Global Meliponiculture: Challenges and Opportunities

Apidologie 37 (2006) 275–292 275 c INRA/DIB-AGIB/ EDP Sciences, 2006 DOI: 10.1051/apido:2006027 Review article

Global meliponiculture: challenges and opportunities

Marilda C-La,VeraLuciaI-Fb, DavidWardRc, Anne Dd,TimHe, Ingrid Af,GiorgioC. Vg, Connal Eh,PauloN-Na

a Ecology Dept., Biosciences Institute, S. Paulo University, Brazil b Biology Dept., F.F.C.L. Ribeirão Preto, S. Paulo University, Av. Bandeirantes 3900, 14040-901 Ribeirão Preto, Brazil c Smithsonian Tropical Research Institute, Balboa, Republic of Panama d Australian Native Bee Research Centre, PO Box 74, North Richmond New South Wales 2754, Australia e CSIRO Entomology, 120 Meiers Rd, Indooroopilly Queensland 4068, Australia f CINAT Centro de Investigaciones Apícolas Tropicales, Universidad Nacional, 475-3000 Heredia, Costa Rica g EMBRAPA Amazônia Oriental, PO Box 48, Belém-PA, CEP 66.017-970, Brazil h ARC Plant Protection Research Institute, Private BagX134, Queenswood, 0121, South Africa

Received 19 January 2006 – Accepted 15 February 2006

Abstract – Stingless bees are social bees that live in tropical and subtropical areas of the world. All species produce honey, which has been appreciated by humans since ancient times. Here, the general panorama of meliponiculture is presented. Deforestation and poor management are the main problems faced by this incipient industry. For a proﬁtable meliponiculture, much more biological information is needed, as well as ﬁeld studies in natural conditions. In the near future, we suggest that the successful use of these pollinators will promote the development of new breeding techniques and commercialization possibilities, which must be designed to be sustainable.

meliponiculture / stingless bees / breeding / honey / wax / pollinators / Apidae / Meliponini

Africa, and a little beyond the tropic of Cancer 1. INTRODUCTION in the Northern hemisphere. Stingless bee beekeeping is known as Stingless bees together with honeybees are meliponiculture. This activity, generally un- eusocial insects. Stingless bees have been dertaken by traditional communities, has local known for their honey and pollen production, characteristics according to regional and tradi- but nowadays their role as the providers of tional knowledge. Honey and a waxy material ecosystem services is also recognized. Today, (the cerumen of stingless bee nests is not pure over 600 species in 56 named genera live in wax, but a wax/plant resin mixture) were the tropical and subtropical areas of the world; 400 traditional products. Resin is also occasion- known species exist in the Neotropical region, ally an important income source for the stin- and it is estimated that there are more than gless bee beekeeper, as well as the renting of 100 to be described (Camargo, pers. comm.). ◦ ◦ colonies for pollination service. The stingless bees reach 32 Sto38 S in South America and Australia, respectively, 28◦Sin In this paper, the regional status of meliponiculture worldwide is considered. The use of Corresponding author: M. Cortopassi-Laurino, stingless bees as crop pollinators has opened a [email protected] new economic possibility for meliponiculture. Present address: Biological Science Centre, Santa Large-scale rearing of stingless bees is a cur- Catarina Federal University, 88040-900, Florinopo- rent challenge as it must be aligned with sus- lis SC, Brazil. tainable development. Article published by EDP Sciences and available at http://www.edpsciences.org/apido or http://dx.doi.org/10.1051/apido:2006027 276 M. Cortopassi-Laurino et al.

2. ORIGINS OF MELIPONICULTURE Kayapó universe was taken from social insects, including stingless bees, ants and wasps. Although diverse indigenous peoples lived In their religious ceremonies, meliponine wax in Mexico when the Spanish conquerors ar- or cerumen was used. Camargo and Posey rived in sixteenth century, the ancient Maya (1990) mention that some Kayapó individu- people held stingless bees in high regard and als knew many details of bee behavior, dis- considered them an integral part of social and tribution, nest types and their respective eco- religious life. At that time, stingless bees pro- logical zones. The bee specialists in Gorotire vided a vital small-scale economy, due to their were all shamans: they looked for bee nests honey (used as medicine, as sweetener and during the night (they were not afraid of spir- for mead), their waxes, and resins. The cul- its), and could find them by the noise from tural importance of a single central species, nest ventilation, which they recognized for called Xunan-Kab (Melipona beecheii), was each bee species (see also Kajobe and Roubik, recorded in documents written with ideograms 2006). They also could follow the odor that by the Maya, recorded in their codices. Only bees used to mark nesting sites. The person three of these codices have survived and one, that found the nest became its owner and the Tro Codex, contains most data on bees and could exploit it, using several techniques ac- their meaning to society. cording to the species. Indigenous people use The honey produced by Xunan-Kab was smoke for managing aggressive bees, such considered sacred by the Maya and was also as Oxytrigona, and also use a common toxic traded. Lopes de Gomara (1552) wrote that liana (Tanaecium nocturnum, Bignoniaceae) stingless bee honey occurred in different col- to manage the bees, which causes bees to sleep ors, flavors, and fluidity, and indicated that it after 1 or 2 minutes. All knowledge on bees was an important medicinal product. Honey was passed orally through generations. The wasalsousedtopaytaxestotheAztecs,and traditional way of finding nests by the noise Inca (Cobo, 1653; Roubik, 2000). In Mex- made by worker bees during nest ventilation ico, there was a god of honey, while honey is still a method used by many native people and cerumen of M. beecheii were used in from different communities in Eastern Ama- sacred ceremonies. Even today, in catholic zon. These collectors usually mark the trees churches in Puebla, Mexico, it is possible to with a nest inside during the night. Later, dur- see wax objects offered to God. Besides the ing the day, the nests are collected. modern cultural changes, the stingless bees, The Kayapó exploit stingless bee nests especially the Xunan-Kab, are among the few (Melipona seminigra pernigra, M. melanoven- native organisms that possess a widely ac- ter, M. rufiventris flavolineata, Scaptotrigona knowledged status as a cultural, ecological nigrohirta, S. polysticta, among others) dur- and economic icon (De Jong, 2001; Villanueva ing several consecutive years. When they open et al., 2005b). However, beekeeping with stin- natural nests to collect food, some brood, gless bees has sharply declined in the last half pollen and honey is left for the spirit “Bep- century. Villanueva et al. (2005a) made the kororoti” and this allows recovery of the first census of meliponiculture in Mayan com- colony. Some species like Tetragonisca an- munities in Yucatan, and verified that this ac- gustula, Trigona cilipes and T. dallatorre- tivity is almost extinct (90% of colonies have ana are brought into houses in baskets made been lost in the last 25 years). with banana leaves, and thus are under semi- In Brazil, stingless bees play an important domestication. The Kayapó seem to be aware role in the ethnobiology of the Kayapó (Posey of the role of stingless bees as crop pollinators and Camargo, 1985; Camargo and Posey, (Slaa et al., 2006); they tend to plant bee plants 1990). They named and classified 34 species near their crops to attract the bees. of stingless bees, of which 9 were consid- In Africa, Byarugaba (2004) discussed the ered managed or semi-domesticated. Resin Abayandas pygmy indigenous knowledge on and cerumen were used in their artifacts and stingless bees in the Bwindi forest reserve, as medicine. The natural world model of the Uganda. He explained that Bwindi is a forest Global meliponiculture: challenges and opportunities 277

