"><span class="tocnumber">3.6</span> <span class="toctext">Natural enemies</span></a> <ul> <li class="toclevel-3 tocsection-24"><a href="#Invertebrates"><span class="tocnumber">3.6.1</span> <span class="toctext">Invertebrates</span></a></li> <li class="toclevel-3 tocsection-25"><a href="#Vertebrates"><span class="tocnumber">3.6.2</span> <span class="toctext">Vertebrates</span></a></li> <li class="toclevel-3 tocsection-26"><a href="#Defense"><span class="tocnumber">3.6.3</span> <span class="toctext">Defense</span></a></li> </ul> </li> <li class="toclevel-2 tocsection-27"><a href="#Role_differentiation"><span class="tocnumber">3.7</span> <span class="toctext">Role differentiation</span></a></li> </ul> </li> <li class="toclevel-1 tocsection-28"><a href="#Interaction_with_humans"><span class="tocnumber">4</span> <span class="toctext">Interaction with humans</span></a> <ul> <li class="toclevel-2 tocsection-29"><a href="#Pollination"><span class="tocnumber">4.1</span> <span class="toctext">Pollination</span></a></li> </ul> </li> <li class="toclevel-1 tocsection-30"><a href="#Worldwide_overview"><span class="tocnumber">5</span> <span class="toctext">Worldwide overview</span></a> <ul> <li class="toclevel-2 tocsection-31"><a href="#Africa"><span class="tocnumber">5.1</span> <span class="toctext">Africa</span></a></li> <li class="toclevel-2 tocsection-32"><a href="#Australia"><span class="tocnumber">5.2</span> <span class="toctext">Australia</span></a> <ul> <li class="toclevel-3 tocsection-33"><a href="#Pollination_2"><span class="tocnumber">5.2.1</span> <span class="toctext">Pollination</span></a></li> </ul> </li> <li class="toclevel-2 tocsection-34"><a href="#Brazil"><span class="tocnumber">5.3</span> <span class="toctext">Brazil</span></a> <ul> <li class="toclevel-3 tocsection-35"><a href="#Honey_production"><span class="tocnumber">5.3.1</span> <span class="toctext">Honey production</span></a></li> <li class="toclevel-3 tocsection-36"><a href="#Bees_as_pets"><span class="tocnumber">5.3.2</span> <span class="toctext">Bees as pets</span></a></li> </ul> </li> <li class="toclevel-2 tocsection-37"><a href="#Central_America"><span class="tocnumber">5.4</span> <span class="toctext">Central America</span></a> <ul> <li class="toclevel-3 tocsection-38"><a href="#History"><span class="tocnumber">5.4.1</span> <span class="toctext">History</span></a></li> <li class="toclevel-3 tocsection-39"><a href="#Tulum"><span class="tocnumber">5.4.2</span> <span class="toctext">Tulum</span></a></li> <li class="toclevel-3 tocsection-40"><a href="#Economic_uses"><span class="tocnumber">5.4.3</span> <span class="toctext">Economic uses</span></a></li> <li class="toclevel-3 tocsection-41"><a href="#Future"><span class="tocnumber">5.4.4</span> <span class="toctext">Future</span></a></li> </ul> </li> </ul> </li> <li class="toclevel-1 tocsection-42"><a href="#References"><span class="tocnumber">6</span> <span class="toctext">References</span></a></li> <li class="toclevel-1 tocsection-43"><a href="#Bibliography"><span class="tocnumber">7</span> <span class="toctext">Bibliography</span></a> <ul> <li class="toclevel-2 tocsection-44"><a href="#Articles_and_publications"><span class="tocnumber">7.1</span> <span class="toctext">Articles and publications</span></a></li> <li class="toclevel-2 tocsection-45"><a href="#Books"><span class="tocnumber">7.2</span> <span class="toctext">Books</span></a></li> </ul> </li> <li class="toclevel-1 tocsection-46"><a href="#External_links"><span class="tocnumber">8</span> <span class="toctext">External links</span></a></li> </ul> </div> <div class="mw-heading mw-heading2"><h2 id="Geographical_distribution">Geographical distribution</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=1" title="Edit section&#039;s source code: Geographical distribution"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Stingless bees can be found in most tropical or subtropical regions of the world, such as the <a href="/enwiki/wiki/Africa" title="Africa">African continent</a> (<a href="/enwiki/wiki/Afrotropical_realm" title="Afrotropical realm">Afrotropical</a> region), <a href="/enwiki/wiki/Southeast_Asia" title="Southeast Asia">Southeast Asia</a> and <a href="/enwiki/wiki/Australia" title="Australia">Australia</a> (<a href="/enwiki/wiki/Indomalayan_realm" title="Indomalayan realm">Indo-Malayan</a> and <a href="/enwiki/wiki/Australasian_realm" title="Australasian realm">Australasian</a> region), and tropical <a href="/enwiki/wiki/Americas" title="Americas">America</a> (<a href="/enwiki/wiki/Neotropical_realm" title="Neotropical realm">Neotropical</a> region).<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">&#91;</span>17<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:0_18-0" class="reference"><a href="#cite_note-:0-18"><span class="cite-bracket">&#91;</span>18<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">&#91;</span>19<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">&#91;</span>20<span class="cite-bracket">&#93;</span></a></sup> </p><p>The majority of <a href="/enwiki/wiki/Indigenous_(ecology)" class="mw-redirect" title="Indigenous (ecology)">native</a> <a href="/enwiki/wiki/Eusocial" class="mw-redirect" title="Eusocial">eusocial</a> bees of <a href="/enwiki/wiki/Central_America" title="Central America">Central</a> and <a href="/enwiki/wiki/South_America" title="South America">South America</a> are SB, although only a few of them produce honey on a scale such that they are farmed by humans.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">&#91;</span>21<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">&#91;</span>22<span class="cite-bracket">&#93;</span></a></sup> The Neotropics, with approximately 426 species, boast the highest abundance and species richness, ranging from Cuba and Mexico in the north to Argentina in the south.<sup id="cite_ref-:0_18-1" class="reference"><a href="#cite_note-:0-18"><span class="cite-bracket">&#91;</span>18<span class="cite-bracket">&#93;</span></a></sup> </p><p>They are also quite diverse in <a href="/enwiki/wiki/Africa" title="Africa">Africa</a>, including <a href="/enwiki/wiki/Madagascar" title="Madagascar">Madagascar</a>,<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">&#91;</span>23<span class="cite-bracket">&#93;</span></a></sup> and are farmed there also. Around 36 species exist on the continent. The equatorial regions harbor the greatest diversity, with the <a href="/enwiki/wiki/Sahara" title="Sahara">Sahara Desert</a> acting as a natural barrier to the north. The range extends southward to <a href="/enwiki/wiki/South_Africa" title="South Africa">South Africa</a> and southern Madagascar, with most African species inhabiting <a href="/enwiki/wiki/Tropical_forest" title="Tropical forest">tropical forests</a> or both tropical forests and <a href="/enwiki/wiki/Savanna" title="Savanna">savannahs</a>.<sup id="cite_ref-:0_18-2" class="reference"><a href="#cite_note-:0-18"><span class="cite-bracket">&#91;</span>18<span class="cite-bracket">&#93;</span></a></sup> </p><p>Meliponine honey is prized as a medicine in many African communities, as well as in South America. Some cultures use SB honey against <a href="/enwiki/wiki/Human_digestive_system" title="Human digestive system">digestive</a>, <a href="/enwiki/wiki/Respiratory_system" title="Respiratory system">respiratory</a>, <a href="/enwiki/wiki/Eye" title="Eye">ocular</a> and <a href="/enwiki/wiki/Reproductive_system" title="Reproductive system">reproductive</a> problems, although more research is needed to disclose evidence that supports these practices.<sup id="cite_ref-:1_9-1" class="reference"><a href="#cite_note-:1-9"><span class="cite-bracket">&#91;</span>9<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">&#91;</span>24<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">&#91;</span>25<span class="cite-bracket">&#93;</span></a></sup> </p><p>In Asia and Australia, approximately 90 species of stingless bees span from <a href="/enwiki/wiki/India" title="India">India</a> in the west to the <a href="/enwiki/wiki/Solomon_Islands" title="Solomon Islands">Solomon Islands</a> in the east, and from <a href="/enwiki/wiki/Nepal" title="Nepal">Nepal</a>, China (<a href="/enwiki/wiki/Yunnan" title="Yunnan">Yunnan</a>, <a href="/enwiki/wiki/Hainan" title="Hainan">Hainan</a>), and <a href="/enwiki/wiki/Taiwan" title="Taiwan">Taiwan</a> in the north to Australia in the south. </p> <figure class="mw-halign-center" typeof="mw:File/Thumb"><a href="/enwiki/wiki/File:Geographic_distribution_of_stingless_bees.png" class="mw-file-description"><img src="/upwiki/wikipedia/commons/thumb/3/34/Geographic_distribution_of_stingless_bees.png/399px-Geographic_distribution_of_stingless_bees.png" decoding="async" width="399" height="196" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/3/34/Geographic_distribution_of_stingless_bees.png/599px-Geographic_distribution_of_stingless_bees.png 1.5x, /upwiki/wikipedia/commons/3/34/Geographic_distribution_of_stingless_bees.png 2x" data-file-width="743" data-file-height="365" /></a><figcaption>Geographic distribution of stingless bees.</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Origin_and_dispersion">Origin and dispersion</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=2" title="Edit section&#039;s source code: Origin and dispersion"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Phylogenetic analyses reveal three distinct groups in the evolutionary history of <b>Meliponini</b>: the Afrotropical, the Indo-Malay/Australasia, and the Neotropical lineages. The evolutionary origin of the Meliponini is Neotropical. Studies observing contemporary <a href="/enwiki/wiki/Species_richness" title="Species richness">species richness</a> show that it remains highest in the Neotropics.<sup id="cite_ref-:2_26-0" class="reference"><a href="#cite_note-:2-26"><span class="cite-bracket">&#91;</span>26<span class="cite-bracket">&#93;</span></a></sup> </p><p>The hypothesis proposes the potential dispersion of stingless bees from what is now <a href="/enwiki/wiki/North_America" title="North America">North America</a>. According to this scenario, these bees would have then traveled to Asia by crossing the <a href="/enwiki/wiki/Bering_Strait" title="Bering Strait">Bering Strait</a> (<a href="/enwiki/wiki/Beringia" title="Beringia">Beringia</a> route) and reached <a href="/enwiki/wiki/Europe" title="Europe">Europe</a> through <a href="/enwiki/wiki/Greenland" title="Greenland">Greenland</a> (<a href="/enwiki/wiki/Thule_Land_Bridge" title="Thule Land Bridge">Thulean route</a>).<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">&#91;</span>27<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:2_26-1" class="reference"><a href="#cite_note-:2-26"><span class="cite-bracket">&#91;</span>26<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">&#91;</span>28<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">&#91;</span>29<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Evolution_and_phylogeny">Evolution and phylogeny</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=3" title="Edit section&#039;s source code: Evolution and phylogeny"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Meliponines form a <a href="/enwiki/wiki/Clade" title="Clade">clade</a> within the corbiculate bees, characterized by unique pollen-carrying structures known as corbiculae (<a href="/enwiki/wiki/Pollen_basket" title="Pollen basket">pollen baskets</a>) located on their hind legs. This group also includes other three tribes: honey bees (<a href="/enwiki/wiki/Apini" class="mw-redirect" title="Apini">Apini</a>), bumble bees (<a href="/enwiki/wiki/Bombini" title="Bombini">Bombini</a>), and orchid bees (<a href="/enwiki/wiki/Euglossini" title="Euglossini">Euglossini</a>). The concept of higher <a href="/enwiki/wiki/Eusociality" title="Eusociality">eusociality</a>, defined by the presence of distinct queen and worker castes and characterized by features such as perennial colony lifestyles and extensive food sharing among adults, is particularly relevant in understanding the social structure of these tribes. Both Meliponini and Apini tribes are considered higher eusocial, while Bombini is considered to be primitively eusocial.<sup id="cite_ref-:29_30-0" class="reference"><a href="#cite_note-:29-30"><span class="cite-bracket">&#91;</span>30<span class="cite-bracket">&#93;</span></a></sup> </p><p>The phylogenetic relationships among the four tribes of corbiculate bees have been a topic of considerable debate within the scientific community. Two primary questions arise: the relationship of stingless bees to honey bees and bumble bees, and whether their eusocial behavior evolved independently or from a <a href="/enwiki/wiki/Common_descent" title="Common descent">common ancestor</a>. Morphological and behavioral studies have suggested that Meliponini and Apini are sister groups, indicating a single origin of higher eusociality. In contrast, molecular studies often support a relationship between Meliponini and Bombini, proposing independent origins of higher eusociality in both Apini and Meliponini.<sup id="cite_ref-:29_30-1" class="reference"><a href="#cite_note-:29-30"><span class="cite-bracket">&#91;</span>30<span class="cite-bracket">&#93;</span></a></sup> </p><p>A morphological, behavioral, and molecular data analysis provided strong support for the latter hypothesis of dual origins of higher eusociality. Subsequent research has reinforced the idea that stingless bees and honey bees evolved their eusocial lifestyles independently, resulting in distinct adaptive strategies for colony reproduction, brood rearing, foraging communication, and colony defense. This divergence helps explain the varied ecological and social solutions developed by these two groups of bees, such as <a href="/enwiki/wiki/Foraging" title="Foraging">foraging</a> communication, colony defense/reproduction and brood rearing.<sup id="cite_ref-:29_30-2" class="reference"><a href="#cite_note-:29-30"><span class="cite-bracket">&#91;</span>30<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">&#91;</span>31<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="clade"><style data-mw-deduplicate="TemplateStyles:r1261294616">body.skin-vector-2022 .mw-parser-output div.clade,body.skin-minerva .mw-parser-output div.clade{overflow-x:auto;overflow-y:hidden}body.skin-minerva .mw-parser-output div.clade p{font-size:inherit}.mw-parser-output table.clade{border-spacing:0;margin:0;font-size:100%;line-height:100%;border-collapse:separate;width:auto;display:table}.mw-parser-output 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td.clade-slabel.last{border-left:none;border-right:none}.mw-parser-output table.clade td.clade-slabel.reverse{border-left:none;border-right:1px solid}.mw-parser-output table.clade td.clade-bar{vertical-align:middle;text-align:left;padding:0 0.5em;position:relative}.mw-parser-output table.clade td.clade-bar.reverse{text-align:right;position:relative}.mw-parser-output table.clade td.clade-leaf{border:0;padding:0;text-align:left}.mw-parser-output table.clade td.clade-leaf p{padding-right:5px;padding-left:2px}.mw-parser-output table.clade td.clade-leafR{border:0;padding:0;text-align:right}.mw-parser-output table.clade td.clade-leafR p{padding-left:5px;padding-right:2px}.mw-parser-output table.clade td.clade-leaf.reverse{text-align:right}.mw-parser-output table.clade td.clade-leaf.reverse p{padding-left:5px;padding-right:2px}.mw-parser-output table.clade:hover span.linkA{background-color:yellow}.mw-parser-output table.clade:hover span.linkB{background-color:green}</style> <table class="clade" style="font-size:85%;line-height:85%"> <tbody><tr> <td class="clade-label first"><a href="/enwiki/wiki/Anthophila" class="mw-redirect" title="Anthophila">Anthophila</a>&#160;(bees) </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1261294616"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1261294616"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p><a href="/enwiki/wiki/Melittidae" title="Melittidae">Melittidae</a> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1261294616"> <table class="clade"> <tbody><tr> <td class="clade-label first"><div style="display:inline" class="nowrap"><span class="nowrap">long-tongued bees</span></div> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1261294616"> <table class="clade"> <tbody><tr> <td class="clade-label first">&#160;<a href="/enwiki/wiki/Apidae" title="Apidae">Apidae</a>&#160; </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1261294616"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p><a href="/enwiki/wiki/Euglossini" title="Euglossini">Euglossini</a> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"><div style="display:inline" class="nowrap"><span class="nowrap">eusocial</span></div> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1261294616"> <table class="clade"> <tbody><tr> <td class="clade-label first"><div style="display:inline" class="nowrap"><span class="nowrap">h. eusocial</span></div> </td> <td rowspan="2" class="clade-leaf"> <p><a href="/enwiki/wiki/Apini" class="mw-redirect" title="Apini">Apini</a> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1261294616"> <table class="clade"> <tbody><tr> <td class="clade-label first"><div style="display:inline" class="nowrap"><span class="nowrap">p. eusocial</span></div> </td> <td rowspan="2" class="clade-leaf"> <p><a href="/enwiki/wiki/Bombini" title="Bombini">Bombini</a> </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label" style="background-color:#f9a6e8"><div style="display:inline" class="nowrap"><span class="nowrap">h. eusocial</span></div> </td> <td rowspan="2" class="clade-leaf" style="background-color:#f9a6e8"> <p><a href="/enwiki/wiki/Meliponini" class="mw-redirect" title="Meliponini">Meliponini</a> </p> </td></tr> <tr> <td class="clade-slabel last" style="background-color:#f9a6e8"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <p><a href="/enwiki/wiki/Megachilidae" title="Megachilidae">Megachilidae</a> (mason, leafcutter bees) </p> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"><div style="display:inline" class="nowrap"><span class="nowrap">short-tongued bees</span></div> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1261294616"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p><a href="/enwiki/wiki/Andrenidae" title="Andrenidae">Andrenidae</a> (mining bees) </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1261294616"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p><a href="/enwiki/wiki/Halictidae" title="Halictidae">Halictidae</a> (sweat bees) </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1261294616"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <p><a href="/enwiki/wiki/Stenotritidae" title="Stenotritidae">Stenotritidae</a> (large Australian bees) </p> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <p><a href="/enwiki/wiki/Colletidae" title="Colletidae">Colletidae</a> (plasterer bees) </p> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> <div class="mw-heading mw-heading3"><h3 id="Fossil_history">Fossil history</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=4" title="Edit section&#039;s source code: Fossil history"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The <a href="/enwiki/wiki/Fossil" title="Fossil">fossil</a> record for stingless bees is notably robust compared to that of many other bee groups, with twelve extinct species currently identified. Fossils of these bees are primarily found in <a href="/enwiki/wiki/Amber" title="Amber">amber</a> and <a href="/enwiki/wiki/Copal" title="Copal">copal</a>, where excellent preservation typically occurs. This favorable fossilization process may be attributed to the behaviors of stingless bees, which collect significant amounts of tree <a href="/enwiki/wiki/Resin" title="Resin">resin</a> for building nests and defense, increasing the likelihood of entrapment.<sup id="cite_ref-:24_32-0" class="reference"><a href="#cite_note-:24-32"><span class="cite-bracket">&#91;</span>32<span class="cite-bracket">&#93;</span></a></sup> </p><p>Despite this relatively good fossil record, the evolutionary history of stingless bees remains poorly understood, particularly regarding their widespread distribution across various ecological niches around the globe. The oldest known fossil stingless bee is <i><a href="/enwiki/w/index.php?title=Cretotrigona_prisca&amp;action=edit&amp;redlink=1" class="new" title="Cretotrigona prisca (page does not exist)">Cretotrigona prisca</a></i>, a small worker bee approximately 5 mm in body length, discovered in New Jersey amber. This species is believed to have existed during the <a href="/enwiki/wiki/Late_Cretaceous" title="Late Cretaceous">Late Cretaceous</a> period, around 65–70 million years ago, marking it as the oldest confirmed fossil of an apid bee and the earliest fossil evidence of a eusocial bee. <i>C. prisca</i> exhibits striking similarities to extant stingless bees, indicating that the evolutionary lineage of meliponines may date back to this period.<sup id="cite_ref-:24_32-1" class="reference"><a href="#cite_note-:24-32"><span class="cite-bracket">&#91;</span>32<span class="cite-bracket">&#93;</span></a></sup> </p><p>Some researchers suggest that stingless bees likely evolved in the Late Cretaceous, approximately 70–87 million years ago.<sup id="cite_ref-:24_32-2" class="reference"><a href="#cite_note-:24-32"><span class="cite-bracket">&#91;</span>32<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">&#91;</span>33<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">&#91;</span>34<span class="cite-bracket">&#93;</span></a></sup> According to recent studies, corbiculate bees, which include stingless bees, are thought to have appeared around 84–87 million years ago, further supporting the notion of their evolution during this dynamic period in Earth's history.<sup id="cite_ref-:24_32-3" class="reference"><a href="#cite_note-:24-32"><span class="cite-bracket">&#91;</span>32<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">&#91;</span>35<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">&#91;</span>36<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Behaviour,_biology_and_ecology"><span id="Behaviour.2C_biology_and_ecology"></span>Behaviour, biology and ecology</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=5" title="Edit section&#039;s source code: Behaviour, biology and ecology"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Overview">Overview</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=6" title="Edit section&#039;s source code: Overview"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Meliponines, considered highly eusocial insects, exhibit a remarkable caste division. The colonies typically consist of a <a href="/enwiki/wiki/Queen_bee" title="Queen bee">queen</a>, <a href="/enwiki/wiki/Worker_bee" title="Worker bee">workers</a>, and sometimes male <a href="/enwiki/wiki/Drone_(bee)" title="Drone (bee)">drones</a>.<sup id="cite_ref-:4_37-0" class="reference"><a href="#cite_note-:4-37"><span class="cite-bracket">&#91;</span>37<span class="cite-bracket">&#93;</span></a></sup> The queen is responsible for reproduction, while the workers perform various tasks such as foraging, nursing, and defending the colony. Individuals work together with a well-defined division of labor for the overall benefit.<sup id="cite_ref-:5_38-0" class="reference"><a href="#cite_note-:5-38"><span class="cite-bracket">&#91;</span>38<span class="cite-bracket">&#93;</span></a></sup> </p><p>Stingless bees are valuable pollinators and contribute to ecosystem health by producing essential products. These insects collect and store honey, pollen, resin, propolis, and cerumen. Honey serves as their primary <a href="/enwiki/wiki/Carbohydrate" title="Carbohydrate">carbohydrate</a> source, while pollen provides essential <a href="/enwiki/wiki/Protein" title="Protein">proteins</a>. Resin, propolis, and cerumen are used in nest construction and maintenance.<sup id="cite_ref-39" class="reference"><a href="#cite_note-39"><span class="cite-bracket">&#91;</span>39<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">&#91;</span>40<span class="cite-bracket">&#93;</span></a></sup> </p><p>Nesting behavior varies among species and may involve hollow tree trunks, external hives, the soil, termite nest or even urban structures. This adaptability underscores their resilience and ability to coexist with human activities.<sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">&#91;</span>41<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Castes">Castes</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=7" title="Edit section&#039;s source code: Castes"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading4"><h4 id="Workers">Workers</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=8" title="Edit section&#039;s source code: Workers"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/enwiki/wiki/Worker_bee" title="Worker bee">Worker bee</a></div> <p>In a SB colony, workers constitute the predominant segment of the population, serving as the colony's primary workforce. They undertake a multitude of responsibilities crucial for the colony's well-being, including defense, cleaning, handling building materials, and the collection and processing of food. Recognizable by the corbicula - a distinctive structure on their hind legs resembling a small basket - workers efficiently carry pollen, resin, clay, and other materials gathered from the environment. Given their abundance and unique physical feature, workers play a central role in sustaining the colony.