Figure 1. Vertical hive with special ventilation system for stingless bees of tropical areas (from Giorgio C. Venturieri in www.cpatu.embrapa.br/meliponicultura.htm). island, where different types of forests merge, Stingless bees nest mainly in tree hollows. For and the diversity of species in many groups is such species, it is easier to move and maintain higher than in other parts, yet only six species a nest in artificial wooden hives. Subterranean of stingless bees are found there. Batwa pyg- nests, compared to aerial nests, are difficult to mies, Abayanda, Bakiga and Bafumbira peo- maintain in hives. Considering the variety of ple use the honey of Meliponula ferruginea bee species, a large array of boxes, differing in and other bees for medicine, e.g. to alleviate style and size, have been built and tested. In constipation. Brazil, horizontal hives are much more common than vertical ones. Some beekeepers be- lieve that the vertical boxes supply the bees 3. FROM TREE HOLLOWS with the same conditions they have in the logs. TO STANDARD HIVES Vertical hives with a special ventilation system have been built in hot, tropical areas (Fig. 1). The first step in meliponiculture was the Rational hives made of wood, with a central maintenance of nests around a shelter or home place for brood and surrounding area for honey in the original section of a tree trunk or branch. or pollen pots, as suggested by Nogueira-Neto 278 M. Cortopassi-Laurino et al.

Figure 2. Clay Hives of Scaptotrigona mexicana used by native people in Puebla, Mexico (photo by Figure 3. Tray of honey pots from Melipona com- Cortopassi-Laurino). pressipes fasciculata ready to be collect (photo by G.C. Venturieri). (1997), have been modified by beekeepers. Modifications have also been made in hive vol- colony density of Trigona minangkabau in- ume, adapting them for the size of each rearing creased by 2–5 times. In southern Brazil some species. Ceramic clay hives (Fig. 2) are still beekeepers use empty bottles, with an entrance used by indigenous people in Mexico as well made with cerumen, to attract stingless bee as by some traditional populations in Brazil swarms; such artificial nest sites are attractive (Castro, 2005). Mud is often used to close the to Tetragonisca angustula. Empty hollow logs logs and rudimentary hives in the field, after can be used as a hive, as Ratnieks and Hart management, to prevent enemies from enter- (2001) described for a Maya Indian stingless ing the nest (primarily phorid flies and ants). bee beekeeping: In Northeast Brazil, where meliponiculture “To divide a colony, Benancio takes some is relatively developed, beekeepers try to use combs of emerging brood and honey from the same wood for constructing hives as used one hive and places it in an empty log by the nesting wild colonies, for instance hive which has been smeared inside with Commiphora leptophloeos (Burseraceae) for honey. The newly emerged bees then go Melipona subnitida (Martins et al., 2004). to work and one of the newly-emerged queens takes over. During the honey har- 4. ESTABLISHING A MELIPONARY vest he noted colonies with brood suitable for making the divisions. Colonies can also Stingless bees live in colonies that repro- be obtained from the wild by cutting down duce by a process called swarming, but this is the tree in which they are living, but it is not the same process employed by honey bees rarely done” (Nogueira-Neto, 1997). Colonies are long- Other beekeepers prefer to use special hives lived; there are reports of a nest remaining ac- for keeping their bees. According to the tive more than 50 years in a protected log hive, species harvested and region, hives have dif- but is not known how many swarms they is- ferent models and sizes. Silva et al. (2004) sue during their lifetime, nor how long queens verified that colonies established in standard live, but the expected time required for colony hives produce more honey than in “caboclo” reproduction greatly exceeds queen longevity. or rustic hives, and thus teach small farm- One possible way to obtain nests for estab- ers in the Amazon how to construct standard lishing new meliponaries is to capture natu- hives for their local species. The hives fa- ral swarms by offering suitable artificial nests. cilitate nest multiplication, honey collection, Inoue et al. (1993) found that with the intro- colony thermoregulation, and other necessities duction of artificial nest sites in orchards the for meliponiculture (Fig. 3). Global meliponiculture: challenges and opportunities 279