<sup id="cite_ref-42" class="reference"><a href="#cite_note-42"><span class="cite-bracket">&#91;</span>42<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:9_43-0" class="reference"><a href="#cite_note-:9-43"><span class="cite-bracket">&#91;</span>43<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-44" class="reference"><a href="#cite_note-44"><span class="cite-bracket">&#91;</span>44<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Queens">Queens</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=9" title="Edit section&#039;s source code: Queens"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/enwiki/wiki/Queen_bee" title="Queen bee">Queen bee</a></div> <p>The principal egg layer in SB colonies is the queen, distinguished from the workers by differences in both size and shape. Stingless bee queens - except in the case of the <i><a href="/enwiki/wiki/Melipona" title="Melipona">Melipona</a></i> genus, where queens and workers receive similar amounts of food and thus exhibit similar sizes - are generally larger and weigh more than workers (approximately 2–6 times). Post-mating, meliponine queens undergo <a href="/enwiki/wiki/Physogastrism" title="Physogastrism">physogastry</a>, developing a distended abdomen. This physical transformation sets them apart from honey bee queens, and even <i>Melipona</i> queens can be easily identified by their enlarged abdomen after mating.<sup id="cite_ref-:4_37-1" class="reference"><a href="#cite_note-:4-37"><span class="cite-bracket">&#91;</span>37<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:6_45-0" class="reference"><a href="#cite_note-:6-45"><span class="cite-bracket">&#91;</span>45<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:7_46-0" class="reference"><a href="#cite_note-:7-46"><span class="cite-bracket">&#91;</span>46<span class="cite-bracket">&#93;</span></a></sup> </p><p>Stingless bee colonies typically follow a monogynous structure, featuring a single egg-laying queen. An exception is noted in <i><a href="/enwiki/wiki/Melipona_bicolor" title="Melipona bicolor">Melipona bicolor</a></i> colonies, which are often polygynous (large populations may have as many as 5 physogastric queens simultaneously involved in oviposition).<sup id="cite_ref-:7_46-1" class="reference"><a href="#cite_note-:7-46"><span class="cite-bracket">&#91;</span>46<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:5_38-1" class="reference"><a href="#cite_note-:5-38"><span class="cite-bracket">&#91;</span>38<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:8_47-0" class="reference"><a href="#cite_note-:8-47"><span class="cite-bracket">&#91;</span>47<span class="cite-bracket">&#93;</span></a></sup> Depending on the species, queens can lay varying quantities of eggs daily, ranging from a dozen (e.g., <i><a href="/enwiki/w/index.php?title=Plebeia_julianii&amp;action=edit&amp;redlink=1" class="new" title="Plebeia julianii (page does not exist)">Plebeia julianii</a></i>) to several hundred (e.g., <i><a href="/enwiki/w/index.php?title=Trigona_recursa&amp;action=edit&amp;redlink=1" class="new" title="Trigona recursa (page does not exist)">Trigona recursa</a></i>). While information on queen lifespans is limited, available data suggest that queens generally outlive workers, with lifespans usually falling between 1 and 3 years, although some queens may live up to 7 years.<sup id="cite_ref-:7_46-2" class="reference"><a href="#cite_note-:7-46"><span class="cite-bracket">&#91;</span>46<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:8_47-1" class="reference"><a href="#cite_note-:8-47"><span class="cite-bracket">&#91;</span>47<span class="cite-bracket">&#93;</span></a></sup> </p><p>The laying queen assumes the crucial role of producing eggs that give rise to all castes within the colony. Additionally, she plays a pivotal role in organizing the colony, overseeing a complex communication system primarily reliant on the use of <a href="/enwiki/wiki/Pheromone" title="Pheromone">pheromones</a>.<sup id="cite_ref-:6_45-1" class="reference"><a href="#cite_note-:6-45"><span class="cite-bracket">&#91;</span>45<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Males_(drones)"><span id="Males_.28drones.29"></span>Males (drones)</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=10" title="Edit section&#039;s source code: Males (drones)"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/enwiki/wiki/Drone_(bee)" title="Drone (bee)">Drone (bee)</a></div> <p>The primary function of males, or drones, is to mate with queens, performing limited tasks within the nest and leaving at around 2–3 weeks old, never to return. The production of males can vary, occurring continuously, sparsely or in large spurts when numerous drones emerge from brood combs for brief periods. Identifying a male can be challenging due to its similar body size to workers, but distinctive features such as the absence of a corbicula, larger eyes, slightly smaller <a href="/enwiki/wiki/Mandible_(insect_mouthpart)" title="Mandible (insect mouthpart)">mandibles</a>, slightly longer and v-shaped <a href="/enwiki/wiki/Antenna_(biology)" class="mw-redirect" title="Antenna (biology)">antennae</a>, and often a lighter face color distinguish them. Clusters of males, numbering in the hundreds, can be observed outside colonies, awaiting the opportunity to mate with virgin queens.<sup id="cite_ref-:9_43-1" class="reference"><a href="#cite_note-:9-43"><span class="cite-bracket">&#91;</span>43<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-48" class="reference"><a href="#cite_note-48"><span class="cite-bracket">&#91;</span>48<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-49" class="reference"><a href="#cite_note-49"><span class="cite-bracket">&#91;</span>49<span class="cite-bracket">&#93;</span></a></sup> </p><p>Males in a stingless bee colony, either produced mainly by the laying queen or primarily by the workers, play an important role in reproduction. Workers can produce males by laying unfertilized eggs, enabled by the <a href="/enwiki/wiki/Haplodiploidy" title="Haplodiploidy">haplodiploidy</a> system, where males are <a href="/enwiki/wiki/Ploidy" title="Ploidy">haploid</a>, having only one set of <a href="/enwiki/wiki/Chromosome" title="Chromosome">chromosomes</a>, while workers are diploid and incapable of producing female eggs due to their inability to mate. This sex determination system is common to all <a href="/enwiki/wiki/Hymenoptera" title="Hymenoptera">hymenopterans</a>.<sup id="cite_ref-50" class="reference"><a href="#cite_note-50"><span class="cite-bracket">&#91;</span>50<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Soldiers">Soldiers</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=11" title="Edit section&#039;s source code: Soldiers"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>While the existence of a soldier caste is well known in <a href="/enwiki/wiki/Ant" title="Ant">ants</a> and <a href="/enwiki/wiki/Termite" title="Termite">termites</a>, the phenomenon was unknown among bees until 2012, when some stingless bees were found to have a similar caste of defensive specialists that help guard the nest entrance against intruders.<sup id="cite_ref-51" class="reference"><a href="#cite_note-51"><span class="cite-bracket">&#91;</span>51<span class="cite-bracket">&#93;</span></a></sup> To date, at least 10 species have been documented to possess such "soldiers", including <i><a href="/enwiki/wiki/Tetragonisca_angustula" title="Tetragonisca angustula">Tetragonisca angustula</a></i>, <i><a href="/enwiki/w/index.php?title=Tetragonisca_fiebrigi&amp;action=edit&amp;redlink=1" class="new" title="Tetragonisca fiebrigi (page does not exist)">T. fiebrigi</a></i>, and <i><a href="/enwiki/w/index.php?title=Frieseomelitta_longipes&amp;action=edit&amp;redlink=1" class="new" title="Frieseomelitta longipes (page does not exist)">Frieseomelitta longipes</a></i>, with the guards not only larger, but also sometimes a different color from ordinary workers.<sup id="cite_ref-52" class="reference"><a href="#cite_note-52"><span class="cite-bracket">&#91;</span>52<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-53" class="reference"><a href="#cite_note-53"><span class="cite-bracket">&#91;</span>53<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Division_of_labour">Division of labour</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=12" title="Edit section&#039;s source code: Division of labour"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>When the young worker bees emerge from their cells, they tend to initially remain inside the hive, performing different jobs. As workers age, they become guards or foragers. Unlike the <a href="/enwiki/wiki/Larva" title="Larva">larvae</a> of <a href="/enwiki/wiki/Honey_bee" title="Honey bee">honey bees</a> and many social wasps, meliponine larvae are not actively fed by adults (<a href="/enwiki/wiki/Progressive_provisioning" title="Progressive provisioning">progressive provisioning</a>). Pollen and nectar are placed in a cell, within which an <a href="/enwiki/wiki/Egg" title="Egg">egg</a> is laid, and the cell is sealed until the adult bee emerges after <a href="/enwiki/wiki/Pupa" title="Pupa">pupation</a> (<a href="/enwiki/wiki/Mass_provisioning" title="Mass provisioning">mass provisioning</a>). </p><p>At any one time, hives can contain from 300 to more than 100,000 workers (with some authors claiming to calculate more than 150,000 workers, but with no methodology explanation), depending on species.<sup id="cite_ref-54" class="reference"><a href="#cite_note-54"><span class="cite-bracket">&#91;</span>54<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Products_and_materials">Products and materials</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=13" title="Edit section&#039;s source code: Products and materials"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The industrious nature of stingless bees extends to their building activities. Unlike honey bees, they do not use pure wax for construction but combine it with resin to create cerumen, a material employed in constructing nest structures such as brood cells, food pots, and the protective involucrum. <a href="/enwiki/wiki/Beeswax" title="Beeswax">Wax</a> is secreted by young bees through glands located on the top of their abdomen and this mixture not only provides structural strength but also offers antimicrobial properties, inhibiting the growth of fungi and bacteria. The creation of batumen involves combining cerumen with additional resin, mud, plant material, and sometimes even animal feces. Batumen, a stronger material, forms protective layers covering the walls of the nesting space, ensuring the safety of the colony.<sup id="cite_ref-:10_55-0" class="reference"><a href="#cite_note-:10-55"><span class="cite-bracket">&#91;</span>55<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:11_56-0" class="reference"><a href="#cite_note-:11-56"><span class="cite-bracket">&#91;</span>56<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:12_57-0" class="reference"><a href="#cite_note-:12-57"><span class="cite-bracket">&#91;</span>57<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:13_58-0" class="reference"><a href="#cite_note-:13-58"><span class="cite-bracket">&#91;</span>58<span class="cite-bracket">&#93;</span></a></sup> </p><p>On the other hand, <a href="/enwiki/wiki/Clay" title="Clay">clay</a>, sourced from the wild and exhibiting diverse colors based on its mineral origin, serves as another essential raw material for SB. While it can be used in its pure form, it is more common to combine clay with vegetable <a href="/enwiki/wiki/Resin" title="Resin">resins</a> to produce geopropolis. The inclusion of clay in this mixture enhances the durability and structural integrity of the resulting substance.<sup id="cite_ref-:10_55-1" class="reference"><a href="#cite_note-:10-55"><span class="cite-bracket">&#91;</span>55<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:11_56-1" class="reference"><a href="#cite_note-:11-56"><span class="cite-bracket">&#91;</span>56<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:12_57-1" class="reference"><a href="#cite_note-:12-57"><span class="cite-bracket">&#91;</span>57<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:13_58-1" class="reference"><a href="#cite_note-:13-58"><span class="cite-bracket">&#91;</span>58<span class="cite-bracket">&#93;</span></a></sup> </p><p>Vegetable resin, gathered from a variety of plant species in the wild, is an essential raw material brought back to the hive. Stored in small, sticky clumps in peripheral areas of the colony, it is often mistakenly treated as a synonym for <a href="/enwiki/wiki/Propolis" title="Propolis">propolis</a>. However, in beekeeping terminology, propolis refers to a mixture of resin, wax, enzymes, and possibly other substances. Stingless bees go beyond the classic propolis by producing various derivatives from resins and wax, sometimes using pure resins for sealing or defense, a behavior not observed in <i>Apis</i> bees. Understanding these distinctions is vital for effective production and value addition to the meliponiculture activity.<sup id="cite_ref-:10_55-2" class="reference"><a href="#cite_note-:10-55"><span class="cite-bracket">&#91;</span>55<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:11_56-2" class="reference"><a href="#cite_note-:11-56"><span class="cite-bracket">&#91;</span>56<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:12_57-2" class="reference"><a href="#cite_note-:12-57"><span class="cite-bracket">&#91;</span>57<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:13_58-2" class="reference"><a href="#cite_note-:13-58"><span class="cite-bracket">&#91;</span>58<span class="cite-bracket">&#93;</span></a></sup> </p><p><a href="/enwiki/wiki/Honey" title="Honey">Honey</a>, a prized product of bee colonies, is crafted through the processing of <a href="/enwiki/wiki/Nectar" title="Nectar">nectars</a>, <a href="/enwiki/wiki/Honeydew_(secretion)" title="Honeydew (secretion)">honeydews</a>, and fruit juices by worker bees. They store these collected substances in an extension of their gut called a <a href="/enwiki/wiki/Crop_(anatomy)" title="Crop (anatomy)">crop</a>. Back at the hive, the bees ripen or dehydrate the nectar droplets by spinning them inside their mouthparts until honey is formed. Ripening concentrates the nectar and increases the sugar content, though it is not nearly as concentrated as the honey from <i>Apis mellifera</i>. Stored in food pots, meliponines' honey is often referred to as pot-honey due to its distinctive storage method. Stingless bee honeys differ from <i>A. mellifera</i> honey in terms of color, texture, and flavor, being more liquid with a higher water content. Rich in <a href="/enwiki/wiki/Mineral" title="Mineral">minerals</a>, <a href="/enwiki/wiki/Amino_acid" title="Amino acid">amino acids</a>, and <a href="/enwiki/wiki/Flavonoid" title="Flavonoid">flavonoid</a> compounds, the composition of honey varies among colonies of the same species, influenced by factors such as season, habitat, and collected resources.<sup id="cite_ref-:10_55-3" class="reference"><a href="#cite_note-:10-55"><span class="cite-bracket">&#91;</span>55<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:11_56-3" class="reference"><a href="#cite_note-:11-56"><span class="cite-bracket">&#91;</span>56<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:12_57-3" class="reference"><a href="#cite_note-:12-57"><span class="cite-bracket">&#91;</span>57<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:13_58-3" class="reference"><a href="#cite_note-:13-58"><span class="cite-bracket">&#91;</span>58<span class="cite-bracket">&#93;</span></a></sup> </p><p>Special methods are being developed to harvest moderate amounts of honey from stingless bees in these areas without causing harm. For honey production, the bees need to be kept in a box specially designed to make the honey stores accessible without damaging the rest of the nest structure. Some recent box designs for honey production provide a separate compartment for the honey stores so the honey pots can be removed without spilling honey into other areas of the nest. Unlike a hive of commercial honeybees, which can produce 75 kg (165 lbs) of honey a year, a hive of Australian stingless bees produces less than 1 kg (2 lbs). Stingless bee honey has a distinctive "bush" taste—a mix of sweet and sour with a hint of fruit.<sup id="cite_ref-59" class="reference"><a href="#cite_note-59"><span class="cite-bracket">&#91;</span>59<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-60" class="reference"><a href="#cite_note-60"><span class="cite-bracket">&#91;</span>60<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-61" class="reference"><a href="#cite_note-61"><span class="cite-bracket">&#91;</span>61<span class="cite-bracket">&#93;</span></a></sup> The taste comes from plant resins—which the bees use to build their hives and honey pots—and varies at different times of year depending on the flowers and trees visited. </p><p>In 2020 researchers at the <a href="/enwiki/wiki/University_of_Queensland" title="University of Queensland">University of Queensland</a> found that some species of stingless bee in Australia, Malaysia, and Brazil produce honey that has <a href="/enwiki/wiki/Trehalulose" title="Trehalulose">trehalulose</a>—a sugar with an unusually low <a href="/enwiki/wiki/Glycemic_index" title="Glycemic index">glycaemic index</a> (GI) compared to that of glucose and fructose, the main sugars composing conventional honey.<sup id="cite_ref-62" class="reference"><a href="#cite_note-62"><span class="cite-bracket">&#91;</span>62<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:25_63-0" class="reference"><a href="#cite_note-:25-63"><span class="cite-bracket">&#91;</span>63<span class="cite-bracket">&#93;</span></a></sup> Such low glycaemic index honey is beneficial for humans because its consumption does not cause <a href="/enwiki/wiki/Blood_sugar_level" title="Blood sugar level">blood sugar</a> to spike, forcing the body to make more insulin in response. Honey with trehalulose is also beneficial as it this sugar cannot nourish the lactic acid-producing bacteria that cause <a href="/enwiki/wiki/Tooth_decay" title="Tooth decay">tooth decay</a>. The university's findings supported the long-standing claims of <a href="/enwiki/wiki/Indigenous_Australian_people" class="mw-redirect" title="Indigenous Australian people">Indigenous Australian people</a> that native honey is beneficial to human health.<sup id="cite_ref-:25_63-1" class="reference"><a href="#cite_note-:25-63"><span class="cite-bracket">&#91;</span>63<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-64" class="reference"><a href="#cite_note-64"><span class="cite-bracket">&#91;</span>64<span class="cite-bracket">&#93;</span></a></sup> This type of honey is scientifically supported as providing therapeutic value to humans as well.<sup id="cite_ref-:25_63-2" class="reference"><a href="#cite_note-:25-63"><span class="cite-bracket">&#91;</span>63<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-65" class="reference"><a href="#cite_note-65"><span class="cite-bracket">&#91;</span>65<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-66" class="reference"><a href="#cite_note-66"><span class="cite-bracket">&#91;</span>66<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-67" class="reference"><a href="#cite_note-67"><span class="cite-bracket">&#91;</span>67<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-68" class="reference"><a href="#cite_note-68"><span class="cite-bracket">&#91;</span>68<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Nest">Nest</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=14" title="Edit section&#039;s source code: Nest"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/enwiki/wiki/File:Natural_cavity_stingless_bee_nest.png" class="mw-file-description"><img src="/upwiki/wikipedia/commons/thumb/d/d3/Natural_cavity_stingless_bee_nest.png/220px-Natural_cavity_stingless_bee_nest.png" decoding="async" width="220" height="302" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/d/d3/Natural_cavity_stingless_bee_nest.png/330px-Natural_cavity_stingless_bee_nest.png 1.5x, /upwiki/wikipedia/commons/thumb/d/d3/Natural_cavity_stingless_bee_nest.png/440px-Natural_cavity_stingless_bee_nest.png 2x" data-file-width="1105" data-file-height="1515" /></a><figcaption>General main elements of stingless bees cavity nest colonies.</figcaption></figure> <p>Stingless bees, as a collective group, display remarkable adaptability to diverse nesting sites. They can be found in exposed nests in trees, from living in ant and termite nests above and below ground to cavities in trees, trunks, branches, rocks, or even human constructions.<sup id="cite_ref-:14_69-0" class="reference"><a href="#cite_note-:14-69"><span class="cite-bracket">&#91;</span>69<span class="cite-bracket">&#93;</span></a></sup> </p><p>Many <a href="/enwiki/wiki/Beekeeper" title="Beekeeper">beekeepers</a> keep the bees in their original log <a href="/enwiki/wiki/Beehive" title="Beehive">hive</a> or transfer them to a wooden box, as this makes controlling the hive easier. Some beekeepers put them in bamboos, flowerpots, coconut shells, and other recycling containers such as a water jug, a broken guitar, and other safe and closed containers.<sup id="cite_ref-70" class="reference"><a href="#cite_note-70"><span class="cite-bracket">&#91;</span>70<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-71" class="reference"><a href="#cite_note-71"><span class="cite-bracket">&#91;</span>71<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:26_72-0" class="reference"><a href="#cite_note-:26-72"><span class="cite-bracket">&#91;</span>72<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Exposed_nests">Exposed nests</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=15" title="Edit section&#039;s source code: Exposed nests"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/enwiki/wiki/File:Arboreal_stingless_bee_nest_(Trigona_sp.)_Flores.jpg" class="mw-file-description"><img src="/upwiki/wikipedia/commons/thumb/1/1d/Arboreal_stingless_bee_nest_%28Trigona_sp.%29_Flores.jpg/220px-Arboreal_stingless_bee_nest_%28Trigona_sp.%29_Flores.jpg" decoding="async" width="220" height="220" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/1/1d/Arboreal_stingless_bee_nest_%28Trigona_sp.%29_Flores.jpg/330px-Arboreal_stingless_bee_nest_%28Trigona_sp.%29_Flores.jpg 1.5x, /upwiki/wikipedia/commons/thumb/1/1d/Arboreal_stingless_bee_nest_%28Trigona_sp.%29_Flores.jpg/440px-Arboreal_stingless_bee_nest_%28Trigona_sp.%29_Flores.jpg 2x" data-file-width="3888" data-file-height="3888" /></a><figcaption><i>Trigona</i> sp. exposed nest in a tree fork near <a href="/enwiki/wiki/Flores,_Pet%C3%A9n" title="Flores, Petén">Flores</a>, <a href="/enwiki/wiki/Guatemala" title="Guatemala">Guatemala</a>.</figcaption></figure> <p>Notably, certain species, such as the African <i>Dactylurina</i>, construct hanging nests from the undersides of large branches for protection against adverse weather conditions. Additionally, some American <i><a href="/enwiki/wiki/Trigona" title="Trigona">Trigona</a></i> species, including <i><a href="/enwiki/wiki/Trigona_corvina" title="Trigona corvina">T. corvina</a></i>, <i><a href="/enwiki/wiki/Trigona_spinipes" title="Trigona spinipes">T. spinipes</a></i>, and T. nigerrima, as well as <i>Tetragonisca weyrauchi</i>, build fully exposed nests.<sup id="cite_ref-:14_69-1" class="reference"><a href="#cite_note-:14-69"><span class="cite-bracket">&#91;</span>69<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Ground_nests">Ground nests</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=16" title="Edit section&#039;s source code: Ground nests"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/enwiki/wiki/File:Paratrigona_ground_nest.jpg" class="mw-file-description"><img src="/upwiki/wikipedia/commons/thumb/3/3b/Paratrigona_ground_nest.jpg/220px-Paratrigona_ground_nest.jpg" decoding="async" width="220" height="293" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/3/3b/Paratrigona_ground_nest.jpg/330px-Paratrigona_ground_nest.jpg 1.5x, /upwiki/wikipedia/commons/thumb/3/3b/Paratrigona_ground_nest.jpg/440px-Paratrigona_ground_nest.jpg 2x" data-file-width="3024" data-file-height="4032" /></a><figcaption><i>Paratrigona</i> sp. ground nest in <a href="/enwiki/wiki/Bras%C3%ADlia" title="Brasília">Brasília</a>, <a href="/enwiki/wiki/Brazil" title="Brazil">Brazil</a>.</figcaption></figure> <p>A significant minority of meliponine species, belonging to genera like <i><a href="/enwiki/wiki/Camargoia" title="Camargoia">Camargoia</a></i>, <i><a href="/enwiki/wiki/Geotrigona" title="Geotrigona">Geotrigona</a></i>, <i><a href="/enwiki/wiki/Melipona" title="Melipona">Melipona</a></i>, <i><a href="/enwiki/w/index.php?title=Mourella&amp;action=edit&amp;redlink=1" class="new" title="Mourella (page does not exist)">Mourella</a></i>, <i><a href="/enwiki/wiki/Nogueirapis" title="Nogueirapis">Nogueirapis</a></i>, <i><a href="/enwiki/wiki/Paratrigona" title="Paratrigona">Paratrigona</a></i>, <i><a href="/enwiki/wiki/Partamona" title="Partamona">Partamona</a></i>, <i><a href="/enwiki/wiki/Schwarziana" title="Schwarziana">Schwarziana</a></i>, and others, opt for ground nests. These species take advantage of cavities in the ground, often utilizing abandoned nests of ants, termites, or rodents. Unlike some other cavity-nesting bees, stingless bees in this category do not excavate their own cavities but may enlarge existing ones.