5. LOGGING AND STINGLESS BEES Stingless bee nests in nature are in danger of depletion by logging practices. In Malaysia (Eltz et al., 2003) and in the Brazilian Amazon (Venturieri, unpubl. data) it has been shown that logging reduces stingless bees nests and, as a consequence, native pollinators, which has implications for forest recovery or restora- tion. Villanueva et al. (2005a) remarked that logging also reduces forest habitat that contains suitable, unoccupied nesting sites. Eltz et al. (2003) considered that the loss of pollinators occurs even if the rules for certified wood Figure 4. Balls of cerumen sold in street fair in management are taken into account. In Brazil the Northeast of Brazil (photo by M. Cortopassi- a Resolution approved by the Environmen- Laurino). tal Ministry National Council (CONAMA) in July 2004, which considers the protection as elsewhere, stingless bee honey is used much and use of wild bees and establishment of more as medicine than as a sweetener. meliponaries. The commercialization of native Some farmers comment that they obtain ex- bees will be allowed only if produced from ceptional honey production from old estab- propagation of hived colonies, or those colo- lished colonies. The average yearly production nized naturally in artificial nests. This should is nearly one or two liters per colony, in stan- help to eliminate the destruction of trees for dard hives (Tab. II). For M. subnitida,anex- extraction of living bee colonies. Nevertheless, ceptional production of nearly 6.0 liters/year the Brazilian government has not yet paid at- occurred in a few colonies out of 80 during tention to the need for general pollinator con- the rainy season in Caatinga regions of Brazil servation among the rules now being estab- (Cortopassi-Laurino and Macêdo, 1998). lished for forest management. ff The tradition of selling stingless bee honey Samejima et al. (2004) studied the e ects of in Brazilian markets has become cost effec- human disturbance (logging and shifting cul- tive only recently. Small jars indicating sting- tivation) on a stingless bee community in a less bee species of origin can be found in local tropical rainforest, in Sarawak, Malaysia. Nest markets in production areas. This honey car- density was positively related to the density ries a higher price than the honey of Apis.In of large trees. The bait results indicated that specialty shops in big cities the price is around some species were abundant in the primary US $40 per liter. forests, whereas others were found mainly in Today, although some beekeepers have disturbed forests. These authors emphasize the more than 500 colonies, they have problems role of nest site availability as a limiting factor preserving the honey because it is watery and for nest density. In Borneo, the distribution of ferments very quickly. In a survey of dif- stingless bee nests is often clumped (Roubik, ferent methods used to collect and conserve 1996, 2006; Nagamitsu and Inoue, 1997; Eltz stingless bee honey, Silva (2005) concluded et al., 2002). that the pasteurization method proposed by Nogueira-Neto (1997) could maintain most 6. HONEY PRODUCTION of the honey’s original characteristics for six months. This method is thus an effective and The extraction of honey is often the sole in- simple way to conserve stingless bee honey come for keepers of stingless bees, followed for commercialization. The way honey is ex- by cerumen (Fig. 4) and resin, but a few years tracted and the use of sanitary measure nec- ago, commercial use of stingless bees in polli- essary play an important role in its final nation began. In all Latin American countries, use and price if selected for table or foods 280 M. Cortopassi-Laurino et al.

Table I. Region and country where stingless bees are being reared.

Region where bee is reared Species of stingless bees Country and or biome M. compressipes manaoensis M. seminigra merrillae Amazon (Amazonas) Brazil1 M. rufiventris paraensis Frieseomellita sp. M. compressipes fasciculata M. compressipes manaoensis M.rufiventris flavolineata M. melanoventer M. seminigra pernigra Amazon (Pará, Maranhão) Brazil1 M. seminigra (Tapajós sub sp.) Scaptotrigona nigrohirta Tetragona clavipes Tetragonisca angustula M. crinita M. eburnea fuscopilosa M. flavolineata Amazon (Acre) Brazil1 M. grandis Tetragonisca weyrauchi M. compressipes fasciculata Amazon (Amapá) Brazil1 M. fulva M. asilvai Scaptotrigona sp. Caatinga* (northeast) Brazil1 M. subnitida M. rufiventris M. scutellaris Atlantic Rain Forest (northeast and southeast) Brazil1 M. quadrifasciata Tetragonisca angustula Frieseomelitta varia M. rufiventris Savanna (northeast) Brazil1 Scaptotrigona spp. M. mandacaia M. beecheii Frieseomelitta nigra Yucatan Mexico2 Scaptotrigona pectoralis Scaptotrigona mexicana Puebla Mexico2 Melipona fasciata Tetragonisca angustula Melipona beecheii Guanacaste/Puntarenas Costa Rica1 Scaptotrigona pectoralis Scaptotrigona luteipennis Melipona eburnea Loreto Tetragonisca angustula San Martin Peru3 Tetragonisca weyrauchi MadredeDios Scaptotrigona aff. polysticta Ibiato/Nguiray Bolivia4 Scaptotrigona postica Scaptotrigona depilis Montevideo Paraguay5 Tetragonisca angustula Tachira Venezuela6 Melipona favosa Trigona carbonaria Trigona hockingsi Queensland and New South Wales Australia1 Austroplebeia spp.

Data from 1: current authors; 2: Cortopassi-Laurino (2002); 3: Rassmussem and Castilho (2003); 4: Pinto pers. comm.; 5: Shevelev (2003); 6: Moreno et al. (2005). Caatinga* a stunt sparse semi-arid region found in Northeast of Brazil. Global meliponiculture: challenges and opportunities 281

Table II. Production of honey/colony/year (liters) in Brazil.