<sup id="cite_ref-:14_69-2" class="reference"><a href="#cite_note-:14-69"><span class="cite-bracket">&#91;</span>69<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Termite_and_ant_shared_nests">Termite and ant shared nests</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=17" title="Edit section&#039;s source code: Termite and ant shared nests"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Numerous stingless bee species have evolved to coexist with termites. They inhabit parts of ant or termite nests, both above and below ground. These nests are often associated with various ant species, such as <i><a href="/enwiki/wiki/Azteca_(ant)" title="Azteca (ant)">Azteca</a></i>, <i><a href="/enwiki/wiki/Carpenter_ant" title="Carpenter ant">Camponotus</a></i>, or <i><a href="/enwiki/wiki/Crematogaster" title="Crematogaster">Crematogaster</a></i>, and termite species like <i><a href="/enwiki/wiki/Nasutitermes" title="Nasutitermes">Nasutitermes</a></i>, <i><a href="/enwiki/wiki/Constrictotermes" title="Constrictotermes">Constrictotermes</a></i>, <i><a href="/enwiki/wiki/Macrotermes" title="Macrotermes">Macrotermes</a></i>, <i>Microcerotermes</i>, <i><a href="/enwiki/wiki/Odontotermes" title="Odontotermes">Odontotermes</a></i>, or <i>Pseudocanthotermes</i>. This strategy allows SB to utilize pre-existing cavities without the need for extensive excavation.<sup id="cite_ref-:14_69-3" class="reference"><a href="#cite_note-:14-69"><span class="cite-bracket">&#91;</span>69<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Cavity_nests">Cavity nests</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=18" title="Edit section&#039;s source code: Cavity nests"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The majority of stingless bees favor nesting in pre-existing cavities within tree trunks or branches. Nesting heights vary, with some colonies positioned close to the ground, typically below 5 meters, while others, like <i>Trigona</i> and <i><a href="/enwiki/wiki/Oxytrigona" title="Oxytrigona">Oxytrigona</a></i>, may nest at higher elevations, ranging from 10 to 25 meters. Some species, such as <i><a href="/enwiki/w/index.php?title=Melipona_nigra&amp;action=edit&amp;redlink=1" class="new" title="Melipona nigra (page does not exist)">Melipona nigra</a></i>, exhibit unique nesting habits at the foot of a tree in root cavities or between roots. The choice of nesting height has implications for predation pressure and the microclimate experienced by the colony.<sup id="cite_ref-:14_69-4" class="reference"><a href="#cite_note-:14-69"><span class="cite-bracket">&#91;</span>69<span class="cite-bracket">&#93;</span></a></sup> </p><p>The majority of stingless bee species exhibit a non-specific preference when it comes to selecting tree species for nesting. Instead, they opportunistically exploit whatever nesting sites are available This adaptability underscores the versatility of SB in adapting to various arboreal environments. Furthermore, cavity-nesting species can opportunistically utilize human constructions, nesting under roofs, in hollow spaces in walls, electricity boxes, or even metal tubes. In few cases, specific tree species, like <i><a href="/enwiki/wiki/Caryocar_brasiliense" title="Caryocar brasiliense">Caryocar brasiliense</a></i>, may be preferred by certain stingless bee species (<i><a href="/enwiki/wiki/Melipona_quadrifasciata" title="Melipona quadrifasciata">Melipona quadrifasciata</a></i>), illustrating a degree of selectivity in nesting choices among different groups.<sup id="cite_ref-:14_69-5" class="reference"><a href="#cite_note-:14-69"><span class="cite-bracket">&#91;</span>69<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-73" class="reference"><a href="#cite_note-73"><span class="cite-bracket">&#91;</span>73<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Entrances">Entrances</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=19" title="Edit section&#039;s source code: Entrances"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Entrance tubes showcase a spectrum of characteristics, from being hard and brittle to soft and flexible. In many situations, the portion near the opening remains soft and flexible, aiding workers in sealing the entrance during the night. The tubes may also feature perforations and a coating of resin droplets, adding to the complexity of their design.<sup id="cite_ref-:15_74-0" class="reference"><a href="#cite_note-:15-74"><span class="cite-bracket">&#91;</span>74<span class="cite-bracket">&#93;</span></a></sup> </p><p>The entrances serve as essential visual landmarks for returning bees, and they are often the first structures constructed at a new nest site. The diversity in entrance size influences foraging traffic, with larger entrances facilitating smoother traffic but potentially necessitating more entrance guards to ensure adequate defense.<sup id="cite_ref-:15_74-1" class="reference"><a href="#cite_note-:15-74"><span class="cite-bracket">&#91;</span>74<span class="cite-bracket">&#93;</span></a></sup> </p><p>Some <i><a href="/enwiki/wiki/Partamona" title="Partamona">Partamona</a></i> species exhibit a distinctive entrance architecture, where workers of <i><a href="/enwiki/wiki/Partamona_helleri" title="Partamona helleri">P. helleri</a></i> construct a large outer mud entrance leading to a smaller adjacent entrance. This unique design enables foragers to enter with high speed, bouncing off the ceiling of the outer entrance towards the smaller inner entrance. The peculiar appearance of this entrance has led to local names such as "toad mouth", highlighting the intriguing adaptations found in stingless bee nest entrances.<sup id="cite_ref-:15_74-2" class="reference"><a href="#cite_note-:15-74"><span class="cite-bracket">&#91;</span>74<span class="cite-bracket">&#93;</span></a></sup> </p> <ul class="gallery mw-gallery-traditional" style="max-width: 812px;"> <li class="gallerycaption">Different species nest entrances in Brazilian stingless bees</li> <li class="gallerybox" style="width: 195px"> <div class="thumb" style="width: 190px; height: 150px;"><span typeof="mw:File"><a href="/enwiki/wiki/File:Abelha_lim%C3%A3o.jpg" class="mw-file-description" title="Lestrimelitta limao"><img alt="Lestrimelitta limao" src="/upwiki/wikipedia/commons/thumb/2/29/Abelha_lim%C3%A3o.jpg/90px-Abelha_lim%C3%A3o.jpg" decoding="async" width="90" height="120" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/2/29/Abelha_lim%C3%A3o.jpg/135px-Abelha_lim%C3%A3o.jpg 1.5x, /upwiki/wikipedia/commons/thumb/2/29/Abelha_lim%C3%A3o.jpg/180px-Abelha_lim%C3%A3o.jpg 2x" data-file-width="3024" data-file-height="4032" /></a></span></div> <div class="gallerytext"><i><a href="/enwiki/wiki/Lestrimelitta_limao" title="Lestrimelitta limao">Lestrimelitta limao</a></i></div> </li> <li class="gallerybox" style="width: 195px"> <div class="thumb" style="width: 190px; height: 150px;"><span typeof="mw:File"><a href="/enwiki/wiki/File:Frieseomelitta_varia_nest_entrance.jpg" class="mw-file-description" title="Frieseomelitta varia"><img alt="Frieseomelitta varia" src="/upwiki/wikipedia/commons/thumb/5/5b/Frieseomelitta_varia_nest_entrance.jpg/157px-Frieseomelitta_varia_nest_entrance.jpg" decoding="async" width="157" height="120" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/5/5b/Frieseomelitta_varia_nest_entrance.jpg/236px-Frieseomelitta_varia_nest_entrance.jpg 1.5x, /upwiki/wikipedia/commons/thumb/5/5b/Frieseomelitta_varia_nest_entrance.jpg/314px-Frieseomelitta_varia_nest_entrance.jpg 2x" data-file-width="1080" data-file-height="825" /></a></span></div> <div class="gallerytext"><i><a href="/enwiki/wiki/Frieseomelitta_varia" title="Frieseomelitta varia">Frieseomelitta varia</a></i></div> </li> <li class="gallerybox" style="width: 195px"> <div class="thumb" style="width: 190px; height: 150px;"><span typeof="mw:File"><a href="/enwiki/wiki/File:Partamona_helleri_colmeia.jpg" class="mw-file-description" title="Partamona helleri"><img alt="Partamona helleri" src="/upwiki/wikipedia/commons/thumb/a/a7/Partamona_helleri_colmeia.jpg/67px-Partamona_helleri_colmeia.jpg" decoding="async" width="67" height="120" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/a/a7/Partamona_helleri_colmeia.jpg/101px-Partamona_helleri_colmeia.jpg 1.5x, /upwiki/wikipedia/commons/thumb/a/a7/Partamona_helleri_colmeia.jpg/135px-Partamona_helleri_colmeia.jpg 2x" data-file-width="577" data-file-height="1024" /></a></span></div> <div class="gallerytext"><i><a href="/enwiki/wiki/Partamona_helleri" title="Partamona helleri">Partamona helleri</a></i></div> </li> <li class="gallerybox" style="width: 195px"> <div class="thumb" style="width: 190px; height: 150px;"><span typeof="mw:File"><a href="/enwiki/wiki/File:Entrada_de_ninho_de_Scaptotrigona_postica.jpg" class="mw-file-description" title="Scaptotrigona postica"><img alt="Scaptotrigona postica" src="/upwiki/wikipedia/commons/thumb/8/8c/Entrada_de_ninho_de_Scaptotrigona_postica.jpg/90px-Entrada_de_ninho_de_Scaptotrigona_postica.jpg" decoding="async" width="90" height="120" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/8/8c/Entrada_de_ninho_de_Scaptotrigona_postica.jpg/135px-Entrada_de_ninho_de_Scaptotrigona_postica.jpg 1.5x, /upwiki/wikipedia/commons/thumb/8/8c/Entrada_de_ninho_de_Scaptotrigona_postica.jpg/180px-Entrada_de_ninho_de_Scaptotrigona_postica.jpg 2x" data-file-width="3024" data-file-height="4032" /></a></span></div> <div class="gallerytext"><i><a href="/enwiki/wiki/Scaptotrigona_postica" title="Scaptotrigona postica">Scaptotrigona postica</a></i></div> </li> <li class="gallerybox" style="width: 195px"> <div class="thumb" style="width: 190px; height: 150px;"><span typeof="mw:File"><a href="/enwiki/wiki/File:Parque_Nacional_da_Serra_do_Itaja%C3%AD_-_Liu_Id%C3%A1rraga_Orozco_(86).jpg" class="mw-file-description" title="Oxytrigona tataira"><img alt="Oxytrigona tataira" src="/upwiki/wikipedia/commons/thumb/6/6b/Parque_Nacional_da_Serra_do_Itaja%C3%AD_-_Liu_Id%C3%A1rraga_Orozco_%2886%29.jpg/90px-Parque_Nacional_da_Serra_do_Itaja%C3%AD_-_Liu_Id%C3%A1rraga_Orozco_%2886%29.jpg" decoding="async" width="90" height="120" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/6/6b/Parque_Nacional_da_Serra_do_Itaja%C3%AD_-_Liu_Id%C3%A1rraga_Orozco_%2886%29.jpg/135px-Parque_Nacional_da_Serra_do_Itaja%C3%AD_-_Liu_Id%C3%A1rraga_Orozco_%2886%29.jpg 1.5x, /upwiki/wikipedia/commons/thumb/6/6b/Parque_Nacional_da_Serra_do_Itaja%C3%AD_-_Liu_Id%C3%A1rraga_Orozco_%2886%29.jpg/180px-Parque_Nacional_da_Serra_do_Itaja%C3%AD_-_Liu_Id%C3%A1rraga_Orozco_%2886%29.jpg 2x" data-file-width="1836" data-file-height="2448" /></a></span></div> <div class="gallerytext"><i><a href="/enwiki/wiki/Oxytrigona_tataira" title="Oxytrigona tataira">Oxytrigona tataira</a></i></div> </li> <li class="gallerybox" style="width: 195px"> <div class="thumb" style="width: 190px; height: 150px;"><span typeof="mw:File"><a href="/enwiki/wiki/File:Melipona_quadrifasciata_nest_guard.jpg" class="mw-file-description" title="Melipona quadrifasciata"><img alt="Melipona quadrifasciata" src="/upwiki/wikipedia/commons/thumb/e/ea/Melipona_quadrifasciata_nest_guard.jpg/160px-Melipona_quadrifasciata_nest_guard.jpg" decoding="async" width="160" height="120" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/e/ea/Melipona_quadrifasciata_nest_guard.jpg/240px-Melipona_quadrifasciata_nest_guard.jpg 1.5x, /upwiki/wikipedia/commons/thumb/e/ea/Melipona_quadrifasciata_nest_guard.jpg/320px-Melipona_quadrifasciata_nest_guard.jpg 2x" data-file-width="720" data-file-height="540" /></a></span></div> <div class="gallerytext"><i><a href="/enwiki/wiki/Melipona_quadrifasciata" title="Melipona quadrifasciata">Melipona quadrifasciata</a></i></div> </li> <li class="gallerybox" style="width: 195px"> <div class="thumb" style="width: 190px; height: 150px;"><span typeof="mw:File"><a href="/enwiki/wiki/File:Tetragonisca_angustula_nest_entrance.jpg" class="mw-file-description" title="Tetragonisca angustula"><img alt="Tetragonisca angustula" src="/upwiki/wikipedia/commons/thumb/3/3e/Tetragonisca_angustula_nest_entrance.jpg/160px-Tetragonisca_angustula_nest_entrance.jpg" decoding="async" width="160" height="107" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/3/3e/Tetragonisca_angustula_nest_entrance.jpg/240px-Tetragonisca_angustula_nest_entrance.jpg 1.5x, /upwiki/wikipedia/commons/thumb/3/3e/Tetragonisca_angustula_nest_entrance.jpg/320px-Tetragonisca_angustula_nest_entrance.jpg 2x" data-file-width="6000" data-file-height="4000" /></a></span></div> <div class="gallerytext"><i><a href="/enwiki/wiki/Tetragonisca_angustula" title="Tetragonisca angustula">Tetragonisca angustula</a></i></div> </li> <li class="gallerybox" style="width: 195px"> <div class="thumb" style="width: 190px; height: 150px;"><span typeof="mw:File"><a href="/enwiki/wiki/File:Nannotrigona_testaceicornis_nest_entrance.jpg" class="mw-file-description" title="Nannotrigona testaceicornis"><img alt="Nannotrigona testaceicornis" src="/upwiki/wikipedia/commons/thumb/0/05/Nannotrigona_testaceicornis_nest_entrance.jpg/90px-Nannotrigona_testaceicornis_nest_entrance.jpg" decoding="async" width="90" height="120" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/0/05/Nannotrigona_testaceicornis_nest_entrance.jpg/135px-Nannotrigona_testaceicornis_nest_entrance.jpg 1.5x, /upwiki/wikipedia/commons/thumb/0/05/Nannotrigona_testaceicornis_nest_entrance.jpg/180px-Nannotrigona_testaceicornis_nest_entrance.jpg 2x" data-file-width="2448" data-file-height="3264" /></a></span></div> <div class="gallerytext"><i><a href="/enwiki/wiki/Nannotrigona_testaceicornis" title="Nannotrigona testaceicornis">Nannotrigona testaceicornis</a></i></div> </li> </ul> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/enwiki/wiki/File:Entrant_tubes.png" class="mw-file-description"><img src="/upwiki/wikipedia/commons/thumb/9/9e/Entrant_tubes.png/220px-Entrant_tubes.png" decoding="async" width="220" height="282" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/9/9e/Entrant_tubes.png/330px-Entrant_tubes.png 1.5x, /upwiki/wikipedia/commons/thumb/9/9e/Entrant_tubes.png/440px-Entrant_tubes.png 2x" data-file-width="591" data-file-height="757" /></a><figcaption>Variation of entrant tube in some native stingless bee species found in Southeast Asia: (1) <i><a href="/enwiki/w/index.php?title=Geniotrigona_thoracica&amp;action=edit&amp;redlink=1" class="new" title="Geniotrigona thoracica (page does not exist)">Geniotrigona thoracica</a></i>, (2) <i><a href="/enwiki/w/index.php?title=Homotrigona_fimbriata&amp;action=edit&amp;redlink=1" class="new" title="Homotrigona fimbriata (page does not exist)">Homotrigona fimbriata</a></i>, (3) <i><a href="/enwiki/w/index.php?title=Lophotrigona_canifrons&amp;action=edit&amp;redlink=1" class="new" title="Lophotrigona canifrons (page does not exist)">Lophotrigona canifrons</a></i>, (4) <i><a href="/enwiki/w/index.php?title=Tetragonilla_collina&amp;action=edit&amp;redlink=1" class="new" title="Tetragonilla collina (page does not exist)">Tetragonilla collina</a></i>, (5) <i><a href="/enwiki/w/index.php?title=Pariotrigona_klossi&amp;action=edit&amp;redlink=1" class="new" title="Pariotrigona klossi (page does not exist)">Pariotrigona klossi</a></i>, (6) <i><a href="/enwiki/w/index.php?title=Heterotrigona_itama&amp;action=edit&amp;redlink=1" class="new" title="Heterotrigona itama (page does not exist)">Heterotrigona itama</a></i>, (7) <i><a href="/enwiki/w/index.php?title=Tetragonula_fuscobalteata&amp;action=edit&amp;redlink=1" class="new" title="Tetragonula fuscobalteata (page does not exist)">Tetragonula fuscobalteata</a></i>, (8) <i><a href="/enwiki/w/index.php?title=Lepidotriogona_terminata&amp;action=edit&amp;redlink=1" class="new" title="Lepidotriogona terminata (page does not exist)">Lepidotriogona terminata</a></i>, (9) <i><a href="/enwiki/w/index.php?title=Tetrigona_melanoleuca&amp;action=edit&amp;redlink=1" class="new" title="Tetrigona melanoleuca (page does not exist)">Tetrigona melanoleuca</a></i>, (10) <i><a href="/enwiki/w/index.php?title=Tetrigona_apicalis&amp;action=edit&amp;redlink=1" class="new" title="Tetrigona apicalis (page does not exist)">Tetrigona apicalis</a></i>, and (11) <i><a href="/enwiki/w/index.php?title=Tetragonula_pagdeni&amp;action=edit&amp;redlink=1" class="new" title="Tetragonula pagdeni (page does not exist)">Tetragonula pagdeni</a></i>.</figcaption></figure> <div class="mw-heading mw-heading4"><h4 id="Brood_cell_arrangement">Brood cell arrangement</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=20" title="Edit section&#039;s source code: Brood cell arrangement"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Stingless bee colonies exhibit a diversity of construction patterns of brood cells, primarily composed of soft cerumen, a mixture of wax and resin. Each crafted cell is designed to rear a single individual bee, emphasizing the precision and efficiency of their nest architecture.<sup id="cite_ref-:15_74-3" class="reference"><a href="#cite_note-:15-74"><span class="cite-bracket">&#91;</span>74<span class="cite-bracket">&#93;</span></a></sup> </p><p>The quantity of brood cells within a nest displays significant variation across different stingless bee species. Nest size can range from a few brood cells, as observed in the Asian <i><a href="/enwiki/w/index.php?title=Lisotrigona_carpenteri&amp;action=edit&amp;redlink=1" class="new" title="Lisotrigona carpenteri (page does not exist)">Lisotrigona carpenteri</a></i>, to remarkably expansive colonies with over 80,000 brood cells, particularly in some American <i><a href="/enwiki/wiki/Trigona" title="Trigona">Trigona</a></i> species.<sup id="cite_ref-:15_74-4" class="reference"><a href="#cite_note-:15-74"><span class="cite-bracket">&#91;</span>74<span class="cite-bracket">&#93;</span></a></sup> </p><p>Meliponine colonies exhibit diverse brood cell arrangements, primarily categorized into three main types: horizontal combs, vertical combs, and clustered cells. Despite these primary types, variations and intermediate forms are prevalent, contributing to the flexibility of nest structures.<sup id="cite_ref-:15_74-5" class="reference"><a href="#cite_note-:15-74"><span class="cite-bracket">&#91;</span>74<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-75" class="reference"><a href="#cite_note-75"><span class="cite-bracket">&#91;</span>75<span class="cite-bracket">&#93;</span></a></sup> </p><p> The first type involves horizontal combs, often characterized by a spiral pattern or layers of cells. The presence of spirals may not be consistent within a species, varying among colonies or even within the same colony. Some species, such as <i><a href="/enwiki/wiki/Melipona" title="Melipona">Melipona</a></i>, <i><a href="/enwiki/wiki/Plebeia" title="Plebeia">Plebeia</a></i>, <i>Plebeina</i>, <i><a href="/enwiki/wiki/Nannotrigona" title="Nannotrigona">Nannotrigona</a></i>, <i>Trigona</i>, and <i>Tetragona</i>, may occasionally build spirals alongside other comb structures, as observed in <i><a href="/enwiki/w/index.php?title=Oxytrigona_mellicolor&amp;action=edit&amp;redlink=1" class="new" title="Oxytrigona mellicolor (page does not exist)">Oxytrigona mellicolor</a></i>. As space diminishes for upward construction, workers initiate the creation of a new comb at the bottom of the brood chamber. This innovative approach optimizes the available space when emerging bees vacate older, lower brood combs.<sup id="cite_ref-:15_74-6" class="reference"><a href="#cite_note-:15-74"><span class="cite-bracket">&#91;</span>74<span class="cite-bracket">&#93;</span></a></sup></p><figure class="mw-halign-left" typeof="mw:File/Thumb"><a href="/enwiki/wiki/File:Sarang_lebah_kelulut.jpeg" class="mw-file-description"><img src="/upwiki/wikipedia/commons/thumb/7/7b/Sarang_lebah_kelulut.jpeg/250px-Sarang_lebah_kelulut.jpeg" decoding="async" width="250" height="184" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/7/7b/Sarang_lebah_kelulut.jpeg/375px-Sarang_lebah_kelulut.jpeg 1.5x, /upwiki/wikipedia/commons/thumb/7/7b/Sarang_lebah_kelulut.jpeg/500px-Sarang_lebah_kelulut.jpeg 2x" data-file-width="1732" data-file-height="1272" /></a><figcaption>Hive box containing colony of <i><a href="/enwiki/w/index.php?title=Heterotrigona_itama&amp;action=edit&amp;redlink=1" class="new" title="Heterotrigona itama (page does not exist)">Heterotrigona itama</a></i></figcaption></figure> <p>The second prevalent brood cell arrangement involves clusters of cells held together with thin cerumen connections. This clustered style is observed in various distantly related genera, such as the American <i><a href="/enwiki/wiki/Trigonisca" title="Trigonisca">Trigonisca</a></i>, <i><a href="/enwiki/wiki/Frieseomelitta" title="Frieseomelitta">Frieseomelitta</a></i>, <i><a href="/enwiki/wiki/Leurotrigona" title="Leurotrigona">Leurotrigona</a></i>, the Australian <i><a href="/enwiki/wiki/Austroplebeia" title="Austroplebeia">Austroplebeia</a></i>, and the African <i><a href="/enwiki/w/index.php?title=Hypotrigona&amp;action=edit&amp;redlink=1" class="new" title="Hypotrigona (page does not exist)">Hypotrigona</a></i>. This arrangement is particularly useful for colonies in irregular cavities unsuitable for traditional comb building.<sup id="cite_ref-:15_74-7" class="reference"><a href="#cite_note-:15-74"><span class="cite-bracket">&#91;</span>74<span class="cite-bracket">&#93;</span></a></sup> </p><p>The construction of vertical combs is a distinctive trait found in only two stingless bee species: the African <i><a href="/enwiki/wiki/Dactylurina" title="Dactylurina">Dactylurina</a></i> and the American <i><a href="/enwiki/wiki/Scaura_longula" title="Scaura longula">Scaura longula</a></i>. This vertical arrangement sets these species apart from the more commonly observed horizontal comb structures in other stingless bee genera.<sup id="cite_ref-:15_74-8" class="reference"><a href="#cite_note-:15-74"><span class="cite-bracket">&#91;</span>74<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Brood_rearing">Brood rearing</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=21" title="Edit section&#039;s source code: Brood rearing"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Stingless bee brood rearing is a sophisticated and intricately coordinated process involving various tasks performed by worker bees, closely synchronized with the queen's activities. The sequence begins with the completion of a new brood cell, marking the initiation of mass provisioning.<sup id="cite_ref-:16_76-0" class="reference"><a href="#cite_note-:16-76"><span class="cite-bracket">&#91;</span>76<span class="cite-bracket">&#93;</span></a></sup> </p><p>Upon finishing a brood cell, several workers engage in mass provisioning, regurgitating larval food into the cell. This collective effort is swiftly followed by the queen laying her egg on top of the provided larval food. The immediate sealing of the cell ensues shortly afterward, culminating this important phase of the brood rearing process.<sup id="cite_ref-:16_76-1" class="reference"><a href="#cite_note-:16-76"><span class="cite-bracket">&#91;</span>76<span class="cite-bracket">&#93;</span></a></sup> </p><p>The practice of mass provisioning, oviposition, and cell sealing is considered an ancestral trait, shared with solitary wasps and bees. However, in the context of stingless bees, these actions represent distinct stages of a highly integrated social process. Notably, the queen plays a central role in orchestrating these activities, acting as a pacemaker for the entire colony.<sup id="cite_ref-:16_76-2" class="reference"><a href="#cite_note-:16-76"><span class="cite-bracket">&#91;</span>76<span class="cite-bracket">&#93;</span></a></sup> </p><p>This process diverges significantly from brood rearing in <i>Apis</i> spp. In honeybee colonies, queens lay eggs into reusable empty cells, which are then progressively provisioned over several days before final sealing. The contrasting approaches in brood rearing highlight the unique social dynamics and adaptations within stingless bee colonies.<sup id="cite_ref-:16_76-3" class="reference"><a href="#cite_note-:16-76"><span class="cite-bracket">&#91;</span>76<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Swarming">Swarming</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=22" title="Edit section&#039;s source code: Swarming"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/enwiki/wiki/Swarming_(honey_bee)" title="Swarming (honey bee)">Swarming (honey bee)</a></div> <p>Stingless bees and honey bees, despite encountering a common challenge in establishing daughter colonies, employ contrasting strategies. There are three key differences: reproductive status and age of the queen that leaves the nest, temporal aspects of colony foundation, and communication processes for nest site selection.<sup id="cite_ref-:17_77-0" class="reference"><a href="#cite_note-:17-77"><span class="cite-bracket">&#91;</span>77<span class="cite-bracket">&#93;</span></a></sup> </p><p>In HB (<i>Apis mellifera</i>), the mother queen, accompanied by a swarm of numerous workers, embarks on relocation to a new home once replacement queens have been reared. Conversely, in SB (meliponines), the departure is orchestrated by the unmated ("virgin") queen, leaving the mother queen in the original nest. Mated stingless bee cannot leave the hive due to damaged wings and increased abdominal size post-mating (<a href="/enwiki/wiki/Physogastrism" title="Physogastrism">physogastrism</a>). The queen's weight in species like <i><a href="/enwiki/wiki/Scaptotrigona_postica" title="Scaptotrigona postica">Scaptotrigona postica</a></i> increases, for example, about 250%.<sup id="cite_ref-:17_77-1" class="reference"><a href="#cite_note-:17-77"><span class="cite-bracket">&#91;</span>77<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-78" class="reference"><a href="#cite_note-78"><span class="cite-bracket">&#91;</span>78<span class="cite-bracket">&#93;</span></a></sup> </p><p>Unlike honey bees, stingless bee colonies are unable to perform absconding - a term denoting the abandonment of the nest and migration to a new location - making them reliant on alternative strategies to cope with challenges. Meliponines progressive found new colonies without abandonning their nest abruptly.<sup id="cite_ref-:17_77-2" class="reference"><a href="#cite_note-:17-77"><span class="cite-bracket">&#91;</span>77<span class="cite-bracket">&#93;</span></a></sup> </p><p>These are the stages of stingless bees swarming:<sup id="cite_ref-:17_77-3" class="reference"><a href="#cite_note-:17-77"><span class="cite-bracket">&#91;</span>77<span class="cite-bracket">&#93;</span></a></sup> </p> <ol><li>Reconnaissance and preparation: Scouts inspect potential new nest sites for suitability, considering factors such as cavity size, entrance characteristics, and potential threats. The criteria for determining suitability remain largely unexplored. Some colonies engage in simultaneous preparation of multiple cavities before making a final decision and some others make the initial reconnaissance but do not move into the cavity;</li> <li>Transport of building material and food: workers seal cracks in the chosen cavity using materials like resin, batumen, or mud. They construct an entrance tube, possibly serving as a visual beacon for nestmate workers. Early food pots are built and filled with honey, requiring a growing number of workers to transport cerumen and honey from the mother nest.</li> <li>Progressive establishment and social link: the mother and daughter colony maintain a social link through workers traveling between the two nests. The duration of this link varies among species, ranging from a few days to several months. Stingless bee colonies display a preference for cavities previously used by other colonies, containing remnants of building material and nest structures.</li> <li>Arrival of the queen: after initial preparations, an unmated queen, accompanied by additional workers, arrives at the new nest site.</li> <li>Drone arrival: males (drones) aggregate outside the newly established nest. They often arrive shortly after swarming initiation, even before the completion of nest structures. Males can be observed near the entrance, awaiting further events.</li> <li>Mating flight: males in aggregations do not enter the colony but await the queen's emergence for a mating flight. Although rarely observed, it is assumed that unmated stingless bee queens embark on a single mating flight, utilizing acquired sperm for the entirety of their reproductive life.</li></ol> <div class="mw-heading mw-heading3"><h3 id="Natural_enemies">Natural enemies</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=23" title="Edit section&#039;s source code: Natural enemies"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In meliponiculture, beekeepers need to be aware of the presence of animals that can harm stingless bee colonies. There are several potential enemies, but the most damaging ones to meliponaries are listed below.<sup id="cite_ref-79" class="reference"><a href="#cite_note-79"><span class="cite-bracket">&#91;</span>79<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Invertebrates">Invertebrates</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=24" title="Edit section&#039;s source code: Invertebrates"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/enwiki/wiki/File:For%C3%ADdeo_e_armadilha.png" class="mw-file-description"><img src="/upwiki/wikipedia/commons/thumb/a/a5/For%C3%ADdeo_e_armadilha.png/220px-For%C3%ADdeo_e_armadilha.png" decoding="async" width="220" height="106" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/a/a5/For%C3%ADdeo_e_armadilha.png/330px-For%C3%ADdeo_e_armadilha.png 1.5x, /upwiki/wikipedia/commons/thumb/a/a5/For%C3%ADdeo_e_armadilha.png/440px-For%C3%ADdeo_e_armadilha.png 2x" data-file-width="817" data-file-height="394" /></a><figcaption>Phorid fly and vinegar trap.</figcaption></figure> <p><a href="/enwiki/wiki/Phoridae" title="Phoridae">Phorid</a> flies in the genus <i><a href="/enwiki/w/index.php?title=Pseudohypocera&amp;action=edit&amp;redlink=1" class="new" title="Pseudohypocera (page does not exist)">Pseudohypocera</a></i> pose a significant threat to stingless bee colonies, causing problems for beekeepers. These parasites lay eggs in open cells of pollen and honey, leading to potential extinction if not addressed. Early detection is crucial for manual removal or using <a href="/enwiki/wiki/Vinegar" title="Vinegar">vinegar</a> traps. It's important never to leave an infested box unattended to prevent the cycle from restarting and avoid contaminating other colonies. Careful handling of food jars, especially during swarms transfers, is essential. Prompt removal of broken jars, sealing gaps with wax or tape, and maintaining vigilance during the rainy season for heightened phorid activity are recommended. Combatting these flies usually is a priority, particularly during increased reproductive periods.<sup id="cite_ref-:72_80-0" class="reference"><a href="#cite_note-:72-80"><span class="cite-bracket">&#91;</span>80<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:82_81-0" class="reference"><a href="#cite_note-:82-81"><span class="cite-bracket">&#91;</span>81<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-82" class="reference"><a href="#cite_note-82"><span class="cite-bracket">&#91;</span>82<span class="cite-bracket">&#93;</span></a></sup> </p> <p><a href="/enwiki/wiki/Termite" title="Termite">Termites</a> usually do not attack bees or their food pots. However, they can cause damage to the structure of hive boxes as there are many <a href="/enwiki/wiki/Xylophagy" title="Xylophagy">xylophagous</a> species. While termites do not usually pose major problems for beekeepers, they should still be monitored closely.<sup id="cite_ref-83" class="reference"><a href="#cite_note-83"><span class="cite-bracket">&#91;</span>83<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-84" class="reference"><a href="#cite_note-84"><span class="cite-bracket">&#91;</span>84<span class="cite-bracket">&#93;</span></a></sup></p><figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/enwiki/wiki/File:Hermetia_illucens_Black_soldier_fly_edit1.jpg" class="mw-file-description"><img src="/upwiki/wikipedia/commons/thumb/4/42/Hermetia_illucens_Black_soldier_fly_edit1.jpg/220px-Hermetia_illucens_Black_soldier_fly_edit1.jpg" decoding="async" width="220" height="147" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/4/42/Hermetia_illucens_Black_soldier_fly_edit1.jpg/330px-Hermetia_illucens_Black_soldier_fly_edit1.jpg 1.5x, /upwiki/wikipedia/commons/thumb/4/42/Hermetia_illucens_Black_soldier_fly_edit1.jpg/440px-Hermetia_illucens_Black_soldier_fly_edit1.jpg 2x" data-file-width="1800" data-file-height="1200" /></a><figcaption><a href="/enwiki/wiki/Hermetia_illucens" title="Hermetia illucens">Black soldier fly.</a></figcaption></figure> <figure typeof="mw:File/Thumb"><a href="/enwiki/wiki/File:Caixa_racional_individual.png" class="mw-file-description"><img src="/upwiki/wikipedia/commons/thumb/6/61/Caixa_racional_individual.png/125px-Caixa_racional_individual.png" decoding="async" width="125" height="166" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/6/61/Caixa_racional_individual.png/188px-Caixa_racional_individual.png 1.5x, /upwiki/wikipedia/commons/thumb/6/61/Caixa_racional_individual.png/250px-Caixa_racional_individual.png 2x" data-file-width="412" data-file-height="546" /></a><figcaption>Rational box support with burnt oil to prevent ant attacks.</figcaption></figure> <p><a href="/enwiki/wiki/Ant" title="Ant">Ants</a> are attracted to bee colonies by the smell of food. To prevent ant attacks, it's important to handle the hive boxes carefully and avoid exposing jars of pollen and honey. Although rare, when attacks do occur, there are intense conflicts between ants and bees. Stingless bees usually manage to defend themselves, but the damage to the bee population can be significant. To prevent ant infestations in meliponaries with individual supports, a useful strategy is to impregnate the box supports with burnt oil.<sup id="cite_ref-:92_85-0" class="reference"><a href="#cite_note-:92-85"><span class="cite-bracket">&#91;</span>85<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-86" class="reference"><a href="#cite_note-86"><span class="cite-bracket">&#91;</span>86<span class="cite-bracket">&#93;</span></a></sup> </p><p>Another group of enemy flies are the <a href="/enwiki/wiki/Hermetia_illucens" title="Hermetia illucens">black soldier flies</a> (<i>Hermetia illucens</i>). They lay their eggs in crevices of boxes and can extend the tip of their abdomen during laying, facilitating access to the inside of the hive. Larvae of this species feed on pollen, feces, and other materials found in colonies. In general, healthy bee colonies can coexist peacefully with soldier flies. However, in areas where these insects are prevalent, beekeepers must remain vigilant and protect the gaps in the colonies to prevent potential issues.<sup id="cite_ref-87" class="reference"><a href="#cite_note-87"><span class="cite-bracket">&#91;</span>87<span class="cite-bracket">&#93;</span></a></sup> </p> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/enwiki/wiki/File:Entrada_de_ninho_de_Lestrimelitta_tropica.jpg" class="mw-file-description"><img src="/upwiki/wikipedia/commons/thumb/0/00/Entrada_de_ninho_de_Lestrimelitta_tropica.jpg/220px-Entrada_de_ninho_de_Lestrimelitta_tropica.jpg" decoding="async" width="220" height="304" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/0/00/Entrada_de_ninho_de_Lestrimelitta_tropica.jpg/330px-Entrada_de_ninho_de_Lestrimelitta_tropica.jpg 1.5x, /upwiki/wikipedia/commons/thumb/0/00/Entrada_de_ninho_de_Lestrimelitta_tropica.jpg/440px-Entrada_de_ninho_de_Lestrimelitta_tropica.jpg 2x" data-file-width="4160" data-file-height="5744" /></a><figcaption><i>Lestrimelitta limao</i> hive entrance.</figcaption></figure> <p>Cleptobiosis, also known as <a href="/enwiki/wiki/Kleptoparasitism" title="Kleptoparasitism">cleptoparasitism</a>, is a behaviour observed in various species of stingless bees, with over 30 identified species engaging in nest attacks, including honey bee nests. This behaviour serves the purpose of either resource theft or usurping the nest by swarming into an already occupied cavity and these bees are called robber bees. The Neotropical genus <i><a href="/enwiki/wiki/Lestrimelitta" title="Lestrimelitta">Lestrimelitta</a></i> and the African genus <i><a href="/enwiki/w/index.php?title=Cleptotrigona&amp;action=edit&amp;redlink=1" class="new" title="Cleptotrigona (page does not exist)">Cleptotrigona</a></i> represent bees with an obligate cleptobiotic lifestyle since they do not visit flowers for nectar or pollen. </p><p>Furthermore, other species such as <i><a href="/enwiki/w/index.php?title=Melipona_fuliginosa&amp;action=edit&amp;redlink=1" class="new" title="Melipona fuliginosa (page does not exist)">Melipona fuliginosa</a></i>, <i><a href="/enwiki/wiki/Oxytrigona_tataira" title="Oxytrigona tataira">Oxytrigona tataira</a>,</i> <i><a href="/enwiki/w/index.php?title=Trigona_hyalinata&amp;action=edit&amp;redlink=1" class="new" title="Trigona hyalinata (page does not exist)">Trigona hyalinata</a></i>, <i><a href="/enwiki/wiki/Trigona_spinipes" title="Trigona spinipes">T. spinipes</a></i>, and <i><a href="/enwiki/wiki/Tetragona_clavipes" title="Tetragona clavipes">Tetragona clavipes</a></i> are reported to have comparable habits of pillaging and invading, which emphasises the variety of strategies employed by stingless bees in acquiring resources. </p><p>Other enemies include: jumping spiders (<a href="/enwiki/wiki/Salticidae" class="mw-redirect" title="Salticidae">Salticidae</a>), moths, assassin bugs (<a href="/enwiki/wiki/Reduviidae" title="Reduviidae">Reduviidae</a>), beetles, parasitoid wasps, predatory mites (<i><a href="/enwiki/wiki/Amblyseius" title="Amblyseius">Amblyseius</a></i>), mantises (<a href="/enwiki/wiki/Mantodea" class="mw-redirect" title="Mantodea">Mantodea</a>), robber flies (<a href="/enwiki/wiki/Asilidae" title="Asilidae">Asilidae</a>), etc.<sup id="cite_ref-88" class="reference"><a href="#cite_note-88"><span class="cite-bracket">&#91;</span>88<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Vertebrates">Vertebrates</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=25" title="Edit section&#039;s source code: Vertebrates"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Human activities pose the most significant threat to stingless bees, whether through honey and nest removal, <a href="/enwiki/wiki/Habitat_destruction" title="Habitat destruction">habitat destruction</a>, <a href="/enwiki/wiki/Pesticide" title="Pesticide">pesticide</a> use or introduction of non-native competitors. Large-scale environmental alterations, particularly the conversion of natural habitats into urban or intensively <a href="/enwiki/wiki/Agriculture" title="Agriculture">farmed land</a>, are the most dramatic threats leading to habitat loss, reduced nest densities, and <a href="/enwiki/wiki/Extinction" title="Extinction">species disappearance</a>.<sup id="cite_ref-:18_89-0" class="reference"><a href="#cite_note-:18-89"><span class="cite-bracket">&#91;</span>89<span class="cite-bracket">&#93;</span></a></sup> </p><p><a href="/enwiki/wiki/Primate" title="Primate">Primates</a>, including <a href="/enwiki/wiki/Chimpanzee" title="Chimpanzee">chimpanzees</a>, <a href="/enwiki/wiki/Gorilla" title="Gorilla">gorillas</a>, <a href="/enwiki/wiki/Baboon" title="Baboon">baboons</a>, and various monkey species, are known to threaten stingless bee colonies. <a href="/enwiki/wiki/Elephant" title="Elephant">Elephants</a>, <a href="/enwiki/wiki/Honey_badger" title="Honey badger">honey badgers</a>, <a href="/enwiki/wiki/Sun_bear" title="Sun bear">sun bears</a>, <a href="/enwiki/wiki/Spectacled_bear" title="Spectacled bear">spectacled bears</a>, <a href="/enwiki/wiki/Anteater" title="Anteater">anteaters</a>, <a href="/enwiki/wiki/Hog-nosed_skunk" title="Hog-nosed skunk">hog-nosed skunks</a>, <a href="/enwiki/wiki/Armadillo" title="Armadillo">armadillos</a>, <a href="/enwiki/wiki/Tayra" title="Tayra">tayras</a>, <a href="/enwiki/wiki/Eyra_cat" class="mw-redirect" title="Eyra cat">eyra cats</a>, <a href="/enwiki/wiki/Kinkajou" title="Kinkajou">kinkajous</a>, <a href="/enwiki/wiki/Grisons" title="Grisons">grisons</a>, and <a href="/enwiki/wiki/Coyote" title="Coyote">coyotes</a> are among the mammals that consume or destroy stingless bee nests. Some, like the tayra and eyra cat, have specific preferences for stealing honey. <a href="/enwiki/wiki/Gecko" title="Gecko">Geckos</a>, <a href="/enwiki/wiki/Lizard" title="Lizard">lizards</a>, and <a href="/enwiki/wiki/Toad" title="Toad">toads</a> also pose threats by hunting adult bees or consuming workers at nest entrances. <a href="/enwiki/wiki/Woodpecker" title="Woodpecker">Woodpeckers</a> and various bird species, including <a href="/enwiki/wiki/Bee-eater" title="Bee-eater">bee-eaters</a>, <a href="/enwiki/wiki/Woodcreeper" title="Woodcreeper">woodcreepers</a>, <a href="/enwiki/wiki/Drongo" title="Drongo">drongos</a>, <a href="/enwiki/wiki/Jacamar" title="Jacamar">jacamars</a>, <a href="/enwiki/wiki/Heron" title="Heron">herons</a>, <a href="/enwiki/wiki/Kingbird" title="Kingbird">kingbirds</a>, <a href="/enwiki/wiki/Flycatcher-shrike" title="Flycatcher-shrike">flycatchers</a>, <a href="/enwiki/wiki/Swift_(bird)" title="Swift (bird)">swifts</a>, and <a href="/enwiki/wiki/Honeyeater" title="Honeyeater">honeyeaters</a>, occasionally prey on stingless bees. African <a href="/enwiki/wiki/Honeyguide" title="Honeyguide">honeyguides</a> have developed a mutualism with human honey-hunters, actively guiding them to bee nests for honey extraction and then consuming leftover wax and larvae.<sup id="cite_ref-:18_89-1" class="reference"><a href="#cite_note-:18-89"><span class="cite-bracket">&#91;</span>89<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Defense">Defense</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=26" title="Edit section&#039;s source code: Defense"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Being <a href="/enwiki/wiki/Tropics" title="Tropics">tropical</a>, stingless bees are active all year round, although they are less active in cooler weather, with some species presenting <a href="/enwiki/wiki/Diapause" title="Diapause">diapause</a>.<sup id="cite_ref-:19_90-0" class="reference"><a href="#cite_note-:19-90"><span class="cite-bracket">&#91;</span>90<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-91" class="reference"><a href="#cite_note-91"><span class="cite-bracket">&#91;</span>91<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-92" class="reference"><a href="#cite_note-92"><span class="cite-bracket">&#91;</span>92<span class="cite-bracket">&#93;</span></a></sup> Unlike other eusocial bees, they do not sting, but will defend by biting if their nest is disturbed. In addition, a few (in the genus <i><a href="/enwiki/wiki/Oxytrigona" title="Oxytrigona">Oxytrigona</a></i>) have mandibular secretions, including <a href="/enwiki/wiki/Formic_acid" title="Formic acid">formic acid</a>, that cause painful blisters.<sup id="cite_ref-:3_13-1" class="reference"><a href="#cite_note-:3-13"><span class="cite-bracket">&#91;</span>13<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-93" class="reference"><a href="#cite_note-93"><span class="cite-bracket">&#91;</span>93<span class="cite-bracket">&#93;</span></a></sup> Despite their lack of a <a href="/enwiki/wiki/Sting_(biology)" class="mw-redirect" title="Sting (biology)">sting</a>, stingless bees, being eusocial, may have very large colonies made formidable by the number of defenders.<sup id="cite_ref-94" class="reference"><a href="#cite_note-94"><span class="cite-bracket">&#91;</span>94<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-95" class="reference"><a href="#cite_note-95"><span class="cite-bracket">&#91;</span>95<span class="cite-bracket">&#93;</span></a></sup> </p><p>Stingless bees use other sophisticated defence tactics to protect their colonies and ensure their survival. One important strategy is to choose nesting habitats with fewer natural enemies to reduce the risk of attacks. In addition, they use <a href="/enwiki/wiki/Camouflage" title="Camouflage">camouflage</a> and <a href="/enwiki/wiki/Mimicry" title="Mimicry">mimicry</a> to blend into their surroundings or imitate other animals to avoid detection. An effective strategy is to nest near colonies that provide protection, using collective strength to defend against potential invaders.<sup id="cite_ref-:19_90-1" class="reference"><a href="#cite_note-:19-90"><span class="cite-bracket">&#91;</span>90<span class="cite-bracket">&#93;</span></a></sup> </p><p>Nest entrance guards play a vital role in colony defense by actively preventing unauthorized entry through attacking intruders and releasing alarm <a href="/enwiki/wiki/Pheromone" title="Pheromone">pheromones</a> to recruit additional defenders. It is worth noting that nest guards often carry sticky substances, such as resins and wax, in their corbiculae or mandibles. Stingless bees apply substances to attackers to immobilise them, thus thwarting potential threats to the colony. Some species (<i>Tetragonisca angustula</i> and <i>Nannotrigona testaceicornis</i>, for example) also close their nest entrances with a soft and porous layer of cerumen at night, further enhancing colony security during vulnerable periods. These intricate defence mechanisms demonstrate the adaptability and resilience of stingless bees in safeguarding their nests and resources.<sup id="cite_ref-:19_90-2" class="reference"><a href="#cite_note-:19-90"><span class="cite-bracket">&#91;</span>90<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Role_differentiation">Role differentiation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=27" title="Edit section&#039;s source code: Role differentiation"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In a simplified sense, the sex of each bee depends on the number of <a href="/enwiki/wiki/Chromosome" title="Chromosome">chromosomes</a> it receives. Female bees have two sets of chromosomes (<a href="/enwiki/wiki/Diploid" class="mw-redirect" title="Diploid">diploid</a>)—one set from the <a href="/enwiki/wiki/Queen_(bee)" class="mw-redirect" title="Queen (bee)">queen</a> and another from one of the male bees or <a href="/enwiki/wiki/Drone_(bee)" title="Drone (bee)">drones</a>. Drones have only one set of chromosomes (<a href="/enwiki/wiki/Haploid" class="mw-redirect" title="Haploid">haploid</a>), and are the result of unfertilized eggs, though inbreeding can result in diploid drones. </p><p>Unlike true honey bees, whose female bees may become workers or queens strictly depending on what kind of food they receive as larvae (queens are fed <a href="/enwiki/wiki/Royal_jelly" title="Royal jelly">royal jelly</a> and workers are fed pollen), the caste system in meliponines is variable, and commonly based simply on the amount of pollen consumed; larger amounts of pollen yield queens in the genus <i><a href="/enwiki/wiki/Melipona" title="Melipona">Melipona</a></i>. Also, a genetic component occurs, however, and as much as 25%<sup id="cite_ref-96" class="reference"><a href="#cite_note-96"><span class="cite-bracket">&#91;</span>96<span class="cite-bracket">&#93;</span></a></sup> (typically 5–14%) of the female brood may be queens. Queen cells in the former case can be distinguished from others by their larger size, as they are stocked with more pollen, but in the latter case, the cells are identical to worker cells, and scattered among the worker brood. When the new queens emerge, they typically leave to mate, and most die.<sup id="cite_ref-:28_97-0" class="reference"><a href="#cite_note-:28-97"><span class="cite-bracket">&#91;</span>97<span class="cite-bracket">&#93;</span></a></sup> New nests are not established by swarms, but by a procession of workers that gradually construct a new nest at a secondary location. The nest is then joined by a newly mated queen, at which point many workers take up permanent residence and help the new queen raise her own workers. If a ruling queen is herself weak or dying, then a new queen can replace her. For <i><a href="/enwiki/wiki/Plebeia_quadripunctata" class="mw-redirect" title="Plebeia quadripunctata">Schwarziana quadripunctata</a></i>, although fewer than 1% of female worker cells produce dwarf queens, they comprise six of seven queen bees, and one of five proceed to head colonies of their own. They are reproductively active, but less fecund than large queens.<sup id="cite_ref-:28_97-1" class="reference"><a href="#cite_note-:28-97"><span class="cite-bracket">&#91;</span>97<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Interaction_with_humans">Interaction with humans</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=28" title="Edit section&#039;s source code: Interaction with humans"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Pollination">Pollination</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=29" title="Edit section&#039;s source code: Pollination"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/enwiki/wiki/Pollination" title="Pollination">Pollination</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/enwiki/wiki/File:Bee_pollinating_a_rose.jpg" class="mw-file-description"><img src="/upwiki/wikipedia/commons/thumb/6/63/Bee_pollinating_a_rose.jpg/220px-Bee_pollinating_a_rose.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/6/63/Bee_pollinating_a_rose.jpg/330px-Bee_pollinating_a_rose.jpg 1.5x, /upwiki/wikipedia/commons/thumb/6/63/Bee_pollinating_a_rose.jpg/440px-Bee_pollinating_a_rose.jpg 2x" data-file-width="1024" data-file-height="768" /></a><figcaption>Solitary bee visiting and pollinating a flower.