Bee Average Honey (max) Author M. asilvai 1,0 (2,5) Carvalho et al. (2003) M. fasciculata 2,4 (4,4) n = 19 Venturieri et al. (2003) M. mandacaia 2,0 (4,0) Carvalho et al. (2003) M. quadrifasciata 2,0 (5,0) n = 50 Valdemar Monteiro (pers. comm.) M. rufiventris 3,0 (5,0) n = 28 Lima Verde (pers. comm.) M. subnitida 2,5 (5,6) n = 80 Ezequiel Macedo (pers. comm.) M. scutellaris 3,0 (10,0) n ≥ 100 Francisco Chagas (pers. comm.) Scaptotrigona postica 1,5 (3,0) Carvalho et al. (2003) Scaptotrigona nigrohirta 3,8 (12,0) Venturieri & Imperatriz-Fonseca (2000) T. angustula 1,0 (2,7) n = 100 Jean Julien (pers.comm.) industries. One campaign headed by Patricia Honey production also depends on the qual- Vit, from Venezuela, focuses upon honey quality of environment, competition with other ity and authenticity, strategic impacts in honey bee species for floral resources, and bee man- commercialization, and standard analysis of agement. Villanueva et al. (2005a) discussed honey from stingless bees. Although most this point in relation to honey production by honey comes from Melipona and Tetrago- Melipona beecheii. The destruction of natu- nisca angustula, there are many species of ral areas and the increased number of Apis stingless bees that produce honey but re- mellifera colonies had an impact on colony main understudied (Roubik, 1983; Cortopassi- survival and honey production. These authors Laurino and Gelli, 1991; Vit et al., 1994, 2004; suggested that colonies were overexploited for Grajales et al., 2001; Bazlem, 2002; DeMera honey collection and perished without ade- and Angert, 2004; Cortopassi-Laurino, 2005). quate management. In early times honey was Physico-chemical parameters, botanical ori- harvested 5 times a year, and the production gin, and microbiological conditions should be was between 10 to 20 liters a year (Crane and known at least for the potential species, to Graham, 1985; Crane, 1992). develop the needed legislation to allow the commerce and control of stingless bee honey. Nogueira-Neto proposes the name “irahoney” 7. REGIONAL ACCOUNTS for this product (ira means bee in the native OF MELIPONICULTURE Tupi-Guarani language), to distinguish it from the honey produced by Apis mellifera, whose 7.1. Meliponiculture in Mexico control and commercial rules do not apply to stingless bees. The remarkable feature of stingless bee bee- It has been suggested by Vit et al. (2004) keeping in Mexico is that, historically, it was that honeys from all Melipona species could the only sustainable type of beekeeping any- be included under only one legislation. To ac- where in the world. Nevertheless, the situa- cept this idea, we still have to study and to tion has changed in recent years, mainly con- test the honey of stingless bees that are now cerning beekeeping with Melipona beecheii most widely bred in Brazil, which comprise (Villanueva et al., 2005a). at least 10 species (Tab. I). Also important Before providing some details which help is the consideration that the honey should be explain the relationship between stingless bee harvested under optimal conditions, which in- beekeeping and management in Mexico, a clude ripened honey from closed storage pots brief survey of the bees is useful. The gen- and a special extraction machine, due to the era Melipona (which includes the largest stin- high level of humidity and yeasts that are fre- gless bees), Scaptotrigona and Cephalotrig- quently observed in the honeys from stingless ona occur throughout tropical America (see bees. Ayala, 1999). The first two are apparently the 282 M. Cortopassi-Laurino et al. most widely cultivated for their honey. In Mex- timate of approximately 2 colonies per hectare ico, modern meliponiculture is almost entirely (Kajobe and Roubik, 2006). This data can be limited to Melipona beecheii and occasion- used to ask the next question: What kinds of ally to M. solani in the lowlands, and includes colonies are found? M. fasciata and M. colimana in highland areas. The Neotropical data reveal a strong ten- Colonies of Scaptotrigona mexicana are main- dency for most forests to contain many tained in clay jars in part of Mexico, as are colonies of Scaptotrigona but few of S. hellwegeri and S. pectoralis. Cephalotrig- Melipona. Unpublished nest survey data from ona is next to Melipona in size and has three the Yucatan peninsula (Gonzalez-Acereto, species in Mexico. Log hives of C. zexme- pers. comm.) concur with similar data from niae (sometimes called C. capitata in Cen- Panama, Costa Rica, and Bolivia. Scaptotrig- tral America) are kept by Mayan people in the ona have the most numerous colonies of any Yucatan peninsula. stingless bee species with a relatively large An increasing number of studies establish body and colony size. Furthermore, colonies baseline numerical density and abundance of of Melipona have far fewer workers, perhaps bee colonies in natural habitats (review by 70–80% less, than Scaptotrigona.However, Kajobe and Roubik, 2006). Are the most abun- regarding honey stores in the nest, both genera dant bee colonies those most likely to be uti- can have several liters at any one time. It lized by man? Generally, bee nest surveys of can be ascertained that Scaptotrigona and stingless bees or Apis are made in secondary Melipona, Cephalotrigona,andTetragona are or old primary forests, and occasionally when the only Neotropical stingless bees that store there are obviously many bee colonies in more so much honey. Thus, if Mexican habitats disturbed areas. First of all, the number of stin- have been consistent in the forest bee colonies gless bee colonies in older forests, anywhere in which they contain, either large bees, large the world, is not particularly high. Usually one honey stores, or high abundance – and prob- to four colonies exists per hectare. The low- ably a combination of these variables – have est density has been found in old central Ama- had marked influence on the stingless bee zonian forest where within 100 ha only three species selected for meliponiculture. colonies occurred. However, the numbers of An incentive to humans for propagation of species found nesting was nowhere near the Melipona beecheii was probably a result of its number of local species (Oliveira, 2001), thus low nesting density and prized honey stores. one might conclude that many colonies were In traditional Mayan stingless bee beekeeping, too small or too far from the ground-level sur- up to five honey harvests were taken annually vey to be detected. Secondly, surveys in dis- from log hives of M. beecheii in forested ar- turbed habitats (including cleared areas with eas (Villanueva et al., 2005a). In that long-term buildings), or in very young forests, have study in the heartland of traditional Mayan registered the highest densities of colonies meliponiculture, the original 27 colonies in a (Batista et al., 2003). Unfortunately, because single meliponary were derived from a sin- there is not a uniform density of suitable nest- gle parent or ‘mother’ colony. All those man- ing sites (like the tree hollows in a continuous aged colonies have disappeared during the last forest) it is impossible to translate the densities 20 years. A regional decline of over 90% of colonies per building or in semi-natural nest in managed colonies in the past half century cavities (walls, holes, felled trees) to the den- was also documented. An explanation was that sity of nesting colonies on a habitat-wide ba- colonies are no longer propagated, while other sis. Vertical compression of all potential nest- factors, such as waning traditions and hurri- ing sites (normally reaching above 30 to 40 m cane damage, exacerbate the depressing situ- in the forest canopy) to buildings or vegeta- ation by increasing colony mortality. tion that reaches only one-fourth this height, The first techniques used for colony prop- also makes the equivalence of nest surveys on agation in Mexico are unknown, but prob- a ‘per area’ basis doubtful. Nonetheless, the ably involved simple division of the brood mean of the few dozen surveys provides an es- and stored food into log hives. Log hives are Global meliponiculture: challenges and opportunities 283 hollow trees of an abundant hardwood; those (Kent, 1984). However, it was reported that currently in use in Yucatan are ‘silicote’ the an abundance of honey and wax was pro- same tree of the famed chicle trade Manilkara duced around the beginning of the 20th cen- (Sapotaceae). The ends of the hollowed log tury (Kent, 1984). are sealed with a firm, thick plate of the lo- The practice of the stingless bee beekeeping cal limestone, and the cracks sealed with mud. in Pacific Costa Rica (Nicoya) was considered The stone plugs are a unique asset, which re- popular (Kent, 1984). Nowadays, the practice quire only a short time to produce from lime- of the meliponiculture in that region is less stone rocks scattered on the surface of the prevalent. Other areas belonging to Puntarenas entire Yucatan peninsula. Another unique as- and Guanacaste provinces (Santa Cruz, Hojan- set of meliponiculture with Melipona is that cha, Filadelfia, and Miramar) are known for the colonies continuously produce queens. A traditional meliponiculture. In other regions of newly divided colony can thus usually possess Costa Rica meliponiculture is less common, a laying queen soon after it has been made by compared with Guanacaste and Puntarenas. the beekeeper. Whether there had been a tra- For instance, in the Atlantic region of Limón dition of colony propagation using Scaptotrig- and Alajuela, we have observed few stingless ona or Cephalotrigona is unknown. The cur- beekeeper but many of them have received mo- rent practice of dividing Scaptotrigona,often tivation during our recent workshops. using brood from three donor nests, is possibly The stingless beekeepers usually keep their a recent advance in meliponiculture (Stierlin colonies in log hives, generally hanging un- and Szabo, 2004; and Szabo and Stierlin, in der the roof of their houses. Special places for press). The basic trial-and-error development keeping the colonies are built away from the of husbandry techniques had the best chances house, and in exceptional cases there are more for success using the native Melipona that than 40 colonies. The farmers that do not have were readily hived and whose colonies were land or a higher educational degree constitute divided successfully using a minimum of tech- the main social group involved in meliponicul- nology or scientific knowledge. Although stin- ture. gless bee beekeeping has progressed since the Stingless bee beekeeping in Costa Rica ancient Maya worked out some basic princi- has been practiced at a low technical level, ples, many of the current techniques remain almost without equipment, and the type of essentially unchanged. hive used is mainly a hollow log (Arce et al., 1994). The main species reared are 7.2. Meliponiculture in Costa Rica “jicote gato” (M. beecheii), “jicote bar- cino” (M. fasciata)(Melipona costaricensis), Stingless beekeeping in Central America “mariquita o mariola” (Tetragonisca angus- originated among the Maya of the Yucatan tula), “soncuano” (Scaptotrigona pectoralis Peninsula and spread to other Central Amer- pectoralis) and “tacanique” (Scaptotrigona ican Indian cultures (Kent, 1984). Before the luteipennis) (Ramírez and Ortiz, 1995). The discovery and the conquest of American con- breeding of Frieseomelitta nigra, T. fulviven- tinent by Europeans (indigenous peoples im- tris, Lestrimelitta, and Plebeia tica is less migrated across the Bering Strait at least common. Lestrimellita, a robber bee that does 15 000 years ago), rearing stingless bees was not visit flowers, is not suitable for stingless an important part of the commercial and ali- bee beekeeping (Santana et al., 2004). mentary customs of many indigenous cultures Arce et al. (1994) reported from a sur- of America. The honey of these bees was used vey of 40 stingless bee beekeepers (Gua- as food and medicine, and the preparation of nacaste Province) a total of 9 different do- other products based on cerumen was also im- mesticated species: M. beecheii, M. fas- portant (Kent, 1984; Sommeijer, 1996). As ciata (now known as M. costaricensis) in other Mesoamerican areas, stingless bee- M. fuliginosa, Tetragonisca angustula, Scap- keeping in Costa Rica did not play an impor- totrigona pectoralis, Cephalotrigona capi- tant role in the religion of indigenous cultures tata, Nannotrigona perilampoides, Trigonisca, 284 M. Cortopassi-Laurino et al.