</figcaption></figure> <p>Bees play a critical role in the ecosystem, particularly in the pollination of natural vegetation. This activity is essential for the reproduction of various plant species, particularly in <a href="/enwiki/wiki/Tropical_forest" title="Tropical forest">tropical forests</a> where most tree species rely on <a href="/enwiki/wiki/Entomophily" title="Entomophily">insect pollination</a>. Even in temperate climates, where <a href="/enwiki/wiki/Anemophily" title="Anemophily">wind pollination</a> is prevalent among forest trees, many bushes and herbaceous plants, rely on bees for pollination. The significance of bees extends to arid regions, such as <a href="/enwiki/wiki/Deserts_and_xeric_shrublands" title="Deserts and xeric shrublands">desertic and xeric shrublands</a>, where bee-pollinated plants are essential for preventing <a href="/enwiki/wiki/Erosion" title="Erosion">erosion</a>, supporting <a href="/enwiki/wiki/Wildlife" title="Wildlife">wildlife</a>, and ensuring <a href="/enwiki/wiki/Ecological_stability" title="Ecological stability">ecosystem stability</a>.<sup id="cite_ref-:23_98-0" class="reference"><a href="#cite_note-:23-98"><span class="cite-bracket">&#91;</span>98<span class="cite-bracket">&#93;</span></a></sup> </p><p>The impact of bee pollination on agriculture is substantial. In the late 1980s, certain plants were estimated to contribute between $4.6 to $18.9 billion to the U.S. economy, primarily through insect-pollinated crops. Although some bee-pollinated plants can self-pollinate in the absence of bees, the resulting crops often suffer from inbreeding depression. The quality and quantity of seeds or fruits are significantly enhanced when bees participate in the pollination process. Although estimates of crop pollination attributed to honey bees are uncertain, it is undeniable that bee pollination is a vital and economically valuable activity.<sup id="cite_ref-:23_98-1" class="reference"><a href="#cite_note-:23-98"><span class="cite-bracket">&#91;</span>98<span class="cite-bracket">&#93;</span></a></sup> </p><p>Ramalho (2004) demonstrates that stingless bees amount to approximately 70% of all bees foraging on flowers in the Brazilian Tropical <a href="/enwiki/wiki/Atlantic_Forest" title="Atlantic Forest">Atlantic Rainforest</a> even though they represented only 7% of all bee species.<sup id="cite_ref-99" class="reference"><a href="#cite_note-99"><span class="cite-bracket">&#91;</span>99<span class="cite-bracket">&#93;</span></a></sup> In a habitat in Costa Rica, stingless bees accounted for 50% of the observed foraging bees, despite representing only 16% of the recorded bee species.<sup id="cite_ref-100" class="reference"><a href="#cite_note-100"><span class="cite-bracket">&#91;</span>100<span class="cite-bracket">&#93;</span></a></sup> Following this pattern, Cairns <i>et al</i>. (2005) found that 52% of all bees visiting flowers in Mexican habitats were meliponines.<sup id="cite_ref-101" class="reference"><a href="#cite_note-101"><span class="cite-bracket">&#91;</span>101<span class="cite-bracket">&#93;</span></a></sup> </p><p>Meliponine bees play a crucial role in tropical environments due to their high population rate, morphological diversity, diverse foraging strategies, generalist foraging habits (<a href="/enwiki/wiki/Polylectic" class="mw-redirect" title="Polylectic">polylecty</a>), and flower constancy during foraging trips. Nest densities and colony sizes can result in over a million individual stingless bees inhabiting a square kilometre of tropical habitat. Due to their diverse morphology and behaviour, bees are capable of collecting pollen and nectar from a wide range of flowering plants. Key plant families are reported as most visited by meliponines: <a href="/enwiki/wiki/Fabaceae" title="Fabaceae">Fabaceae</a>, <a href="/enwiki/wiki/Euphorbiaceae" title="Euphorbiaceae">Euphorbiaceae</a>, <a href="/enwiki/wiki/Asteraceae" title="Asteraceae">Asteraceae</a> and <a href="/enwiki/wiki/Myrtaceae" title="Myrtaceae">Myrtaceae</a>.<sup id="cite_ref-:20_102-0" class="reference"><a href="#cite_note-:20-102"><span class="cite-bracket">&#91;</span>102<span class="cite-bracket">&#93;</span></a></sup> </p><p>Grüter compiled some studies about twenty crops that substantially benefit from SB pollination (following table) and also lists seventy-four crops that are at least occasionally or potentially pollinated by stingless bees.<sup id="cite_ref-:20_102-1" class="reference"><a href="#cite_note-:20-102"><span class="cite-bracket">&#91;</span>102<span class="cite-bracket">&#93;</span></a></sup> </p> <table class="wikitable mw-collapsible"> <caption> </caption> <tbody><tr> <th>Common name </th> <th>Scientific name </th> <th>Family </th> <th>Pollinator genus </th> <th>Reference </th></tr> <tr> <td>Annatto, achiote </td> <td><i><a href="/enwiki/wiki/Bixa_orellana" title="Bixa orellana">Bixa orellana</a></i> </td> <td><a href="/enwiki/wiki/Bixaceae" title="Bixaceae">Bixaceae</a> </td> <td><i><a href="/enwiki/wiki/Melipona" title="Melipona">Melipona</a></i> </td> <td><sup id="cite_ref-:21_103-0" class="reference"><a href="#cite_note-:21-103"><span class="cite-bracket">&#91;</span>103<span class="cite-bracket">&#93;</span></a></sup> </td></tr> <tr> <td>Aubergine </td> <td><i><a href="/enwiki/wiki/Solanum_melongena" class="mw-redirect" title="Solanum melongena">Solanum melogena</a></i> </td> <td><a href="/enwiki/wiki/Solanaceae" title="Solanaceae">Solanaceae</a> </td> <td><i>Melipona</i> </td> <td><sup id="cite_ref-104" class="reference"><a href="#cite_note-104"><span class="cite-bracket">&#91;</span>104<span class="cite-bracket">&#93;</span></a></sup> </td></tr> <tr> <td>Avocado </td> <td><i><a href="/enwiki/wiki/Persea_americana" class="mw-redirect" title="Persea americana">Persea americana</a></i> </td> <td><a href="/enwiki/wiki/Lauraceae" title="Lauraceae">Lauraceae</a> </td> <td><i><a href="/enwiki/wiki/Nannotrigona" title="Nannotrigona">Nannotrigona</a>, <a href="/enwiki/wiki/Trigona" title="Trigona">Trigona</a></i> </td> <td><sup id="cite_ref-:21_103-1" class="reference"><a href="#cite_note-:21-103"><span class="cite-bracket">&#91;</span>103<span class="cite-bracket">&#93;</span></a></sup> </td></tr> <tr> <td>Camu-camu </td> <td><i><a href="/enwiki/wiki/Myrciaria_dubia" title="Myrciaria dubia">Myrciaria dubia</a></i> </td> <td><a href="/enwiki/wiki/Myrtaceae" title="Myrtaceae">Myrtaceae</a> </td> <td><i>Melipona, <a href="/enwiki/wiki/Scaptotrigona" title="Scaptotrigona">Scaptotrigona</a></i> </td> <td><sup id="cite_ref-:21_103-2" class="reference"><a href="#cite_note-:21-103"><span class="cite-bracket">&#91;</span>103<span class="cite-bracket">&#93;</span></a></sup> </td></tr> <tr> <td>Carambola </td> <td><i><a href="/enwiki/wiki/Averrhoa_carambola" title="Averrhoa carambola">Averrhoa carambola</a></i> </td> <td><a href="/enwiki/wiki/Oxalidaceae" title="Oxalidaceae">Oxalidaceae</a> </td> <td><i>Trigona</i> </td> <td><sup id="cite_ref-:21_103-3" class="reference"><a href="#cite_note-:21-103"><span class="cite-bracket">&#91;</span>103<span class="cite-bracket">&#93;</span></a></sup> </td></tr> <tr> <td>Chayote, choko </td> <td><i><a href="/enwiki/wiki/Sechium_edule" class="mw-redirect" title="Sechium edule">Sechium edule</a></i> </td> <td><a href="/enwiki/wiki/Cucurbitaceae" title="Cucurbitaceae">Cucubitaceae</a> </td> <td><i>Trigona, <a href="/enwiki/wiki/Partamona" title="Partamona">Partamona</a></i> </td> <td><sup id="cite_ref-:21_103-4" class="reference"><a href="#cite_note-:21-103"><span class="cite-bracket">&#91;</span>103<span class="cite-bracket">&#93;</span></a></sup> </td></tr> <tr> <td>Coconut </td> <td><i><a href="/enwiki/wiki/Cocos_nucifera" class="mw-redirect" title="Cocos nucifera">Cocos nucifera</a></i> </td> <td><a href="/enwiki/wiki/Arecaceae" title="Arecaceae">Arecaceae</a> </td> <td>various genera </td> <td><sup id="cite_ref-:21_103-5" class="reference"><a href="#cite_note-:21-103"><span class="cite-bracket">&#91;</span>103<span class="cite-bracket">&#93;</span></a></sup> </td></tr> <tr> <td>Coffee </td> <td><i><a href="/enwiki/wiki/Coffea_arabica" title="Coffea arabica">Coffea arabica</a></i> </td> <td><a href="/enwiki/wiki/Rubiaceae" title="Rubiaceae">Rubiaceae</a> </td> <td><i><a href="/enwiki/w/index.php?title=Lepidotrigona&amp;action=edit&amp;redlink=1" class="new" title="Lepidotrigona (page does not exist)">Lepidotrigona</a>, Trigona</i> </td> <td><sup id="cite_ref-:21_103-6" class="reference"><a href="#cite_note-:21-103"><span class="cite-bracket">&#91;</span>103<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:22_105-0" class="reference"><a href="#cite_note-:22-105"><span class="cite-bracket">&#91;</span>105<span class="cite-bracket">&#93;</span></a></sup> </td></tr> <tr> <td>Coffee </td> <td><i><a href="/enwiki/wiki/Coffea_canephora" title="Coffea canephora">Coffea canephora</a></i> </td> <td>Rubiaceae </td> <td><i>Lepidotrigona, Trigona</i> </td> <td><sup id="cite_ref-:22_105-1" class="reference"><a href="#cite_note-:22-105"><span class="cite-bracket">&#91;</span>105<span class="cite-bracket">&#93;</span></a></sup> </td></tr> <tr> <td>Cucumber </td> <td><i><a href="/enwiki/wiki/Cucumis_sativus" class="mw-redirect" title="Cucumis sativus">Cucumis sativus</a></i> </td> <td>Curcubitaceae </td> <td><i>Nannotrigona, Scaptotrigona</i> </td> <td><sup id="cite_ref-:21_103-7" class="reference"><a href="#cite_note-:21-103"><span class="cite-bracket">&#91;</span>103<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:22_105-2" class="reference"><a href="#cite_note-:22-105"><span class="cite-bracket">&#91;</span>105<span class="cite-bracket">&#93;</span></a></sup> </td></tr> <tr> <td>Cupuaçu </td> <td><i><a href="/enwiki/wiki/Theobroma_grandiflorum" title="Theobroma grandiflorum">Theobroma grandiflorum</a></i> </td> <td><a href="/enwiki/wiki/Malvaceae" title="Malvaceae">Malvaceae</a> </td> <td><i>Trigona</i> </td> <td><sup id="cite_ref-:21_103-8" class="reference"><a href="#cite_note-:21-103"><span class="cite-bracket">&#91;</span>103<span class="cite-bracket">&#93;</span></a></sup> </td></tr> <tr> <td>Macadamia </td> <td><i><a href="/enwiki/wiki/Macadamia_integrifolia" title="Macadamia integrifolia">Macadamia integrifolia</a></i> </td> <td><a href="/enwiki/wiki/Proteaceae" title="Proteaceae">Proteaceae</a> </td> <td><i>Trigona</i> </td> <td><sup id="cite_ref-:21_103-9" class="reference"><a href="#cite_note-:21-103"><span class="cite-bracket">&#91;</span>103<span class="cite-bracket">&#93;</span></a></sup> </td></tr> <tr> <td>Mango </td> <td><i><a href="/enwiki/wiki/Mangifera_indica" title="Mangifera indica">Mangifera indica</a></i> </td> <td><a href="/enwiki/wiki/Anacardiaceae" title="Anacardiaceae">Anacardiaceae</a> </td> <td><i>Trigona</i> </td> <td><sup id="cite_ref-:21_103-10" class="reference"><a href="#cite_note-:21-103"><span class="cite-bracket">&#91;</span>103<span class="cite-bracket">&#93;</span></a></sup> </td></tr> <tr> <td>Mapati, uvilla </td> <td><i><a href="/enwiki/wiki/Pourouma_cecropiifolia" title="Pourouma cecropiifolia">Pourouma cecropiifolia</a></i> </td> <td><a href="/enwiki/wiki/Moraceae" title="Moraceae">Moraceae</a> </td> <td><i><a href="/enwiki/wiki/Oxytrigona" title="Oxytrigona">Oxytrigona</a>, Trigona</i> </td> <td><sup id="cite_ref-:21_103-11" class="reference"><a href="#cite_note-:21-103"><span class="cite-bracket">&#91;</span>103<span class="cite-bracket">&#93;</span></a></sup> </td></tr> <tr> <td>Mealy sage </td> <td><i><a href="/enwiki/wiki/Salvia_farinacea" title="Salvia farinacea">Salvia farinacea</a></i> </td> <td><a href="/enwiki/wiki/Lamiaceae" title="Lamiaceae">Lamiaceae</a> </td> <td><i>Nannotrigona, <a href="/enwiki/wiki/Tetragonisca" title="Tetragonisca">Tetragonisca</a></i> </td> <td><sup id="cite_ref-:22_105-3" class="reference"><a href="#cite_note-:22-105"><span class="cite-bracket">&#91;</span>105<span class="cite-bracket">&#93;</span></a></sup> </td></tr> <tr> <td>Rambutan </td> <td><i><a href="/enwiki/wiki/Nephelium_lappaceum" class="mw-redirect" title="Nephelium lappaceum">Nephelium lappaceum</a></i> </td> <td><a href="/enwiki/wiki/Sapindaceae" title="Sapindaceae">Sapindaceae</a> </td> <td><i>Scaptotrigona</i> </td> <td><sup id="cite_ref-:21_103-12" class="reference"><a href="#cite_note-:21-103"><span class="cite-bracket">&#91;</span>103<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:22_105-4" class="reference"><a href="#cite_note-:22-105"><span class="cite-bracket">&#91;</span>105<span class="cite-bracket">&#93;</span></a></sup> </td></tr> <tr> <td>Rockmelon </td> <td><i><a href="/enwiki/wiki/Cucumis_melo" title="Cucumis melo">Cucumis melo</a></i> </td> <td>Curcubitaceae </td> <td><i><a href="/enwiki/w/index.php?title=Heterotrigona&amp;action=edit&amp;redlink=1" class="new" title="Heterotrigona (page does not exist)">Heterotrigona</a></i> </td> <td><sup id="cite_ref-106" class="reference"><a href="#cite_note-106"><span class="cite-bracket">&#91;</span>106<span class="cite-bracket">&#93;</span></a></sup> </td></tr> <tr> <td>Strawberry </td> <td><i><a href="/enwiki/wiki/Fragaria" title="Fragaria">Fragaria</a></i> sp. </td> <td><a href="/enwiki/wiki/Rosaceae" title="Rosaceae">Rosaceae</a> </td> <td>various genera </td> <td><sup id="cite_ref-:21_103-13" class="reference"><a href="#cite_note-:21-103"><span class="cite-bracket">&#91;</span>103<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:22_105-5" class="reference"><a href="#cite_note-:22-105"><span class="cite-bracket">&#91;</span>105<span class="cite-bracket">&#93;</span></a></sup> </td></tr> <tr> <td>Sweet pepper </td> <td><i><a href="/enwiki/wiki/Capsicum_annuum" title="Capsicum annuum">Capsicum annuum</a></i> </td> <td>Solanaceae </td> <td><i><a href="/enwiki/wiki/Austroplebeia" title="Austroplebeia">Austroplebeia</a>, Melipona, <a href="/enwiki/wiki/Tetragonula" title="Tetragonula">Tetragonula</a></i> </td> <td><sup id="cite_ref-:21_103-14" class="reference"><a href="#cite_note-:21-103"><span class="cite-bracket">&#91;</span>103<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:22_105-6" class="reference"><a href="#cite_note-:22-105"><span class="cite-bracket">&#91;</span>105<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-107" class="reference"><a href="#cite_note-107"><span class="cite-bracket">&#91;</span>107<span class="cite-bracket">&#93;</span></a></sup> </td></tr> <tr> <td>Tomato </td> <td><i><a href="/enwiki/wiki/Lycopersicon_esculentum" class="mw-redirect" title="Lycopersicon esculentum">Solanum lycopersicum</a></i> </td> <td>Solanaceae </td> <td><i>Melipona, Nannotrigona</i> </td> <td><sup id="cite_ref-:22_105-7" class="reference"><a href="#cite_note-:22-105"><span class="cite-bracket">&#91;</span>105<span class="cite-bracket">&#93;</span></a></sup> </td></tr></tbody></table> <div class="mw-heading mw-heading2"><h2 id="Worldwide_overview">Worldwide overview</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=30" title="Edit section&#039;s source code: Worldwide overview"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Africa">Africa</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=31" title="Edit section&#039;s source code: Africa"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Stingless bees also play a vital ecological role across <a href="/enwiki/wiki/Sub-Saharan_Africa" title="Sub-Saharan Africa">Sub-Saharan Africa</a> and <a href="/enwiki/wiki/Madagascar" title="Madagascar">Madagascar</a>. To understand these insects on the African continent, it's important to consider the prevailing socio-economic and cultural contexts. Despite their ecological significance, the diversity, conservation, and behavior of these bees remain underexplored, particularly compared to better-studied regions such as <a href="/enwiki/wiki/South_America" title="South America">South America</a> and <a href="/enwiki/wiki/Southeast_Asia" title="Southeast Asia">Southeast Asia</a>. Also, honeybees were extensively researched, in contrast to native meliponines.<sup id="cite_ref-:31_108-0" class="reference"><a href="#cite_note-:31-108"><span class="cite-bracket">&#91;</span>108<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:32_109-0" class="reference"><a href="#cite_note-:32-109"><span class="cite-bracket">&#91;</span>109<span class="cite-bracket">&#93;</span></a></sup> </p><p>Africa is home to seven <a href="/enwiki/wiki/Biodiversity_hotspot" title="Biodiversity hotspot">biodiversity hotspots</a>, yet the recorded bee fauna is moderate relative to the continent's size. Madagascar stands out with exceptionally high levels of endemic species, though much of the bee diversity remains undocumented.<sup id="cite_ref-:30_110-0" class="reference"><a href="#cite_note-:30-110"><span class="cite-bracket">&#91;</span>110<span class="cite-bracket">&#93;</span></a></sup><b><sup id="cite_ref-:31_108-1" class="reference"><a href="#cite_note-:31-108"><span class="cite-bracket">&#91;</span>108<span class="cite-bracket">&#93;</span></a></sup></b> Africa is home to aproximately 36 species of meliponines, including seven endemic to Madagascar. Most of these bees are found in equatorian regions (tropical forests and some savannahs).<sup id="cite_ref-111" class="reference"><a href="#cite_note-111"><span class="cite-bracket">&#91;</span>111<span class="cite-bracket">&#93;</span></a></sup> </p><p>Factors such as habitat destruction, pesticide use, and invasive species pose significant threats to these pollinators. Furthermore, high rates of nest mortality, driven by predation and human activity, exacerbate conservation challenges. Research indicates that stingless bees in Africa face greater pressures than their counterparts in the American and Asian tropics, underlining the urgency for targeted conservation measures.<sup id="cite_ref-:30_110-1" class="reference"><a href="#cite_note-:30-110"><span class="cite-bracket">&#91;</span>110<span class="cite-bracket">&#93;</span></a></sup><b><sup id="cite_ref-:31_108-2" class="reference"><a href="#cite_note-:31-108"><span class="cite-bracket">&#91;</span>108<span class="cite-bracket">&#93;</span></a></sup></b> </p><p>Uganda's <a href="/enwiki/wiki/Bwindi_Impenetrable_National_Park" title="Bwindi Impenetrable National Park">Bwindi Impenetrable National Park</a> has shown the presence of at least five stingless bee species in, distributed across two genera: <i><a href="/enwiki/wiki/Meliponula" title="Meliponula">Meliponula</a></i> and <i><a href="/enwiki/w/index.php?title=Hypotrigona&amp;action=edit&amp;redlink=1" class="new" title="Hypotrigona (page does not exist)">Hypotrigona</a></i>.<sup id="cite_ref-:31_108-3" class="reference"><a href="#cite_note-:31-108"><span class="cite-bracket">&#91;</span>108<span class="cite-bracket">&#93;</span></a></sup><b><sup id="cite_ref-:32_109-1" class="reference"><a href="#cite_note-:32-109"><span class="cite-bracket">&#91;</span>109<span class="cite-bracket">&#93;</span></a></sup></b> In Madagascar, there is only one genus of stingless bees: <i><a href="/enwiki/w/index.php?title=Liotrigona&amp;action=edit&amp;redlink=1" class="new" title="Liotrigona (page does not exist)">Liotrigona</a>.</i><sup id="cite_ref-:30_110-2" class="reference"><a href="#cite_note-:30-110"><span class="cite-bracket">&#91;</span>110<span class="cite-bracket">&#93;</span></a></sup> </p><p>Meliponiculture, for example, is practised in Angola and Tanzania, and interest in managing stingless bees is growing in other African countries as well. <sup id="cite_ref-112" class="reference"><a href="#cite_note-112"><span class="cite-bracket">&#91;</span>112<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Australia">Australia</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=32" title="Edit section&#039;s source code: Australia"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Of the 1,600 species of wild bees native to Australia, about 14 are meliponines.<sup id="cite_ref-113" class="reference"><a href="#cite_note-113"><span class="cite-bracket">&#91;</span>113<span class="cite-bracket">&#93;</span></a></sup> "<i>Coot-tha</i>", which derives from "<i>ku-ta</i>", is one of the Aboriginal names for "wild stingless bee honey".<sup id="cite_ref-114" class="reference"><a href="#cite_note-114"><span class="cite-bracket">&#91;</span>114<span class="cite-bracket">&#93;</span></a></sup> These species bear a variety of names, including Australian native honey bees, native bees, sugar-bag bees, and sweat bees (because they land on people's skin to collect sweat).<sup id="cite_ref-115" class="reference"><a href="#cite_note-115"><span class="cite-bracket">&#91;</span>115<span class="cite-bracket">&#93;</span></a></sup> The various stingless species look quite similar, with the two most common species, <i><a href="/enwiki/wiki/Tetragonula_carbonaria" title="Tetragonula carbonaria">Tetragonula carbonaria</a></i> and <i><a href="/enwiki/wiki/Austroplebeia_australis" title="Austroplebeia australis">Austroplebeia australis</a></i>, displaying the greatest variation, as the latter is smaller and less active. Both of these inhabit the area around <a href="/enwiki/wiki/Brisbane" title="Brisbane">Brisbane</a>.<sup id="cite_ref-116" class="reference"><a href="#cite_note-116"><span class="cite-bracket">&#91;</span>116<span class="cite-bracket">&#93;</span></a></sup> </p><p>As stingless bees are usually harmless to humans, they have become an increasingly attractive addition to the suburban backyard. Most meliponine beekeepers do not keep the bees for <a href="/enwiki/wiki/Honey" title="Honey">honey</a>, but rather for the pleasure of conserving native species whose original <a href="/enwiki/wiki/Habitat_(ecology)" class="mw-redirect" title="Habitat (ecology)">habitat</a> is declining due to human development. In return, the bees <a href="/enwiki/wiki/Pollinate" class="mw-redirect" title="Pollinate">pollinate</a> crops, garden flowers, and bushland during their search for nectar and pollen. While a number of beekeepers fill a small niche market for bush honey, native meliponines only produce small amounts and the structure of their hives makes the honey difficult to extract. Only in warm areas of Australia such as <a href="/enwiki/wiki/Queensland" title="Queensland">Queensland</a> and northern <a href="/enwiki/wiki/New_South_Wales" title="New South Wales">New South Wales</a> are favorable for these bees to produce more honey than they need for their own survival. Most bees only come out of the hive when it is above about 18°C (64°F).<sup id="cite_ref-117" class="reference"><a href="#cite_note-117"><span class="cite-bracket">&#91;</span>117<span class="cite-bracket">&#93;</span></a></sup> Harvesting honey from a nest in a cooler area could weaken or even kill the nest. </p> <div class="mw-heading mw-heading4"><h4 id="Pollination_2">Pollination</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=33" title="Edit section&#039;s source code: Pollination"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Australian farmers rely almost exclusively on the introduced <a href="/enwiki/wiki/Western_honey_bee" title="Western honey bee">western honey bee</a> to pollinate their crops. However, native bees may be better pollinators for certain agricultural crops. Stingless bees have been shown to be valuable pollinators of tropical plants such as <a href="/enwiki/wiki/Macadamia" title="Macadamia">macadamias</a> and <a href="/enwiki/wiki/Mango" title="Mango">mangos</a>.<sup id="cite_ref-:21_103-15" class="reference"><a href="#cite_note-:21-103"><span class="cite-bracket">&#91;</span>103<span class="cite-bracket">&#93;</span></a></sup> Their foraging may also benefit strawberries, watermelons, citrus, avocados, <a href="/enwiki/wiki/Lychee" title="Lychee">lychees</a>, and many others.<sup id="cite_ref-:21_103-16" class="reference"><a href="#cite_note-:21-103"><span class="cite-bracket">&#91;</span>103<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:22_105-8" class="reference"><a href="#cite_note-:22-105"><span class="cite-bracket">&#91;</span>105<span class="cite-bracket">&#93;</span></a></sup> Research into the use of stingless bees for crop pollination in Australia is still in its very early stages, but these bees show great potential. Studies at the <a href="/enwiki/wiki/University_of_Western_Sydney" class="mw-redirect" title="University of Western Sydney">University of Western Sydney</a> have shown these bees are effective pollinators even in confined areas, such as glasshouses.<sup id="cite_ref-118" class="reference"><a href="#cite_note-118"><span class="cite-bracket">&#91;</span>118<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Brazil">Brazil</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=34" title="Edit section&#039;s source code: Brazil"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/enwiki/wiki/List_of_stingless_bees_of_Brazil" title="List of stingless bees of Brazil">List of stingless bees of Brazil</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/enwiki/wiki/File:Abelhas_sem_ferrao_brasil.png" class="mw-file-description"><img src="/upwiki/wikipedia/commons/thumb/1/18/Abelhas_sem_ferrao_brasil.png/220px-Abelhas_sem_ferrao_brasil.png" decoding="async" width="220" height="184" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/1/18/Abelhas_sem_ferrao_brasil.png/330px-Abelhas_sem_ferrao_brasil.png 1.5x, /upwiki/wikipedia/commons/thumb/1/18/Abelhas_sem_ferrao_brasil.png/440px-Abelhas_sem_ferrao_brasil.png 2x" data-file-width="738" data-file-height="618" /></a><figcaption>Number of valid species of stingless bees in Brazil. State and region cut-outs. Study published in 2023.<sup id="cite_ref-:27_2-1" class="reference"><a href="#cite_note-:27-2"><span class="cite-bracket">&#91;</span>2<span class="cite-bracket">&#93;</span></a></sup></figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/enwiki/wiki/File:Pollinating_bee_covered_with_pollen.jpg" class="mw-file-description"><img src="/upwiki/wikipedia/commons/thumb/7/7d/Pollinating_bee_covered_with_pollen.jpg/220px-Pollinating_bee_covered_with_pollen.jpg" decoding="async" width="220" height="196" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/7/7d/Pollinating_bee_covered_with_pollen.jpg/330px-Pollinating_bee_covered_with_pollen.jpg 1.5x, /upwiki/wikipedia/commons/thumb/7/7d/Pollinating_bee_covered_with_pollen.jpg/440px-Pollinating_bee_covered_with_pollen.jpg 2x" data-file-width="1885" data-file-height="1677" /></a><figcaption><i><a href="/enwiki/wiki/Trigona_spinipes" title="Trigona spinipes">Trigona spinipes</a>,</i> covered with <a href="/enwiki/wiki/Pollen" title="Pollen">pollen</a>, visiting a flower of the vegetable sponge gourd (<i>Luffa cylindrica</i>) in <a href="/enwiki/wiki/Campinas" title="Campinas">Campinas</a>, <a href="/enwiki/wiki/Brazil" title="Brazil">Brazil</a>.</figcaption></figure> <p>Brazil is home to several species bees belonging to Meliponini, with more than 300 species already identified and probably more yet to be discovered and described. They vary greatly in shape, size, and habits, and 20 to 30 of these species have good potential as honey producers. Although they are still quite unknown by most people, an increasing number of beekeepers (<i>meliponicultores</i>, in Portuguese) have been dedicated to these bees throughout the country.<sup id="cite_ref-:27_2-2" class="reference"><a href="#cite_note-:27-2"><span class="cite-bracket">&#91;</span>2<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-119" class="reference"><a href="#cite_note-119"><span class="cite-bracket">&#91;</span>119<span class="cite-bracket">&#93;</span></a></sup> This activity has experienced significant growth since August 2004, when national laws were changed to allow native bee colonies to be freely marketed, which was previously forbidden in an unsuccessful attempt to protect these species. Nowadays the capture or destruction of existing colonies in nature is still forbidden, and only new colonies formed by the bees themselves in artificial traps can be collected from the wild.<sup id="cite_ref-120" class="reference"><a href="#cite_note-120"><span class="cite-bracket">&#91;</span>120<span class="cite-bracket">&#93;</span></a></sup> Most marketed colonies are artificially produced by authorized beekeepers, through division of already existing captive colonies. Besides honey production, Brazilian stingless bees such as the jataí (<i><a href="/enwiki/wiki/Tetragonisca_angustula" title="Tetragonisca angustula">Tetragonisca angustula</a></i>), mandaguari (<i><a href="/enwiki/wiki/Scaptotrigona_postica" title="Scaptotrigona postica">Scaptotrigona postica</a></i>), and mandaçaia (<i><a href="/enwiki/wiki/Melipona_quadrifasciata" title="Melipona quadrifasciata">Melipona quadrifasciata</a></i>) serve as major pollinators of tropical plants and are considered the ecological equivalent of the honey bee.<sup id="cite_ref-:21_103-17" class="reference"><a href="#cite_note-:21-103"><span class="cite-bracket">&#91;</span>103<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:22_105-9" class="reference"><a href="#cite_note-:22-105"><span class="cite-bracket">&#91;</span>105<span class="cite-bracket">&#93;</span></a></sup> </p><p>Also, much practical and academic work is being done about the best ways of keeping such bees, multiplying their colonies, and exploring the honey they produce.<sup id="cite_ref-121" class="reference"><a href="#cite_note-121"><span class="cite-bracket">&#91;</span>121<span class="cite-bracket">&#93;</span></a></sup> Among many others, species such as jandaíra (<i><a href="/enwiki/wiki/Melipona_subnitida" title="Melipona subnitida">Melipona subnitida</a></i>) and true <i>uruçu</i> (<i><a href="/enwiki/wiki/Melipona_scutellaris" title="Melipona scutellaris">Melipona scutellaris</a></i>) in the northeast of the country, mandaçaia (<i><a href="/enwiki/wiki/Melipona_quadrifasciata" title="Melipona quadrifasciata">Melipona quadrifasciata</a></i>) and yellow uruçu (<i>Melipona rufiventris</i>) in the south-southeast, tiúba or jupará (<i><a href="/enwiki/wiki/Melipona_interrupta" title="Melipona interrupta">Melipona interrupta</a></i>) and canudo (<i><a href="/enwiki/wiki/Scaptotrigona_polysticta" title="Scaptotrigona polysticta">Scaptotrigona polysticta</a></i>) in the north and jataí (<i><a href="/enwiki/wiki/Tetragonisca_angustula" title="Tetragonisca angustula">Tetragonisca angustula</a></i>) throughout the country are increasingly kept by small, medium, and large producers. Many other species as the mandaguari (<i><a href="/enwiki/wiki/Scaptotrigona_postica" title="Scaptotrigona postica">Scaptotrigona postica</a></i>), the guaraipo (<i><a href="/enwiki/wiki/Melipona_bicolor" title="Melipona bicolor">Melipona bicolor</a></i>), marmelada (<i><a href="/enwiki/wiki/Frieseomelitta_varia" title="Frieseomelitta varia">Frieseomelitta varia</a></i>) and the iraí (<i><a href="/enwiki/wiki/Nannotrigona_testaceicornis" title="Nannotrigona testaceicornis">Nannotrigona testaceicornis</a></i>), to mention a few, are also reared.<sup id="cite_ref-:33_122-0" class="reference"><a href="#cite_note-:33-122"><span class="cite-bracket">&#91;</span>122<span class="cite-bracket">&#93;</span></a></sup> </p><p>According to <a href="/enwiki/wiki/Chico_Mendes_Institute_for_Biodiversity_Conservation" title="Chico Mendes Institute for Biodiversity Conservation">ICMBio</a> and the <a href="/enwiki/wiki/Ministry_of_Environment_and_Climate_Change_(Brazil)" title="Ministry of Environment and Climate Change (Brazil)">Ministry of the Environment</a> there are presently four species of Meliponini listed in the <a href="/enwiki/wiki/IUCN_Red_List" title="IUCN Red List">National Red List of Threatened Species</a> in Brazil. <i><a href="/enwiki/wiki/Melipona_capixaba" title="Melipona capixaba">Melipona capixaba</a></i>, <i><a href="/enwiki/wiki/Melipona_rufiventris" title="Melipona rufiventris">Melipona rufiventris</a></i>, <i><a href="/enwiki/wiki/Melipona_scutellaris" title="Melipona scutellaris">Melipona scutellaris</a></i>, and <i><a href="/enwiki/w/index.php?title=Partamona_littoralis&amp;action=edit&amp;redlink=1" class="new" title="Partamona littoralis (page does not exist)">Partamona littoralis</a></i> all listed as Endangered (EN).<sup id="cite_ref-123" class="reference"><a href="#cite_note-123"><span class="cite-bracket">&#91;</span>123<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-124" class="reference"><a href="#cite_note-124"><span class="cite-bracket">&#91;</span>124<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Honey_production">Honey production</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=35" title="Edit section&#039;s source code: Honey production"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Although the colony population of most of these bees is much smaller than that of European bees, the productivity per bee can be quite high. Interestingly, honey production is more connected to the body size, not the colony size. The manduri (<i><a href="/enwiki/w/index.php?title=Melipona_marginata&amp;action=edit&amp;redlink=1" class="new" title="Melipona marginata (page does not exist)">Melipona marginata</a></i>), jandaíra (<i><a href="/enwiki/wiki/Melipona_subnitida" title="Melipona subnitida">Melipona subnitida</a></i>) and the <i>guaraipo</i> (<i><a href="/enwiki/wiki/Melipona_bicolor" title="Melipona bicolor">M. bicolor</a></i>) live in swarms of only around 300 individuals but can still produce up to 5 liters (1.3 US gallon) of honey a year under the right conditions.<sup id="cite_ref-125" class="reference"><a href="#cite_note-125"><span class="cite-bracket">&#91;</span>125<span class="cite-bracket">&#93;</span></a></sup> In large bee farms, only the availability of flowers limits the honey production per colony. However, much larger numbers of beehives are required to produce amounts of honey comparable to that of European bees. Also, due to the fact of those bees storing honey in cerumen pots instead of standardized honeycombs as in the honeybee rearing makes extraction a lot more difficult and laborious.<sup id="cite_ref-126" class="reference"><a href="#cite_note-126"><span class="cite-bracket">&#91;</span>126<span class="cite-bracket">&#93;</span></a></sup> </p><p>The honey from stingless bees has a higher water content, from 25% to 35%, compared to the honey from the genus <i>Apis</i>. This contributes to its less cloying taste but also causes it to spoil more easily. Thus, for marketing, this honey needs to be processed through desiccation, fermentation or pasteurization. In its natural state, it should be kept under refrigeration.<sup id="cite_ref-127" class="reference"><a href="#cite_note-127"><span class="cite-bracket">&#91;</span>127<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Bees_as_pets">Bees as pets</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=36" title="Edit section&#039;s source code: Bees as pets"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/enwiki/wiki/File:Manda%C3%A7aias.jpg" class="mw-file-description"><img src="/upwiki/wikipedia/commons/thumb/f/ff/Manda%C3%A7aias.jpg/220px-Manda%C3%A7aias.jpg" decoding="async" width="220" height="171" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/f/ff/Manda%C3%A7aias.jpg/330px-Manda%C3%A7aias.jpg 1.5x, /upwiki/wikipedia/commons/thumb/f/ff/Manda%C3%A7aias.jpg/440px-Manda%C3%A7aias.jpg 2x" data-file-width="1879" data-file-height="1464" /></a><figcaption>A swarm of mandaçaias bees around an artificial <a href="/enwiki/wiki/Beehive" title="Beehive">beehive</a> installed in a house's backyard in <a href="/enwiki/wiki/Brazil" title="Brazil">Brazil</a></figcaption></figure> <p>Due to the lack of a functional stinger and characteristic nonaggressive behavior of many Brazilian species of stingless bees, they can be reared without problems in densely populated environments (residential buildings, schools, urban parks), provided enough flowers are at their disposal nearby. Some breeders (<i>meliponicultores</i>) can produce honey even in apartments up to the 12th floor.<sup id="cite_ref-128" class="reference"><a href="#cite_note-128"><span class="cite-bracket">&#91;</span>128<span class="cite-bracket">&#93;</span></a></sup> </p><p>The <a href="/enwiki/wiki/Melipona_quadrifasciata" title="Melipona quadrifasciata">mandaçaias</a> (<i>Melipona quadrifasciata</i>) are extremely tame, rarely attacking humans (only when their hives are opened for honey extraction or colony division). They form small, manageable colonies of only 400&#8211;600&#160;individuals. They are fairly large bees, up to 11&#160;mm (7/16") in length, and as a result have better body heat control, allowing them to live in regions where temperatures can drop a little lower than 0&#160;°C (32&#160;°F). However, they are somewhat selective about which flowers they will visit, preferring the flora that occurs in their natural environment. They are thus difficult to keep outside their region of origin (the eastern coast of Brazil). Once very common, the mandaçaia is now rather rare in nature, mainly due to the destruction of their native forests in the of Brazil.<sup id="cite_ref-129" class="reference"><a href="#cite_note-129"><span class="cite-bracket">&#91;</span>129<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:33_122-1" class="reference"><a href="#cite_note-:33-122"><span class="cite-bracket">&#91;</span>122<span class="cite-bracket">&#93;</span></a></sup> </p><p>Other groups of Brazilian stingless bees, genera <i><a href="/enwiki/wiki/Plebeia" title="Plebeia">Plebeia</a></i> and <i><a href="/enwiki/wiki/Leurotrigona" title="Leurotrigona">Leurotrigona</a></i>, are also very tame and much smaller, with one of them (<i><a href="/enwiki/w/index.php?title=Plebeia_minima&amp;action=edit&amp;redlink=1" class="new" title="Plebeia minima (page does not exist)">Plebeia minima</a></i>) reaching no more than 2.5&#160;mm (3/32") in length, and the <i><a href="/enwiki/wiki/Lambe-olhos" class="mw-redirect" title="Lambe-olhos">lambe-olhos</a></i> ("lick-eyes" bee, <i><a href="/enwiki/wiki/Leurotrigona_muelleri" title="Leurotrigona muelleri">Leurotrigona muelleri</a></i>) being even smaller, at no more than 1.5&#160;mm (3/32"). Many of these species are known as mirim (meaning 'small' in the <a href="/enwiki/wiki/Tupi-Guarani_languages" class="mw-redirect" title="Tupi-Guarani languages">Tupi-Guarani languages</a>). As a result, they can be kept in very small artificial hives, thus being of interest for keepers who want them as pollinators in small glasshouses or just for the pleasure of having a 'toy' bee colony at home.<sup id="cite_ref-:33_122-2" class="reference"><a href="#cite_note-:33-122"><span class="cite-bracket">&#91;</span>122<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-130" class="reference"><a href="#cite_note-130"><span class="cite-bracket">&#91;</span>130<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-131" class="reference"><a href="#cite_note-131"><span class="cite-bracket">&#91;</span>131<span class="cite-bracket">&#93;</span></a></sup> Being so tiny, these species produce only a very small amount of honey, typically less than 500&#160;ml (1/2 US pint) a year, so are not interesting for commercial honey production. </p> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/enwiki/wiki/File:Abelha-jatai.jpg" class="mw-file-description"><img src="/upwiki/wikipedia/commons/thumb/0/0e/Abelha-jatai.jpg/220px-Abelha-jatai.jpg" decoding="async" width="220" height="147" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/0/0e/Abelha-jatai.jpg/330px-Abelha-jatai.jpg 1.5x, /upwiki/wikipedia/commons/thumb/0/0e/Abelha-jatai.jpg/440px-Abelha-jatai.jpg 2x" data-file-width="957" data-file-height="638" /></a><figcaption>Entrance pipe of <i>jataís´s</i> colony: The <i>jataí</i> bees build a wax pipe in the entrance of their nest.</figcaption></figure> <p>Belonging to the same group, the jataí (<i><a href="/enwiki/wiki/Tetragonisca_angustula" title="Tetragonisca angustula">Tetragonisca angustula</a></i>), the marmelada (<i><a href="/enwiki/wiki/Frieseomelitta_varia" title="Frieseomelitta varia">Frieseomelitta varia</a></i>), and the moça-branca (<i><a href="/enwiki/w/index.php?title=Frieseomelitta_doederleini&amp;action=edit&amp;redlink=1" class="new" title="Frieseomelitta doederleini (page does not exist)">Frieseomelitta doederleini</a></i>) are intermediate in size between those very small species and the European bee. They are very adaptable species; the jataí, and can be reared in many different regions and environments, being quite common in most Brazilian cities. The jataí can bite when disturbed, but its jaws are weak, and in practice they are harmless, while the marmelada and moça-branca usually deposit propolis on their aggressors. Jataí is one of the first species to be kept by home beekeepers. Their nests can be easily identified in trees or wall cavities by the yellow wax pipe they build at the entrance, usually guarded by some soldier bees, which are stronger than regular worker bees. The marmelada and moça-branca make a little less honey, but it is denser and sweeter than most from other stingless bees and is considered very tasty.<sup id="cite_ref-:33_122-3" class="reference"><a href="#cite_note-:33-122"><span class="cite-bracket">&#91;</span>122<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-132" class="reference"><a href="#cite_note-132"><span class="cite-bracket">&#91;</span>132<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Central_America">Central America</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=37" title="Edit section&#039;s source code: Central America"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/enwiki/wiki/File:Colmena_melipona.jpg" class="mw-file-description"><img src="/upwiki/wikipedia/commons/thumb/5/5a/Colmena_melipona.jpg/220px-Colmena_melipona.jpg" decoding="async" width="220" height="88" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/5/5a/Colmena_melipona.jpg/330px-Colmena_melipona.jpg 1.5x, /upwiki/wikipedia/commons/thumb/5/5a/Colmena_melipona.jpg/440px-Colmena_melipona.jpg 2x" data-file-width="640" data-file-height="256" /></a><figcaption>A Maya stingless bee hive: A piece of hollow log provides a home for meliponine bees in <a href="/enwiki/wiki/Belize" title="Belize">Belize</a>.</figcaption></figure> <p>The stingless bees <i><a href="/enwiki/wiki/Melipona_beecheii" title="Melipona beecheii">Melipona beecheii</a></i> and <i><a href="/enwiki/w/index.php?title=Melipona_yucatanica&amp;action=edit&amp;redlink=1" class="new" title="Melipona yucatanica (page does not exist)">M. yucatanica</a></i> are the primary native bees cultured in Central America, though a few other species are reported as being occasionally managed (e.g., <i><a href="/enwiki/wiki/Trigona_fulviventris" title="Trigona fulviventris">Trigona fulviventris</a></i> and <i><a href="/enwiki/wiki/Scaptotrigona_mexicana" title="Scaptotrigona mexicana">Scaptotrigona mexicana</a></i>).<sup id="cite_ref-133" class="reference"><a href="#cite_note-133"><span class="cite-bracket">&#91;</span>133<span class="cite-bracket">&#93;</span></a></sup> They were extensively cultured by the <a href="/enwiki/wiki/Maya_civilization" title="Maya civilization">Maya civilization</a> for honey, and regarded as sacred. They continue to be cultivated by the modern <a href="/enwiki/wiki/Maya_peoples" title="Maya peoples">Maya peoples</a>, although these bees are endangered due to massive deforestation, altered <a href="/enwiki/wiki/Agriculture" title="Agriculture">agricultural</a> practices (especially overuse of <a href="/enwiki/wiki/Insecticide" title="Insecticide">insecticides</a>), and changing beekeeping practices with the arrival of the <a href="/enwiki/wiki/Africanized_honey_bee" class="mw-redirect" title="Africanized honey bee">Africanized honey bee</a>, which produces much greater honey crops.<sup id="cite_ref-:26_72-1" class="reference"><a href="#cite_note-:26-72"><span class="cite-bracket">&#91;</span>72<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="History">History</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=38" title="Edit section&#039;s source code: History"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Native meliponines (<i>M. beecheii</i> being the most common) have been kept by the lowland Maya for thousands of years. The <a href="/enwiki/wiki/Yucatec_Maya_language" title="Yucatec Maya language">Yucatec Maya language</a> name for this bee is <i>xunan kab</i>, meaning "(royal, noble) lady bee".<sup id="cite_ref-134" class="reference"><a href="#cite_note-134"><span class="cite-bracket">&#91;</span>134<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-135" class="reference"><a href="#cite_note-135"><span class="cite-bracket">&#91;</span>135<span class="cite-bracket">&#93;</span></a></sup> The bees were once the subject of religious ceremonies and were a symbol of the bee-god <a href="/enwiki/wiki/Ah-Muzen-Cab" title="Ah-Muzen-Cab">Ah-Muzen-Cab</a>, known from the <i><a href="/enwiki/wiki/Madrid_Codex_(Maya)" title="Madrid Codex (Maya)">Madrid Codex</a></i>.<sup id="cite_ref-:34_136-0" class="reference"><a href="#cite_note-:34-136"><span class="cite-bracket">&#91;</span>136<span class="cite-bracket">&#93;</span></a></sup> </p><p>The bees were, and still are, treated as pets. Families would have one or many log-hives hanging in and around their houses. Although they are stingless, the bees do bite and can leave welts similar to a mosquito bite. The traditional way to gather bees, still favored among the locals, is find a wild hive, then the branch is cut around the hive to create a portable log, enclosing the colony. With proper maintenance, hives have been recorded as lasting over 80 years, being passed down through generations. In the archaeological record of Mesoamerica, stone discs have been found that are generally considered to be the caps of long-disintegrated logs that once housed the beehives.<sup id="cite_ref-:34_136-1" class="reference"><a href="#cite_note-:34-136"><span class="cite-bracket">&#91;</span>136<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-:35_137-0" class="reference"><a href="#cite_note-:35-137"><span class="cite-bracket">&#91;</span>137<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Tulum">Tulum</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=39" title="Edit section&#039;s source code: Tulum"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/enwiki/wiki/Tulum" title="Tulum">Tulum</a>, the site of a <a href="/enwiki/wiki/Pre-Columbian" class="mw-redirect" title="Pre-Columbian">pre-Columbian</a> Maya city on the Caribbean coast 130&#160;km (81&#160;mi) south of Cancun, has a god depicted repeatedly all over the site. Upside down, he appears as a small figure over many doorways and entrances. One of the temples, the Temple of the Descending God (<i>Templo del Dios Descendente</i>), stands just left of the central plaza. Speculation is that he may be the "Bee God", Ah Muzen Cab, as seen in the <i>Madrid Codex</i>. It is possible that this was a religious/trade center with emphasis on <i>xunan kab</i>, the "royal lady".<sup id="cite_ref-:35_137-1" class="reference"><a href="#cite_note-:35-137"><span class="cite-bracket">&#91;</span>137<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Economic_uses">Economic uses</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=40" title="Edit section&#039;s source code: Economic uses"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/enwiki/wiki/Balch%C3%A9" title="Balché">Balché</a>, a traditional <a href="/enwiki/wiki/Mesoamerica" title="Mesoamerica">Mesoamerican</a> alcoholic beverage similar to <a href="/enwiki/wiki/Mead" title="Mead">mead</a>, was made from fermented honey and the bark of the leguminous<i> balché</i> tree (<i><a href="/enwiki/wiki/Lonchocarpus_violaceus" title="Lonchocarpus violaceus">Lonchocarpus violaceus</a></i>), hence its name. It was traditionally brewed in a canoe. The drink was known to have <a href="/enwiki/wiki/Entheogen" title="Entheogen">entheogenic</a> properties, that is, to produce mystical experiences, and was consumed in medicinal and ritual practices. Beekeepers would place the nests near the psychoactive plant <i><a href="/enwiki/wiki/Turbina_corymbosa" class="mw-redirect" title="Turbina corymbosa">Turbina corymbosa</a></i> and possibly near <i>balché</i> trees, forcing the bees to use nectar from these plants to make their honey. Additionally, brewers would add extracts of the bark of the <i>balché</i> tree to the honey mixture before fermentation. The resulting beverage is responsible for psychotropic effects when consumed, due to the ergoline compounds in the pollen of the <i>T. corymbosa</i>, the <i>Melipona</i> nectar gathered from the <i>balché</i> flowers, or the hallucinogenic compounds of the <i>balché</i> tree bark.<sup id="cite_ref-138" class="reference"><a href="#cite_note-138"><span class="cite-bracket">&#91;</span>138<span class="cite-bracket">&#93;</span></a></sup> </p><p><a href="/enwiki/wiki/Lost-wax_casting" title="Lost-wax casting">Lost-wax casting</a>, a common metalworking method typically found where the inhabitants keep bees, was also used by the Maya. The wax from <i>Melipona</i> is soft and easy to work, especially in the humid Maya lowland. This allowed the Maya to create smaller works of art, jewelry, and other metalsmithing that would be difficult to forge. It also makes use of the leftovers from honey extraction. If the hive was damaged beyond repair, the whole of the comb could be used, thus using all of the hive. With experienced keepers, though, only the honey pot could be removed, the honey extracted, and the wax used for casting or other purposes.<sup id="cite_ref-139" class="reference"><a href="#cite_note-139"><span class="cite-bracket">&#91;</span>139<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-140" class="reference"><a href="#cite_note-140"><span class="cite-bracket">&#91;</span>140<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Future">Future</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=41" title="Edit section&#039;s source code: Future"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The outlook for meliponines in Mesoamerica is uncertain. The number of active Meliponini beekeepers is shy in comparison with the Africanized <i><a href="/enwiki/wiki/Apis_mellifera" class="mw-redirect" title="Apis mellifera">Apis mellifera</a></i> breeders. The high honey yield, 100&#160;kg (220&#160;lbs) or more annually, along with the ease of hive care and ability to create new hives from existing stock, commonly outweighs the negative consequences of "killer bee" hive maintenance.<sup id="cite_ref-:36_141-0" class="reference"><a href="#cite_note-:36-141"><span class="cite-bracket">&#91;</span>141<span class="cite-bracket">&#93;</span></a></sup> </p><p>An additional blow to the art of meliponine beekeeping is that many of the <i>meliponicultores</i> are now elderly, and their hives may not be cared for once they die. The hives are considered similar to an old family collection, to be parted out once the collector dies or to be buried in whole or part along with the beekeeper upon death. In fact, a survey of a once-popular area of the Maya lowlands shows the rapid decline of beekeepers, down to around 70 in 2004 from thousands in the late 1980s. Conservation efforts are underway in several parts of Mesoamerica.