Frieseomelitta, Oxytrigona mellicolor,and Veen (1995), in Guanacaste Province Costa Tetragona. Some farmers obtained production Rica, where fewer species were collected at of up to 6 liters and, in the case of M. beecheii, higher altitudes. It is well known that sting- the production average was 2.63 liters per hive. less bee species richness declines dramatically According to van Veen et al. (1990), above 1000 m in Central America. They are meliponiculture in Costa Rica is practiced ba- more abundant at 200–500 m and rare at 700– sically in two ways: (1) maintaining the nests 1000 m. van Veen et al. (2004) studied in Costa in tree trunks, from which the honey, by a lat- Rica the production of queens and drones for eral opening, is extracted, generally used for M. beecheii in relation to food storage, brood M. beecheii; and (2) maintaining the colonies and adult populations. They found that the in small boxes, pieces of bamboo or hollow colonies grow at the end of the rainy season, gourds, ordinarily with T. angustula. Although which is a period characterized by poor forag- meliponiculture occurs throughout Costa Rica ing conditions. On the other hand they found and even in urban areas (Sommeijer, 1996), the a significant correlation between the amount local people do not understand the reproduc- of pollen stored and the production of queens tive biology of the bees. This is reflected in the and drones. Finally, more action should be fear the people have in dividing nests. When taken to continue the work initiated by Wille carried out in a haphazard way, nest division (1961) on the biology, biodiversity conserva- results in parasite attack (phorid flies) and loss tion and management of the stingless bees in of the colony. Costa Rica. In relation with the design and dimensions A cooperative programme for research, ex- of hive boxes, van Veen et al. (1993) recom- tension and training related to tropical bees mended for M. beecheii a hive with a volume (PRAM) involving Utrecht University and the of 10 liters with internal dimensions of 15 cm Universidad Nacional de Costa Rica began in height, 15 cm width and 45 cm length. For 1990 (www.una.ac.cr/cnat/conven2.htm) and T. angustula the recommended box measures more recently El Salvador joined the group. 15 cm × 15 cm × 20 cm long, which gives a volume of 4.5 liters. In its application we have 7.3. Meliponiculture in Brazil observed that the stingless beekeepers mod- ify the dimensions of hives according to the Meliponiculture in Brazil was studied and species and the size of the colony. summarized by Nogueira-Neto (1953, 1970, Thanks to different extension projects 1997, 2002) and Kerr et al. (1996), who pro- and workshops carried out by the Tropical moted development of this activity. Besides Beekeeping Research Center (CINAT) Uni- advancing the activity with guidebooks, which versidad Nacional, Heredia, Costa Rica the provide a unique literature in Portuguese, these interest in this honey and in stingless bees has investigators’ research resulted in a group of increased over the past few years. Today com- scientists that also work with meliponiculture merce of this honey commands high prices; and Brazilian stingless bees. one liter sells for US $20–50 and small con- Theories of genetic variability (Kerr and tainers of 10 mL cost US $2.0 due to an in- Vencovski, 1982; Kerr, 1985) predicted that to creasing interest in its medicinal use (mainly successfully breed stingless bees it was nec- used against cataracts). essary to have at least 44 colonies, to avoid Research carried out by CINAT confirmed diploid males created by excessive inbreed- the importance of Nannotrigona perilam- ing in the population. From 1990 to 2001, poides and Tetragonisca angustula as pollina- Nogueira-Neto (2002) performed 287 colony tors of the ornamental plant Salvia farinacea divisions, starting with 28 foundress colonies. (Slaa et al., 2000). This stingless bee can be Seven species of stingless bees (and one sub- considered an alternative to honey bees for species) were inbred in five localities inside commercial crop pollination. Also, there is a their extensive native habitats, and in two lo- study on the influence of altitude on the distri- calities far from these habitats. On the whole, bution of stingless bees by Ortiz-Mora and van the number of colonies increased and the loss Global meliponiculture: challenges and opportunities 285 of colonies over the years was small. Diploid also adjusted the measurements of the box for males were not found in nests after several di- each bee species. visions. This investigator also evaluated how The way a stingless bee beekeeper divides long a queen lived in some species, by mark- and propagates colonies varies according to ing physogastric queens with a dot of perma- species and husbandry knowledge (Nogueira- nent paint. Most colonies changed queens ev- Neto, 1997). A curiosity to mention here is the ery year. The experiments continue, and it is beekeeper that successfully divides the nests clear that we need more research on the genet- of M. scutellaris using a large knife, cutting ics of stingless bee populations. Furthermore, the nests vertically in two parts. Three main the survival of small populations is a conser- ways of splitting the nests are known, using vation concern. Nogueira-Neto is testing man- one, two or more colonies in order to form agement techniques that are locally adapted to a new one, although the adult workers come stingless bees. Phorid flies are controlled with from only one nest (Cortopassi-Laurino and the use of vinegar traps, first constructed by Macêdo, 1998). Imperatriz-Fonseca. These are often very ef- Most Brazilian stingless bee beekeepers are fective in control. Nowadays they can be used members of associations and cooperatives, and inside the nest, as well as outside. some of them have their own homepages. The In early times, beekeepers collected oldest is CAPEL, in Pernambuco state. Blogs stingless bee nests in nature, as did their or discussion websites are also available for relatives and parents. More recently they are stingless bee beekeepers, generally used for learning how to divide the colonies, as well beginners, which facilitates the spread of genas become more conscious of environmental eral beekeeping techniques. concerns. In Brazil, with few exceptions, bee- In Brazil, a dozen beekeepers started rear- keepers maintain a low technical level, almost ing stingless bees as part of social and govern- without equipment. The harvest of honey is mental projects. The largest is Iraquara Project often made by shaking the hives upside-down, in Amazonas state, which produced 3 tons of thereby destroying young larvae that fall from honey in 2004 (Villas-Bôas and Malaspina, their provisions, and also attracting phorid 2005). In the last three years, the Brazilian flies that damage and parasitize the nest. Agricultural Research Company (EMBRAPA) Honey, new colonies and nuclei (small in- has trained more than 450 rural workers, stu- cipient colonies) are the main products of dents, and technicians in meliponiculture in Brazilian meliponiculture. Stingless bee bee- Northeastern Amazon. After implementation keeping as a hobby and for educational pur- of these courses, the number of bee boxes in- poses is another strong reason to keep these creased from 281 in 2003 to over 900 in 2005 bees in urban or rural areas. by the same group of beekeepers. Many of Surveys carried out in 2001 by Rosso et al. these beekeepers also commercialized part of (2001). in Brazil showed that meliponiculture their colony divisions, and thus helped spread is growing rapidly, and for most people it is the meliponiculture practice. a secondary economic activity. Most of these Research with stingless bees for use as beekeepers rear stingless bees from their nat- pollinators exists in Brazil. Malagodi-Braga ural regions, but for only one person surveyed and Kleinert (2004) verified the importance of the sale of new colonies or nuclei, and honey, Tetragonisca angustula as effective pollinators was the only family income. One beekeeper (a for some strawberry cultivars. Additional stud- breeder of Melipona scutellaris) increased the ies concerning stingless bees as pollinators are number of his colonies from 200 (in 2001) to in place (Venturieri et al., 2005). Slaa et al. 750 (in 2006). Another breeder from Central (2006) reviewed this point. Amazon, keeping Scaptotrigona sp. and other five species of Meliponini, increased the num- 7.4. Meliponiculture in Africa ber of colonies from 45 in 2000 to 350 in 2005 after adoption of vertical rational hives (simi- Stingless bee taxonomy has recently been lar to the one shown in Fig. 1). This beekeeper reviewed by Eardley (2005). There are over 286 M. Cortopassi-Laurino et al.