<sup id="cite_ref-:36_141-1" class="reference"><a href="#cite_note-:36-141"><span class="cite-bracket">&#91;</span>141<span class="cite-bracket">&#93;</span></a></sup><sup id="cite_ref-142" class="reference"><a href="#cite_note-142"><span class="cite-bracket">&#91;</span>142<span class="cite-bracket">&#93;</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=42" title="Edit section&#039;s source code: References"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width reflist-columns-2"> <ol class="references"> <li id="cite_note-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, pp.&#160;46–47)</span> </li> <li id="cite_note-:27-2"><span class="mw-cite-backlink">^ <a href="#cite_ref-:27_2-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:27_2-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-:27_2-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text"><a href="#CITEREFNogueira2023">Nogueira (2023)</a></span> </li> <li id="cite_note-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-3">^</a></b></span> <span class="reference-text"><a href="#CITEREFMichener2000">Michener (2000</a>, p.&#160;803)</span> </li> <li id="cite_note-4"><span class="mw-cite-backlink"><b><a href="#cite_ref-4">^</a></b></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, p.&#160;1)</span> </li> <li id="cite_note-5"><span class="mw-cite-backlink"><b><a href="#cite_ref-5">^</a></b></span> <span class="reference-text"><a href="#CITEREFSilveira2002">Silveira (2002</a>, p.&#160;253)<span class="error harv-error" style="display: none; font-size:100%"> harvtxt error: no target: CITEREFSilveira2002 (<a href="/enwiki/wiki/Category:Harv_and_Sfn_template_errors" title="Category:Harv and Sfn template errors">help</a>)</span></span> </li> <li id="cite_note-6"><span class="mw-cite-backlink"><b><a href="#cite_ref-6">^</a></b></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, p.&#160;43)</span> </li> <li id="cite_note-7"><span class="mw-cite-backlink"><b><a href="#cite_ref-7">^</a></b></span> <span class="reference-text"><a href="#CITEREFRoubik1989">Roubik (1989</a>, p.&#160;8)</span> </li> <li id="cite_note-8"><span class="mw-cite-backlink"><b><a href="#cite_ref-8">^</a></b></span> <span class="reference-text"><a href="#CITEREFKajobe2006">Kajobe (2006)</a></span> </li> <li id="cite_note-:1-9"><span class="mw-cite-backlink">^ <a href="#cite_ref-:1_9-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:1_9-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><a href="#CITEREFChakuyaGandiwaMubokoMuposhi2022">Chakuya et al. (2022)</a></span> </li> <li id="cite_note-10"><span class="mw-cite-backlink"><b><a href="#cite_ref-10">^</a></b></span> <span class="reference-text"><a href="#CITEREFMichener2000">Michener (2000</a>, p.&#160;111)</span> </li> <li id="cite_note-11"><span class="mw-cite-backlink"><b><a href="#cite_ref-11">^</a></b></span> <span class="reference-text"><a href="#CITEREFSarchet2014">Sarchet (2014)</a></span> </li> <li id="cite_note-12"><span class="mw-cite-backlink"><b><a href="#cite_ref-12">^</a></b></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, pp.&#160;7 &amp; 16)</span> </li> <li id="cite_note-:3-13"><span class="mw-cite-backlink">^ <a href="#cite_ref-:3_13-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:3_13-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, p.&#160;65)</span> </li> <li id="cite_note-14"><span class="mw-cite-backlink"><b><a href="#cite_ref-14">^</a></b></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, p.&#160;4)</span> </li> <li id="cite_note-15"><span class="mw-cite-backlink"><b><a href="#cite_ref-15">^</a></b></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, pp.&#160;25–27)</span> </li> <li id="cite_note-16"><span class="mw-cite-backlink"><b><a href="#cite_ref-16">^</a></b></span> <span class="reference-text"><a href="#CITEREFVillas-Bôas2018">Villas-Bôas (2018</a>, pp.&#160;14–15)<span class="error harv-error" style="display: none; font-size:100%"> harvtxt error: no target: CITEREFVillas-Bôas2018 (<a href="/enwiki/wiki/Category:Harv_and_Sfn_template_errors" title="Category:Harv and Sfn template errors">help</a>)</span></span> </li> <li id="cite_note-17"><span class="mw-cite-backlink"><b><a href="#cite_ref-17">^</a></b></span> <span class="reference-text"><a href="#CITEREFMichener2000">Michener (2000</a>, p.&#160;803)</span> </li> <li id="cite_note-:0-18"><span class="mw-cite-backlink">^ <a href="#cite_ref-:0_18-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:0_18-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-:0_18-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, pp.&#160;47–49)</span> </li> <li id="cite_note-19"><span class="mw-cite-backlink"><b><a href="#cite_ref-19">^</a></b></span> <span class="reference-text"><a href="#CITEREFSilveira2002">Silveira (2002</a>, p.&#160;31)<span class="error harv-error" style="display: none; font-size:100%"> harvtxt error: no target: CITEREFSilveira2002 (<a href="/enwiki/wiki/Category:Harv_and_Sfn_template_errors" title="Category:Harv and Sfn template errors">help</a>)</span></span> </li> <li id="cite_note-20"><span class="mw-cite-backlink"><b><a href="#cite_ref-20">^</a></b></span> <span class="reference-text"><a href="#CITEREFRasmussenThomasEngel2017">Rasmussen, Thomas &amp; Engel (2017)</a></span> </li> <li id="cite_note-21"><span class="mw-cite-backlink"><b><a href="#cite_ref-21">^</a></b></span> <span class="reference-text"><a href="#CITEREFCortopassi-LaurinoImperatriz-FonsecaRoubik2006">Cortopassi-Laurino, Imperatriz-Fonseca &amp; Roubik (2006)</a><span class="error harv-error" style="display: none; font-size:100%"> harvtxt error: no target: CITEREFCortopassi-LaurinoImperatriz-FonsecaRoubik2006 (<a href="/enwiki/wiki/Category:Harv_and_Sfn_template_errors" title="Category:Harv and Sfn template errors">help</a>)</span></span> </li> <li id="cite_note-22"><span class="mw-cite-backlink"><b><a href="#cite_ref-22">^</a></b></span> <span class="reference-text"><a href="#CITEREFVenturieriRaiolPareira2003">Venturieri, Raiol &amp; Pareira (2003)</a></span> </li> <li id="cite_note-23"><span class="mw-cite-backlink"><b><a href="#cite_ref-23">^</a></b></span> <span class="reference-text"><a href="#CITEREFKoch2010">Koch (2010)</a></span> </li> <li id="cite_note-24"><span class="mw-cite-backlink"><b><a href="#cite_ref-24">^</a></b></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, pp.&#160;28–29)</span> </li> <li id="cite_note-25"><span class="mw-cite-backlink"><b><a href="#cite_ref-25">^</a></b></span> <span class="reference-text"><a href="#CITEREFSouzaYuyamaAguiarOliveira2004">Souza et al. (2004)</a></span> </li> <li id="cite_note-:2-26"><span class="mw-cite-backlink">^ <a href="#cite_ref-:2_26-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:2_26-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><a href="#CITEREFRoubik2023a">Roubik (2023a)</a><span class="error harv-error" style="display: none; font-size:100%"> harvtxt error: no target: CITEREFRoubik2023a (<a href="/enwiki/wiki/Category:Harv_and_Sfn_template_errors" title="Category:Harv and Sfn template errors">help</a>)</span></span> </li> <li id="cite_note-27"><span class="mw-cite-backlink"><b><a href="#cite_ref-27">^</a></b></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, pp.&#160;54–56)</span> </li> <li id="cite_note-28"><span class="mw-cite-backlink"><b><a href="#cite_ref-28">^</a></b></span> <span class="reference-text"><a href="#CITEREFRasmussenCameron2009">Rasmussen &amp; Cameron (2009)</a></span> </li> <li id="cite_note-29"><span class="mw-cite-backlink"><b><a href="#cite_ref-29">^</a></b></span> <span class="reference-text"><a href="#CITEREFJalil2014">Jalil (2014</a>, p.&#160;126)</span> </li> <li id="cite_note-:29-30"><span class="mw-cite-backlink">^ <a href="#cite_ref-:29_30-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:29_30-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-:29_30-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, pp.&#160;43–44)</span> </li> <li id="cite_note-31"><span class="mw-cite-backlink"><b><a href="#cite_ref-31">^</a></b></span> <span class="reference-text"><a href="#CITEREFPayne2014">Payne (2014)</a></span> </li> <li id="cite_note-:24-32"><span class="mw-cite-backlink">^ <a href="#cite_ref-:24_32-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:24_32-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-:24_32-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-:24_32-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text"><a href="#CITEREFGrüter2010">Grüter (2010</a>, pp.&#160;49–51)<span class="error harv-error" style="display: none; font-size:100%"> harvtxt error: no target: CITEREFGrüter2010 (<a href="/enwiki/wiki/Category:Harv_and_Sfn_template_errors" title="Category:Harv and Sfn template errors">help</a>)</span></span> </li> <li id="cite_note-33"><span class="mw-cite-backlink"><b><a href="#cite_ref-33">^</a></b></span> <span class="reference-text"><a href="#CITEREFRasmussenCameron2009">Rasmussen &amp; Cameron (2009)</a></span> </li> <li id="cite_note-34"><span class="mw-cite-backlink"><b><a href="#cite_ref-34">^</a></b></span> <span class="reference-text"><a href="#CITEREFCardinalDanforth2011">Cardinal &amp; Danforth (2011)</a></span> </li> <li id="cite_note-35"><span class="mw-cite-backlink"><b><a href="#cite_ref-35">^</a></b></span> <span class="reference-text"><a href="#CITEREFCardinalDanforth2013">Cardinal &amp; Danforth (2013)</a></span> </li> <li id="cite_note-36"><span class="mw-cite-backlink"><b><a href="#cite_ref-36">^</a></b></span> <span class="reference-text"><a href="#CITEREFMartinsMeloRenner2014">Martins, Melo &amp; Renner (2014)</a></span> </li> <li id="cite_note-:4-37"><span class="mw-cite-backlink">^ <a href="#cite_ref-:4_37-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:4_37-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><a href="#CITEREFNogueira-Neto1997">Nogueira-Neto (1997</a>, p.&#160;78)<span class="error harv-error" style="display: none; font-size:100%"> harvtxt error: no target: CITEREFNogueira-Neto1997 (<a href="/enwiki/wiki/Category:Harv_and_Sfn_template_errors" title="Category:Harv and Sfn template errors">help</a>)</span></span> </li> <li id="cite_note-:5-38"><span class="mw-cite-backlink">^ <a href="#cite_ref-:5_38-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:5_38-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><a href="#CITEREFNogueira-Neto1997">Nogueira-Neto (1997</a>, p.&#160;82)<span class="error harv-error" style="display: none; font-size:100%"> harvtxt error: no target: CITEREFNogueira-Neto1997 (<a href="/enwiki/wiki/Category:Harv_and_Sfn_template_errors" title="Category:Harv and Sfn template errors">help</a>)</span></span> </li> <li id="cite_note-39"><span class="mw-cite-backlink"><b><a href="#cite_ref-39">^</a></b></span> <span class="reference-text"><a href="#CITEREFNogueira-Neto1997">Nogueira-Neto (1997</a>, pp.&#160;40–46)<span class="error harv-error" style="display: none; font-size:100%"> harvtxt error: no target: CITEREFNogueira-Neto1997 (<a href="/enwiki/wiki/Category:Harv_and_Sfn_template_errors" title="Category:Harv and Sfn template errors">help</a>)</span></span> </li> <li id="cite_note-40"><span class="mw-cite-backlink"><b><a href="#cite_ref-40">^</a></b></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, pp.&#160;12–15)</span> </li> <li id="cite_note-41"><span class="mw-cite-backlink"><b><a href="#cite_ref-41">^</a></b></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, pp.&#160;87–121)</span> </li> <li id="cite_note-42"><span class="mw-cite-backlink"><b><a href="#cite_ref-42">^</a></b></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, pp.&#160;7–11)</span> </li> <li id="cite_note-:9-43"><span class="mw-cite-backlink">^ <a href="#cite_ref-:9_43-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:9_43-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><a href="#CITEREFVillas-Bôas2018">Villas-Bôas (2018</a>, p.&#160;23)<span class="error harv-error" style="display: none; font-size:100%"> harvtxt error: no target: CITEREFVillas-Bôas2018 (<a href="/enwiki/wiki/Category:Harv_and_Sfn_template_errors" title="Category:Harv and Sfn template errors">help</a>)</span></span> </li> <li id="cite_note-44"><span class="mw-cite-backlink"><b><a href="#cite_ref-44">^</a></b></span> <span class="reference-text"><a href="#CITEREFNogueira-Neto1997">Nogueira-Neto (1997</a>, pp.&#160;82–83)<span class="error harv-error" style="display: none; font-size:100%"> harvtxt error: no target: CITEREFNogueira-Neto1997 (<a href="/enwiki/wiki/Category:Harv_and_Sfn_template_errors" title="Category:Harv and Sfn template errors">help</a>)</span></span> </li> <li id="cite_note-:6-45"><span class="mw-cite-backlink">^ <a href="#cite_ref-:6_45-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:6_45-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><a href="#CITEREFVillas-Bôas2018">Villas-Bôas (2018</a>, p.&#160;19)<span class="error harv-error" style="display: none; font-size:100%"> harvtxt error: no target: CITEREFVillas-Bôas2018 (<a href="/enwiki/wiki/Category:Harv_and_Sfn_template_errors" title="Category:Harv and Sfn template errors">help</a>)</span></span> </li> <li id="cite_note-:7-46"><span class="mw-cite-backlink">^ <a href="#cite_ref-:7_46-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:7_46-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-:7_46-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, pp.&#160;7 &amp; 5)</span> </li> <li id="cite_note-:8-47"><span class="mw-cite-backlink">^ <a href="#cite_ref-:8_47-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:8_47-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><a href="#CITEREFImperatriz-FonsecaZucchi1995">Imperatriz-Fonseca &amp; Zucchi (1995)</a></span> </li> <li id="cite_note-48"><span class="mw-cite-backlink"><b><a href="#cite_ref-48">^</a></b></span> <span class="reference-text"><a href="#CITEREFNogueira-Neto1997">Nogueira-Neto (1997</a>, pp.&#160;83–85)<span class="error harv-error" style="display: none; font-size:100%"> harvtxt error: no target: CITEREFNogueira-Neto1997 (<a href="/enwiki/wiki/Category:Harv_and_Sfn_template_errors" title="Category:Harv and Sfn template errors">help</a>)</span></span> </li> <li id="cite_note-49"><span class="mw-cite-backlink"><b><a href="#cite_ref-49">^</a></b></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, p.&#160;11)</span> </li> <li id="cite_note-50"><span class="mw-cite-backlink"><b><a href="#cite_ref-50">^</a></b></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, p.&#160;11)</span> </li> <li id="cite_note-51"><span class="mw-cite-backlink"><b><a href="#cite_ref-51">^</a></b></span> <span class="reference-text"><a href="#CITEREFGrüterMenezesImperatriz-FonsecaRatnieks2012">Grüter et al. (2012)</a></span> </li> <li id="cite_note-52"><span class="mw-cite-backlink"><b><a href="#cite_ref-52">^</a></b></span> <span class="reference-text"><a href="#CITEREFGrüterSegersMenezesVollet-Neto2017">Grüter et al. (2017)</a></span> </li> <li id="cite_note-53"><span class="mw-cite-backlink"><b><a href="#cite_ref-53">^</a></b></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, p.&#160;258)</span> </li> <li id="cite_note-54"><span class="mw-cite-backlink"><b><a href="#cite_ref-54">^</a></b></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, pp.&#160;17–24)</span> </li> <li id="cite_note-:10-55"><span class="mw-cite-backlink">^ <a href="#cite_ref-:10_55-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:10_55-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-:10_55-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-:10_55-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text"><a href="#CITEREFNogueira-Neto1997">Nogueira-Neto (1997</a>, pp.&#160;40–46)<span class="error harv-error" style="display: none; font-size:100%"> harvtxt error: no target: CITEREFNogueira-Neto1997 (<a href="/enwiki/wiki/Category:Harv_and_Sfn_template_errors" title="Category:Harv and Sfn template errors">help</a>)</span></span> </li> <li id="cite_note-:11-56"><span class="mw-cite-backlink">^ <a href="#cite_ref-:11_56-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:11_56-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-:11_56-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-:11_56-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, pp.&#160;12–15)</span> </li> <li id="cite_note-:12-57"><span class="mw-cite-backlink">^ <a href="#cite_ref-:12_57-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:12_57-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-:12_57-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-:12_57-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text"><a href="#CITEREFVillas-Bôas2018">Villas-Bôas (2018</a>, pp.&#160;24–26)<span class="error harv-error" style="display: none; font-size:100%"> harvtxt error: no target: CITEREFVillas-Bôas2018 (<a href="/enwiki/wiki/Category:Harv_and_Sfn_template_errors" title="Category:Harv and Sfn template errors">help</a>)</span></span> </li> <li id="cite_note-:13-58"><span class="mw-cite-backlink">^ <a href="#cite_ref-:13_58-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:13_58-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-:13_58-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-:13_58-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text"><a href="#CITEREFVenturieri2004">Venturieri (2004</a>, pp.&#160;23–30)</span> </li> <li id="cite_note-59"><span class="mw-cite-backlink"><b><a href="#cite_ref-59">^</a></b></span> <span class="reference-text"><a href="#CITEREFMdudaHusseinMuruke2023">Mduda, Hussein &amp; Muruke (2023)</a></span> </li> <li id="cite_note-60"><span class="mw-cite-backlink"><b><a href="#cite_ref-60">^</a></b></span> <span class="reference-text"><a href="#CITEREFFerreiraLencioniBenassiBarth2009">Ferreira et al. 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font-size:100%"> harvtxt error: no target: CITEREFNogueira-Neto1997 (<a href="/enwiki/wiki/Category:Harv_and_Sfn_template_errors" title="Category:Harv and Sfn template errors">help</a>)</span></span> </li> <li id="cite_note-84"><span class="mw-cite-backlink"><b><a href="#cite_ref-84">^</a></b></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, p.&#160;236)</span> </li> <li id="cite_note-:92-85"><span class="mw-cite-backlink"><b><a href="#cite_ref-:92_85-0">^</a></b></span> <span class="reference-text"><a href="#CITEREFVillas-Bôas2018">Villas-Bôas (2018</a>, p.&#160;106)<span class="error harv-error" style="display: none; font-size:100%"> harvtxt error: no target: CITEREFVillas-Bôas2018 (<a href="/enwiki/wiki/Category:Harv_and_Sfn_template_errors" title="Category:Harv and Sfn template errors">help</a>)</span></span> </li> <li id="cite_note-86"><span class="mw-cite-backlink"><b><a href="#cite_ref-86">^</a></b></span> <span class="reference-text"><a href="#CITEREFGrüter2020">Grüter (2020</a>, p.&#160;235)</span> </li> <li id="cite_note-87"><span class="mw-cite-backlink"><b><a href="#cite_ref-87">^</a></b></span> <span class="reference-text"><a href="#CITEREFVillas-Bôas2018">Villas-Bôas (2018</a>, p.&#160;105)<span class="error harv-error" style="display: none; 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Retrieved <span class="nowrap">12 March</span> 2016</span>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=unknown&amp;rft.btitle=Meliponario+Capixaba%3A+%C3%89+POSS%C3%8DVEL+CRIAR+ABELHAS+EM+APARTAMENTOS%3F&amp;rft.au=Jo%C3%A3o+Luiz&amp;rft_id=http%3A%2F%2Fmeliponariocapixaba.blogspot.com.br%2F2009%2F11%2Fe-possivel-criar-abelhas-em.html&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AStingless+bee" class="Z3988"></span></span> </li> <li id="cite_note-129"><span class="mw-cite-backlink"><b><a href="#cite_ref-129">^</a></b></span> <span class="reference-text"><a href="#CITEREFBritoFaquinelloPaula-LeiteCarvalho1997">Brito et al. (1997)</a><span class="error harv-error" style="display: none; font-size:100%"> harvtxt error: no target: CITEREFBritoFaquinelloPaula-LeiteCarvalho1997 (<a href="/enwiki/wiki/Category:Harv_and_Sfn_template_errors" title="Category:Harv and Sfn template errors">help</a>)</span></span> </li> <li id="cite_note-130"><span class="mw-cite-backlink"><b><a href="#cite_ref-130">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://sites.usp.br/jardimdabotanicausprp/lambe-olhos-leurotrigona-muelleri/">"Lambe-olhos – Leurotrigona muelleri – Laboratório de Sistemática de Plantas"</a>. <i>sites.usp.br</i><span class="reference-accessdate">. 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Retrieved <span class="nowrap">2024-12-31</span></span>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=unknown&amp;rft.jtitle=sites.usp.br&amp;rft.atitle=Mirim+%E2%80%93+Plebeia+droryana+%E2%80%93+Laborat%C3%B3rio+de+Sistem%C3%A1tica+de+Plantas&amp;rft_id=https%3A%2F%2Fsites.usp.br%2Fjardimdabotanicausprp%2Fmirim-plebeia-droryana%2F&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AStingless+bee" class="Z3988"></span></span> </li> <li id="cite_note-132"><span class="mw-cite-backlink"><b><a href="#cite_ref-132">^</a></b></span> <span class="reference-text"><a href="#CITEREFCastanheiraContel2005">Castanheira &amp; Contel (2005)</a></span> </li> <li id="cite_note-133"><span class="mw-cite-backlink"><b><a href="#cite_ref-133">^</a></b></span> <span class="reference-text"><a href="#CITEREFKent1984">Kent (1984)</a></span> </li> <li id="cite_note-134"><span class="mw-cite-backlink"><b><a href="#cite_ref-134">^</a></b></span> <span class="reference-text"><a href="#CITEREFGrüter2010">Grüter (2010</a>, p.&#160;54)<span class="error harv-error" style="display: none; font-size:100%"> harvtxt error: no target: CITEREFGrüter2010 (<a href="/enwiki/wiki/Category:Harv_and_Sfn_template_errors" title="Category:Harv and Sfn template errors">help</a>)</span></span> </li> <li id="cite_note-135"><span class="mw-cite-backlink"><b><a href="#cite_ref-135">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1 cs1-prop-foreign-lang-source"><a rel="nofollow" class="external text" href="http://www.uqroo.mx/libros/maya/diccionario.pdf">"Diccionario Introductorio"</a> <span class="cs1-format">(PDF)</span>. <i>uqroo.mx</i> (in Spanish). Universidad De Quintana Roo<span class="reference-accessdate">. Retrieved <span class="nowrap">27 March</span> 2014</span>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=unknown&amp;rft.jtitle=uqroo.mx&amp;rft.atitle=Diccionario+Introductorio&amp;rft_id=http%3A%2F%2Fwww.uqroo.mx%2Flibros%2Fmaya%2Fdiccionario.pdf&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AStingless+bee" class="Z3988"></span></span> </li> <li id="cite_note-:34-136"><span class="mw-cite-backlink">^ <a href="#cite_ref-:34_136-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:34_136-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://ecomercioagrario.com/en/people-and-bees-mayan-bees-in-the-madrid-codex/">"People and bees. Mayan bees in the Madrid Codex"</a>. <i>eComercio Agrario</i>. 2016-07-17<span class="reference-accessdate">. Retrieved <span class="nowrap">2024-12-31</span></span>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=unknown&amp;rft.jtitle=eComercio+Agrario&amp;rft.atitle=People+and+bees.+Mayan+bees+in+the+Madrid+Codex&amp;rft.date=2016-07-17&amp;rft_id=https%3A%2F%2Fecomercioagrario.com%2Fen%2Fpeople-and-bees-mayan-bees-in-the-madrid-codex%2F&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AStingless+bee" class="Z3988"></span></span> </li> <li id="cite_note-:35-137"><span class="mw-cite-backlink">^ <a href="#cite_ref-:35_137-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:35_137-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKennedyArghiris2019" class="citation web cs1">Kennedy, Jennifer; Arghiris, Richard (2019-01-31). <a rel="nofollow" class="external text" href="https://news.mongabay.com/2019/01/house-of-the-royal-lady-bee-maya-revive-native-bees-and-ancient-beekeeping/">"House of the Royal Lady Bee: Maya revive native bees and ancient beekeeping"</a>. <i>Mongabay Environmental News</i><span class="reference-accessdate">. Retrieved <span class="nowrap">2024-12-31</span></span>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=unknown&amp;rft.jtitle=Mongabay+Environmental+News&amp;rft.atitle=House+of+the+Royal+Lady+Bee%3A+Maya+revive+native+bees+and+ancient+beekeeping&amp;rft.date=2019-01-31&amp;rft.aulast=Kennedy&amp;rft.aufirst=Jennifer&amp;rft.au=Arghiris%2C+Richard&amp;rft_id=https%3A%2F%2Fnews.mongabay.com%2F2019%2F01%2Fhouse-of-the-royal-lady-bee-maya-revive-native-bees-and-ancient-beekeeping%2F&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AStingless+bee" class="Z3988"></span></span> </li> <li id="cite_note-138"><span class="mw-cite-backlink"><b><a href="#cite_ref-138">^</a></b></span> <span class="reference-text"><a href="#CITEREFOtt1998">Ott (1998)</a></span> </li> <li id="cite_note-139"><span class="mw-cite-backlink"><b><a href="#cite_ref-139">^</a></b></span> <span class="reference-text"><a href="#CITEREFPitses2018">Pitses (2018)</a><span class="error harv-error" style="display: none; font-size:100%"> harvtxt error: no target: CITEREFPitses2018 (<a href="/enwiki/wiki/Category:Harv_and_Sfn_template_errors" title="Category:Harv and Sfn template errors">help</a>)</span></span> </li> <li id="cite_note-140"><span class="mw-cite-backlink"><b><a href="#cite_ref-140">^</a></b></span> <span class="reference-text"><a href="#CITEREFSchwarz1945">Schwarz (1945)</a></span> </li> <li id="cite_note-:36-141"><span class="mw-cite-backlink">^ <a href="#cite_ref-:36_141-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:36_141-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><a href="#CITEREFVillanueva-GRoubikColli-Ucán2005">Villanueva-G, Roubik &amp; Colli-Ucán (2005)</a></span> </li> <li id="cite_note-142"><span class="mw-cite-backlink"><b><a href="#cite_ref-142">^</a></b></span> <span class="reference-text">A comprehensive conservation guide can be found in the June 2005 issue of <i>Bee World</i>.