20 stingless bee species in Africa, but A survey of stingless bee keeping in Aus- much work remains to be done. Stingless tralia in 1998–99 concluded that 62 bee- bee studies were conducted by Bassindale keepers kept 317 hives of stingless bees for (1955), who studied Hypotrigona; Darchen crop pollination (Heard and Dollin, 2000). (1972) studied several species, and also made However, only one beekeeper, with 65 hives, films on African stingless bees, available was offering crop pollination service. Now, in in www.cerimes.education.fr. Breeding and 2005, at least six beekeepers offer stingless bee meliponiculture in Africa was considered by services for crop pollination, usually as their Portugal-Araujo (1957) in Angola, but this secondary financial activity. Those beekeep- practice did not evolve in subsequent years. ers have 15 to 150 hives. Many crop grow- African stingless bee honey is mostly col- ers are also purchasing their own stingless bee lected by harvesting from feral colonies, which colonies. The main species used for crop pol- subsequently destroys them. A few commu- lination are Trigona carbonaria and T. hock- nities use hollow logs or clay pots as hives, ingsi. and they harvest the honey in a more sustain- Macadamia nut (Macadamia integrifolia), able way. In at least Tanzania, and Angola, native to eastern Australia, is the most com- meliponiculture exists and an interest to de- mon crop pollinated with stingless bees. With velop meliponiculture has been identified in over 800 farms, Australia is the world’s largest Ghana, Kenya, Botswana and South Africa. In producer of macadamia nuts. Stingless bees Angola, Meliponula bocandei, the largest stin- mainly collect pollen from the flowers and gless bee, produces 10–15 kg of honey in a are efficient pollinators (Heard, 1994). Many season (Armor, 2005). other crops may be pollinated by stingless bees (Heard, 1999) and in Australia, successful results for lychee, melon and avocado have been 7.5. Stingless beekeeping in Australia reported. Wild stingless bees often provide a free pollination service (Heard and Exley, In Australia 20 years ago, few beekeepers 1994) but where the bees are rare because of had more than one or two stingless bee hives. land clearing, hives need to be introduced. Today, there is a growing interest in nest con- Australian stingless bees prefer to forage servation. Crop pollination services are devel- close to their hives, thus tend to stay within the oping, honey and cerumen are being harvested, crop area better than does Apis (Carter, pers. and a stingless bee industry is being developed comm.). One beekeeper reports that the effec- by the indigenous community. Bees also are tive foraging range of stingless bees within being used for public education. a macadamia crop is 25 m (Adcock, pers. There is a growing interest in conserving comm.). Stingless bees also cope better than wild stingless bee colonies. Many individual Apis in crops covered by netting (Felhaber, beekeepers relocate nests found in fallen trees pers. comm.). However, stingless bees can be and artificial sites. In Ipswich, Queensland, a quite sensitive to pesticides, which can re- rescue group led by C. Heather actively iden- sult in substantial bee losses (Grosskopf, pers. tifies land about to be cleared for development comm.). and seeks permission to remove nests that Demand for crop pollination services with would otherwise be destroyed. Over a seven- stingless bees in Australia should continue year period this group has rescued 200 sting- to increase as farmers become informed of less bee nests. Once wild colonies are captured the benefits and more hives become avail- in hives, they can be allowed to reproduce. In- able. Small hive beetles (Aethina tumida) deed, the increase in popularity of the bees that are progressively spreading through New in Australia over the last 20 years is partly South Wales and Queensland may deplete due to the development of a husbandry method feral Apis populations, increasing demand for (Heard, 1988). Colony multiplication is infre- paid pollination services. Varroa mite is also quent, typically every one to two years, but threatening to enter Australia and will also re- population growth is exponential. duce Apis populations. Global meliponiculture: challenges and opportunities 287