</span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="Bibliography">Bibliography</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=43" title="Edit section&#039;s source code: Bibliography"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Articles_and_publications">Articles and publications</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=44" title="Edit section&#039;s source code: Articles and publications"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAlvesImperatriz-FonsecaSantos-Filho2009" class="citation journal cs1">Alves, D A; Imperatriz-Fonseca, V L; Santos-Filho, P S. (2009). <a rel="nofollow" class="external text" href="https://doi.org/10.4238%2Fvol8-2kerr030">"Production of workers, queens and males in <i>Plebeia remota</i> colonies (Hymenoptera, Apidae, Meliponini), a stingless bee with reproductive diapause"</a>. <i>Genetics and Molecular Research</i>. <b>8</b> (2): <span class="nowrap">672–</span>683. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.4238%2Fvol8-2kerr030">10.4238/vol8-2kerr030</a></span>. <a href="/enwiki/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&#160;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/19554766">19554766</a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.jtitle=Genetics+and+Molecular+Research&amp;rft.atitle=Production+of+workers%2C+queens+and+males+in+Plebeia+remota+colonies+%28Hymenoptera%2C+Apidae%2C+Meliponini%29%2C+a+stingless+bee+with+reproductive+diapause&amp;rft.volume=8&amp;rft.issue=2&amp;rft.pages=%3Cspan+class%3D%22nowrap%22%3E672-%3C%2Fspan%3E683&amp;rft.date=2009&amp;rft_id=info%3Adoi%2F10.4238%2Fvol8-2kerr030&amp;rft_id=info%3Apmid%2F19554766&amp;rft.aulast=Alves&amp;rft.aufirst=D+A&amp;rft.au=Imperatriz-Fonseca%2C+V+L&amp;rft.au=Santos-Filho%2C+P+S.&amp;rft_id=https%3A%2F%2Fdoi.org%2F10.4238%252Fvol8-2kerr030&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AStingless+bee" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAntoniniMartins2003" class="citation journal cs1">Antonini, Yasmine; Martins, Rogério P. 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(June 2005). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1626201">"Working-class royalty: Bees beat the caste system"</a>. <i><a href="/enwiki/wiki/Biology_Letters" title="Biology Letters">Biology Letters</a></i>. <b>1</b> (2): <span class="nowrap">125–</span>128. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1098%2Frsbl.2004.0281">10.1098/rsbl.2004.0281</a>. <a href="/enwiki/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a>&#160;<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1626201">1626201</a></span>. <a href="/enwiki/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&#160;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/17148145">17148145</a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.jtitle=Biology+Letters&amp;rft.atitle=Working-class+royalty%3A+Bees+beat+the+caste+system&amp;rft.volume=1&amp;rft.issue=2&amp;rft.pages=%3Cspan+class%3D%22nowrap%22%3E125-%3C%2Fspan%3E128&amp;rft.date=2005-06&amp;rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1626201%23id-name%3DPMC&amp;rft_id=info%3Apmid%2F17148145&amp;rft_id=info%3Adoi%2F10.1098%2Frsbl.2004.0281&amp;rft.aulast=Wenseleers&amp;rft.aufirst=T.&amp;rft.au=Ratnieks%2C+F.+L.+W.&amp;rft.au=Ribeiro%2C+M.+D.+F.&amp;rft.au=Alves%2C+D.+D.+A.&amp;rft.au=Imperatriz-Fonseca%2C+V.+L.&amp;rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1626201&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AStingless+bee" class="Z3988"></span><span style="position:relative; top: -2px;"><span typeof="mw:File"><a href="/enwiki/wiki/Open_access" title="open access publication – free to read"><img alt="Open access icon" src="/upwiki/wikipedia/commons/thumb/7/77/Open_Access_logo_PLoS_transparent.svg/9px-Open_Access_logo_PLoS_transparent.svg.png" decoding="async" width="9" height="14" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/7/77/Open_Access_logo_PLoS_transparent.svg/14px-Open_Access_logo_PLoS_transparent.svg.png 1.5x, /upwiki/wikipedia/commons/thumb/7/77/Open_Access_logo_PLoS_transparent.svg/18px-Open_Access_logo_PLoS_transparent.svg.png 2x" data-file-width="640" data-file-height="1000" /></a></span></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWitterBlochteinSantos2007" class="citation journal cs1">Witter, Sidia; Blochtein, Betina; Santos, Camila dos (August 2007). <a rel="nofollow" class="external text" href="https://www.agricultura.rs.gov.br/upload/arquivos/202105/11142006-boletim-15.pdf">"Abelhas sem ferrão do Rio Grande do Sul: manejo e conservação"</a> <span class="cs1-format">(PDF)</span>. <i>Boletim FEPAGRO</i> (15).</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.jtitle=Boletim+FEPAGRO&amp;rft.atitle=Abelhas+sem+ferr%C3%A3o+do+Rio+Grande+do+Sul%3A+manejo+e+conserva%C3%A7%C3%A3o&amp;rft.issue=15&amp;rft.date=2007-08&amp;rft.aulast=Witter&amp;rft.aufirst=Sidia&amp;rft.au=Blochtein%2C+Betina&amp;rft.au=Santos%2C+Camila+dos&amp;rft_id=https%3A%2F%2Fwww.agricultura.rs.gov.br%2Fupload%2Farquivos%2F202105%2F11142006-boletim-15.pdf&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AStingless+bee" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFZulkhairi_AminSabriMohammadIsmail2018" class="citation journal cs1">Zulkhairi Amin, Fatin Aina; Sabri, Suriana; Mohammad, Salma Malihah; Ismail, Maznah; Chan, Kim Wei; Ismail, Norsharina; Norhaizan, Mohd Esa; Zawawi, Norhasnida (2018-12-26). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6327266">"Therapeutic Properties of Stingless Bee Honey in Comparison with European Bee Honey"</a>. <i>Advances in Pharmacological and Pharmaceutical Sciences</i>. <b>2018</b>: e6179596. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1155%2F2018%2F6179596">10.1155/2018/6179596</a></span>. <a href="/enwiki/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a>&#160;<a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/2633-4682">2633-4682</a>. <a href="/enwiki/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a>&#160;<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6327266">6327266</a></span>. <a href="/enwiki/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&#160;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/30687402">30687402</a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.jtitle=Advances+in+Pharmacological+and+Pharmaceutical+Sciences&amp;rft.atitle=Therapeutic+Properties+of+Stingless+Bee+Honey+in+Comparison+with+European+Bee+Honey&amp;rft.volume=2018&amp;rft.pages=e6179596&amp;rft.date=2018-12-26&amp;rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC6327266%23id-name%3DPMC&amp;rft.issn=2633-4682&amp;rft_id=info%3Apmid%2F30687402&amp;rft_id=info%3Adoi%2F10.1155%2F2018%2F6179596&amp;rft.aulast=Zulkhairi+Amin&amp;rft.aufirst=Fatin+Aina&amp;rft.au=Sabri%2C+Suriana&amp;rft.au=Mohammad%2C+Salma+Malihah&amp;rft.au=Ismail%2C+Maznah&amp;rft.au=Chan%2C+Kim+Wei&amp;rft.au=Ismail%2C+Norsharina&amp;rft.au=Norhaizan%2C+Mohd+Esa&amp;rft.au=Zawawi%2C+Norhasnida&amp;rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC6327266&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AStingless+bee" class="Z3988"></span></li></ul> <div class="mw-heading mw-heading3"><h3 id="Books">Books</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=45" title="Edit section&#039;s source code: Books"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGrüter2020" class="citation book cs1">Grüter, Christoph (2020). <a rel="nofollow" class="external text" href="http://link.springer.com/10.1007/978-3-030-60090-7"><i>Stingless Bees: Their Behaviour, Ecology and Evolution</i></a>. Fascinating Life Sciences. Cham: Springer International Publishing. <a href="/enwiki/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2F978-3-030-60090-7">10.1007/978-3-030-60090-7</a>. <a href="/enwiki/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a>&#160;<a href="/enwiki/wiki/Special:BookSources/978-3-030-60089-1" title="Special:BookSources/978-3-030-60089-1"><bdi>978-3-030-60089-1</bdi></a>. <a href="/enwiki/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a>&#160;<a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:227250633">227250633</a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=book&amp;rft.btitle=Stingless+Bees%3A+Their+Behaviour%2C+Ecology+and+Evolution&amp;rft.place=Cham&amp;rft.series=Fascinating+Life+Sciences&amp;rft.pub=Springer+International+Publishing&amp;rft.date=2020&amp;rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A227250633%23id-name%3DS2CID&amp;rft_id=info%3Adoi%2F10.1007%2F978-3-030-60090-7&amp;rft.isbn=978-3-030-60089-1&amp;rft.aulast=Gr%C3%BCter&amp;rft.aufirst=Christoph&amp;rft_id=http%3A%2F%2Flink.springer.com%2F10.1007%2F978-3-030-60090-7&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AStingless+bee" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFJalil2014" class="citation book cs1 cs1-prop-foreign-lang-source">Jalil, Abu Hassan (2014). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=fUh9BAAAQBAJ"><i>Beescape for Meliponines: Conservation of Indo-Malayan Stingless Bees</i></a> (in Arabic). Partridge Publishing Singapore. <a href="/enwiki/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a>&#160;<a href="/enwiki/wiki/Special:BookSources/978-1-4828-2362-2" title="Special:BookSources/978-1-4828-2362-2"><bdi>978-1-4828-2362-2</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=book&amp;rft.btitle=Beescape+for+Meliponines%3A+Conservation+of+Indo-Malayan+Stingless+Bees&amp;rft.pub=Partridge+Publishing+Singapore&amp;rft.date=2014&amp;rft.isbn=978-1-4828-2362-2&amp;rft.aulast=Jalil&amp;rft.aufirst=Abu+Hassan&amp;rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DfUh9BAAAQBAJ&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AStingless+bee" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKajobe2008" class="citation book cs1">Kajobe, R. (2008). <a rel="nofollow" class="external text" href="https://books.google.com.br/books/about/Foraging_Behaviour_of_Equatorial_Afrotro.html?id=bMuezQEACAAJ&amp;redir_esc=y"><i>Foraging Behaviour of Equatorial Afrotropical Stingless Bees: Habitat Selection and Competition for Resources</i></a>. Utrecht University.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=book&amp;rft.btitle=Foraging+Behaviour+of+Equatorial+Afrotropical+Stingless+Bees%3A+Habitat+Selection+and+Competition+for+Resources&amp;rft.pub=Utrecht+University&amp;rft.date=2008&amp;rft.aulast=Kajobe&amp;rft.aufirst=R.&amp;rft_id=https%3A%2F%2Fbooks.google.com.br%2Fbooks%2Fabout%2FForaging_Behaviour_of_Equatorial_Afrotro.html%3Fid%3DbMuezQEACAAJ%26redir_esc%3Dy&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AStingless+bee" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMichener2000" class="citation book cs1">Michener, Charles D. (2000). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=bu_1gmY13FIC"><i>The Bees of the World</i></a>. Baltimore, Md.: Johns Hopkins Univ. Press. <a href="/enwiki/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a>&#160;<a href="/enwiki/wiki/Special:BookSources/978-0-8018-6133-8" title="Special:BookSources/978-0-8018-6133-8"><bdi>978-0-8018-6133-8</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=book&amp;rft.btitle=The+Bees+of+the+World&amp;rft.place=Baltimore%2C+Md.&amp;rft.pub=Johns+Hopkins+Univ.+Press&amp;rft.date=2000&amp;rft.isbn=978-0-8018-6133-8&amp;rft.aulast=Michener&amp;rft.aufirst=Charles+D.&amp;rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3Dbu_1gmY13FIC&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AStingless+bee" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRoubik1989" class="citation book cs1">Roubik, David W. (1989). <a rel="nofollow" class="external text" href="https://www.cambridge.org/core/books/ecology-and-natural-history-of-tropical-bees/01B9D0798E5CBA8A68865ED0CC166A4A"><i>Ecology and natural history of tropical bees</i></a>. Cambridge tropical biology series. Cambridge: Cambridge Univ. Pr. <a href="/enwiki/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a>&#160;<a href="/enwiki/wiki/Special:BookSources/978-0-521-26236-1" title="Special:BookSources/978-0-521-26236-1"><bdi>978-0-521-26236-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=book&amp;rft.btitle=Ecology+and+natural+history+of+tropical+bees&amp;rft.place=Cambridge&amp;rft.series=Cambridge+tropical+biology+series&amp;rft.pub=Cambridge+Univ.+Pr&amp;rft.date=1989&amp;rft.isbn=978-0-521-26236-1&amp;rft.aulast=Roubik&amp;rft.aufirst=David+W.&amp;rft_id=https%3A%2F%2Fwww.cambridge.org%2Fcore%2Fbooks%2Fecology-and-natural-history-of-tropical-bees%2F01B9D0798E5CBA8A68865ED0CC166A4A&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AStingless+bee" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSilveiraMeloAlmeida2002" class="citation book cs1">Silveira, Fernando A.; Melo, Gabriel A. R.; Almeida, Eduardo A. B. (2002). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=2-JMAAAAYAAJ&amp;q=Abelhas+brasileiras:+sistem%C3%A1tica+e+identifica%C3%A7%C3%A3o"><i>Abelhas brasileiras: sistemática e identificação</i></a> (1st&#160;ed.). Belo Horizonte: Silveira. <a href="/enwiki/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a>&#160;<a href="/enwiki/wiki/Special:BookSources/978-85-903034-1-1" title="Special:BookSources/978-85-903034-1-1"><bdi>978-85-903034-1-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=book&amp;rft.btitle=Abelhas+brasileiras%3A+sistem%C3%A1tica+e+identifica%C3%A7%C3%A3o&amp;rft.place=Belo+Horizonte&amp;rft.edition=1st&amp;rft.pub=Silveira&amp;rft.date=2002&amp;rft.isbn=978-85-903034-1-1&amp;rft.aulast=Silveira&amp;rft.aufirst=Fernando+A.&amp;rft.au=Melo%2C+Gabriel+A.+R.&amp;rft.au=Almeida%2C+Eduardo+A.+B.&amp;rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3D2-JMAAAAYAAJ%26q%3DAbelhas%2Bbrasileiras%3A%2Bsistem%25C3%25A1tica%2Be%2Bidentifica%25C3%25A7%25C3%25A3o&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AStingless+bee" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFVenturieri2004" class="citation journal cs1">Venturieri, G C. (2004). <a rel="nofollow" class="external text" href="https://www.infoteca.cnptia.embrapa.br/infoteca/bitstream/doc/410121/1/livroabelha2014.pdf">"Meliponicultura: Criação de Abelhas Indígenas Sem Ferrão"</a> <span class="cs1-format">(PDF)</span>. <i>Comunicado Técnico Embrapa Amazônia Oriental</i>. <b>118</b>: <span class="nowrap">1–</span>4.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.jtitle=Comunicado+T%C3%A9cnico+Embrapa+Amaz%C3%B4nia+Oriental&amp;rft.atitle=Meliponicultura%3A+Cria%C3%A7%C3%A3o+de+Abelhas+Ind%C3%ADgenas+Sem+Ferr%C3%A3o&amp;rft.volume=118&amp;rft.pages=%3Cspan+class%3D%22nowrap%22%3E1-%3C%2Fspan%3E4&amp;rft.date=2004&amp;rft.aulast=Venturieri&amp;rft.aufirst=G+C.&amp;rft_id=https%3A%2F%2Fwww.infoteca.cnptia.embrapa.br%2Finfoteca%2Fbitstream%2Fdoc%2F410121%2F1%2Flivroabelha2014.pdf&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AStingless+bee" class="Z3988"></span></li> <li>Villas-Bôas, Jerônimo (2018). <i><a rel="nofollow" class="external text" href="https://ispn.org.br/site/wp-content/uploads/2018/10/ManualTecnologicoMel.pdf">Manual Tecnológico de Aproveitamento Integral dos Produtos das Abelhas Nativas sem Ferrão</a></i>. Brasília: Instituto Sociedade, População e Natureza (ISPN), v. 2.</li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFVitPedroRoubik2018" class="citation book cs1">Vit, Patricia; Pedro, Silvia R. M.; Roubik, David W. (2018-03-05). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=NJxPDwAAQBAJ&amp;dq=%22australia%22+%22native%22+%22stingless+bee%22&amp;pg=PA401"><i>Pot-Pollen in Stingless Bee Melittology</i></a>. Springer. <a href="/enwiki/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a>&#160;<a href="/enwiki/wiki/Special:BookSources/978-3-319-61839-5" title="Special:BookSources/978-3-319-61839-5"><bdi>978-3-319-61839-5</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=book&amp;rft.btitle=Pot-Pollen+in+Stingless+Bee+Melittology&amp;rft.pub=Springer&amp;rft.date=2018-03-05&amp;rft.isbn=978-3-319-61839-5&amp;rft.aulast=Vit&amp;rft.aufirst=Patricia&amp;rft.au=Pedro%2C+Silvia+R.+M.&amp;rft.au=Roubik%2C+David+W.&amp;rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DNJxPDwAAQBAJ%26dq%3D%2522australia%2522%2B%2522native%2522%2B%2522stingless%2Bbee%2522%26pg%3DPA401&amp;rfr_id=info%3Asid%2Fen.wikipedia.org%3AStingless+bee" class="Z3988"></span>&lt;</li></ul> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/enwiki/w/index.php?title=Stingless_bee&amp;action=edit&amp;section=46" title="Edit section&#039;s source code: External links"><span>edit source</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output .side-box-flex{display:flex;align-items:center}.mw-parser-output .side-box-text{flex:1;min-width:0}}@media(min-width:720px){.mw-parser-output .side-box{width:238px}.mw-parser-output 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width="30" height="40" class="mw-file-element" srcset="/upwiki/wikipedia/en/thumb/4/4a/Commons-logo.svg/45px-Commons-logo.svg.png 1.5x, /upwiki/wikipedia/en/thumb/4/4a/Commons-logo.svg/59px-Commons-logo.svg.png 2x" data-file-width="1024" data-file-height="1376" /></a></span></div> <div class="side-box-text plainlist">Wikimedia Commons has media related to <span style="font-weight: bold; font-style: italic;"><a href="https://commons.wikimedia.org/wiki/Category:Meliponini" class="extiw" title="commons:Category:Meliponini">Meliponini</a></span>.</div></div> </div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1235681985"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1237033735"><div class="side-box side-box-right plainlinks sistersitebox"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1126788409"> <div class="side-box-flex"> <div class="side-box-image"><span class="noviewer" typeof="mw:File"><a href="/enwiki/wiki/File:Wikispecies-logo.svg" class="mw-file-description"><img alt="" src="/upwiki/wikipedia/commons/thumb/d/df/Wikispecies-logo.svg/34px-Wikispecies-logo.svg.png" decoding="async" width="34" height="40" class="mw-file-element" srcset="/upwiki/wikipedia/commons/thumb/d/df/Wikispecies-logo.svg/51px-Wikispecies-logo.svg.png 1.5x, /upwiki/wikipedia/commons/thumb/d/df/Wikispecies-logo.svg/68px-Wikispecies-logo.svg.png 2x" data-file-width="941" data-file-height="1103" /></a></span></div> <div class="side-box-text plainlist"><a href="/enwiki/wiki/Wikispecies" title="Wikispecies">Wikispecies</a> has information related to <i><b><a href="https://species.wikimedia.org/wiki/Meliponini" class="extiw" title="wikispecies:Meliponini"><i>Meliponini</i></a></b></i>.</div></div> </div> <ul><li><a rel="nofollow" class="external text" href="https://www.ibra.org.uk/">International Bee Research Association</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20140215065345/http://www.ufv.br/dbg/bee/Versao2/stinglessbees.htm">Stingless honey bees of Brazil</a></li> <li><a rel="nofollow" class="external text" href="https://www.sciencedaily.com/releases/2005/06/050615062105.htm">Mayan Stingless Bee Keeping: Going, Going, Gone?</a></li> <li><a rel="nofollow" class="external text" href="http://green.blogs.nytimes.com/2012/02/17/a-different-kind-of-beekeeping-takes-flight/?_r=0">A Different Kind of Beekeeping Takes Flight</a></li></ul> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output 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.navbox-image img{max-width:none!important}@media print{body.ns-0 .mw-parser-output .navbox{display:none!important}}</style></div><div role="navigation" class="navbox" aria-labelledby="Taxon_identifiers" style="padding:3px"><table class="nowraplinks hlist navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Taxon_identifiers" style="font-size:114%;margin:0 4em"><a href="/enwiki/wiki/Help:Taxon_identifiers" title="Help:Taxon identifiers">Taxon identifiers</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: left;"><i>Meliponini</i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><span style="white-space:nowrap;"><a href="/enwiki/wiki/Wikidata" title="Wikidata">Wikidata</a>: <span class="uid"><span class="external"><a href="https://www.wikidata.org/wiki/Q127768" class="extiw" title="wikidata:Q127768">Q127768</a></span></span></span></li> <li><span style="white-space:nowrap;"><a href="/enwiki/wiki/Wikispecies" title="Wikispecies">Wikispecies</a>: <span class="uid"><span class="external"><a href="https://species.wikimedia.org/wiki/Meliponini" class="extiw" title="wikispecies:Meliponini">Meliponini</a></span></span></span></li> <li><span style="white-space:nowrap;"><a href="/enwiki/wiki/Australian_Faunal_Directory" title="Australian Faunal Directory">AFD</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://biodiversity.org.au/afd/taxa/Meliponini">Meliponini</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/enwiki/wiki/Barcode_of_Life_Data_System" title="Barcode of Life Data System">BOLD</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://bench.boldsystems.org/index.php/TaxBrowser_TaxonPage?taxid=891930">891930</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/enwiki/wiki/Catalogue_of_Life" title="Catalogue of Life">CoL</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.catalogueoflife.org/data/taxon/KZ6">KZ6</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/enwiki/wiki/INaturalist" title="INaturalist">iNaturalist</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://inaturalist.org/taxa/200106">200106</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/enwiki/wiki/Integrated_Taxonomic_Information_System" title="Integrated Taxonomic Information System">ITIS</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&amp;search_value=633943">633943</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/enwiki/wiki/National_Center_for_Biotechnology_Information" title="National Center for Biotechnology Information">NCBI</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Info&amp;id=83319">83319</a></span></span></li> <li><span style="white-space:nowrap;"><a href="/enwiki/wiki/Paleobiology_Database" title="Paleobiology Database">Paleobiology Database</a>: <span class="uid"><a rel="nofollow" class="external text" href="https://paleobiodb.org/classic/basicTaxonInfo?taxon_no=193747">193747</a></span></span></li></ul> </div></td></tr></tbody></table></div> <div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox authority-control" aria-labelledby="Authority_control_databases_frameless&amp;#124;text-top&amp;#124;10px&amp;#124;alt=Edit_this_at_Wikidata&amp;#124;link=https&amp;#58;//www.wikidata.org/wiki/Q127768#identifiers&amp;#124;class=noprint&amp;#124;Edit_this_at_Wikidata" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Authority_control_databases_frameless&amp;#124;text-top&amp;#124;10px&amp;#124;alt=Edit_this_at_Wikidata&amp;#124;link=https&amp;#58;//www.wikidata.org/wiki/Q127768#identifiers&amp;#124;class=noprint&amp;#124;Edit_this_at_Wikidata" style="font-size:114%;margin:0 4em"><a href="/enwiki/wiki/Help:Authority_control" title="Help:Authority control">Authority control databases</a> <span class="mw-valign-text-top noprint" typeof="mw:File/Frameless"><a href="https://www.wikidata.org/wiki/Q127768#identifiers" title="Edit this at Wikidata"><img alt="Edit this at Wikidata" src="/upwiki/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/10px-OOjs_UI_icon_edit-ltr-progressive.svg.png" decoding="async" width="10" height="10" class="mw-file-element" srcset="/upwiki/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/15px-OOjs_UI_icon_edit-ltr-progressive.svg.png 1.5x, /upwiki/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/20px-OOjs_UI_icon_edit-ltr-progressive.svg.png 2x" data-file-width="20" data-file-height="20" /></a></span></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">National</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"><ul><li><span class="uid"><a rel="nofollow" class="external text" href="https://id.loc.gov/authorities/sh85128156">United States</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://catalogue.bnf.fr/ark:/12148/cb13574884x">France</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://data.bnf.fr/ark:/12148/cb13574884x">BnF data</a></span></li><li><span class="uid"><a rel="nofollow" class="external text" href="https://www.nli.org.il/en/authorities/987007536424605171">Israel</a></span></li></ul></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"><ul><li><span class="uid"><a rel="nofollow" class="external text" href="https://www.idref.fr/053452216">IdRef</a></span></li></ul></div></td></tr></tbody></table></div></div>'
Whether or not the change was made through a Tor exit node (tor_exit_node)
false
Unix timestamp of change (timestamp)
'1735679669'
Retrieved from "https://en.wikipedia.org/wiki/Special:AbuseLog/39604616"