In addition to crops, native bushland re- Brisbane Botanic Gardens has a demonstration quires pollination services. The pollinators log nest, which they use as part of their educa- may have been driven extinct by human tion program. Workshops on stingless bees at induced pressures. Some interest is being nature education centers are growing in pop- expressed by managers of bushland to re- ularity. The Australian Native Bee Research introduce stingless bees into conservation ar- Centre has a site at www.aussiebee.com. eas. Interest in production and marketing of Australian honey is growing. New hive de- 7.6. Meliponiculture in Asia signs that are suited to the extraction of honey and cerumen are being developed. Interest in Southeast Asia has several honey bee the honey is being expressed by restaurants species and about 45 stingless bee species, that promote native foods. We estimate that the but nowhere is there a standard practice called total current annual production in Australia is meliponiculture. Thailand, like Viet Nam, is a extremely small, possibly less than 100 kg, but very long country, extending for over 1000 km there is potential for rapid growth. The mar- south to north. In the south, there are five Apis ket price of the honey is currently about AU species and at least 30 stingless bee species, $50 per kg, wholesale. To develop commer- while in the north, there are four Apis and less cially, the price needs to increase, because the than 10 stingless bee species. The southern production per hive is low and the cost of pro- Thai name for meliponines is “Oong”, while in duction is high. Challenges include minimiz- the north, they are called “Channa Long”. The ing fighting swarms and nest parasites (Dollin, honey bees are called “Png”. The local peo- 2002). ple only use stingless bee honey for “medicinal Russell and Janine Zabel have been work- purposes” because they say it is a lot of work to ing with the indigenous community of Au- remove the honey from a tree or subterranean rukun in far north Queensland since 2002 to nest of stingless bees. In contrast, they can ob- develop a stingless bee industry. Currently tain several liters of honey from a nest of Apis they have hived over 130 colonies, mostly dorsata or Apis cerana, and many nests of the T. hockingsi. Stingless bee honey, known to former can be harvested on a single tree. There indigenous people as ‘Sugarbag’, is a major is also traditional use of wax from the nests product. Cerumen is also collected for mak- of Apis dorsata in batik fabric design, so that ing traditional artifacts (Zabel, pers. comm.). there is a steady demand for honey hunting, A major potential use for the cerumen is to using both the comb and the honey, from the form the mouthpiece of the “didgeridoo”, a giant honey bee. The local stingless bees have traditional musical instrument. Until recently, so little to offer, by comparison, that it does cerumen was very difficult to obtain and honey not occur to anyone that the keeping of sting- bee wax was used as a substitute. However, less bees in hives is worth the trouble. Sting- stingless bee cerumen is the authentic and su- less bee beekeeping for pollination service is perior product. Demand is expected to grow only now beginning to take root in southern rapidly as awareness increases. Asia (in India) and in SE Asia (Malaysia and Stingless bees are attractive subjects for the Philippines). public nature education. Museums, exhibi- tions, gardens, and schools in Australia are in- creasingly using social bee colonies to illus- 8. LARGE-SCALE BEEKEEPING trate the beauty, complexity and fascination of WITH STINGLESS BEES nature. Topics such as sociality, parasitism and pollination can be demonstrated using these In Brazil a large quantity of colonies has yet insects, which are gentle and easy to keep. The to become available, for instance, to be sold Melbourne Museum maintains a permanent for agricultural use or for use as greenhouse indoor public display by only occasionally ex- pollinators. To have a system similar to that changing the colony with another outside. The of Bombus terrestris (Velthuis and van Doorn, 288 M. Cortopassi-Laurino et al.

2004), additional information on queen pro- In all places where meliponiculture devel- duction is needed, as well as controlled mating ops, bottlenecks for their improvement include and colony propagation techniques under arti- how to keep and conserve their honey, how to ficial conditions. Until now queen emergency rear them in large quantities, how to prevent cells for stingless bees have been found only colonies from being contaminated by agricul- in Frieseomellita varia (Faustino et al., 2002), tural pesticides and maintain the bees, how to which allow the propagation of many colonies use their services and conserve their popula- in this genus. A similar finding for other stin- tions, and how to provide qualified informa- gless bees species, such as Frieseomelitta, tion and training in all levels. which construct brood cells in clusters, would be nothing short of revolutionary. Generally a large number of gynes (po- Résumé – Méliponiculture globale : défis et tential laying queens) is available only for chances. Les abeilles sans aiguillon (Apidae, Meli- Melipona bees, where gynes, workers and ponini) sont des insectes sociaux qui vivent dans les males emerge from similar brood cells. Re- régions tropicales et subtropicales du globe. L’importance de leurs produits en Amérique latine, prin- cent review (see Santos et al., 2006) shows that cipalement le miel (utilisé comme édulcorant et Melipona queens are produced in a smaller comme remède), est souligné. Il existe des docu- number than presumed before, and only a few ments écrits, principalement les codex mayas, qui number of other stingless bees has been stud- montrent le rôle central des abeilles sans aiguillon ied in relation to gyne and male production dans la vie religieuse et culturelle. Melipona beecheii est la première abeille dont l’élevage en grand (Chin and Sommeijer, 2005). Another per- a été mentionné : elle est élevée dans des troncs spective is related to miniature gyne produc- d’arbre regroupés en “ méliponaires ”, ruchers pou- tion (Ribeiro et al., 2006). vant réunir plusieurs centaines de leurs colonies. Maintenir des abeilles dans des troncs d’arbre a été la première étape de l’élevage dans plusieurs régions du monde où les abeilles sans aiguillon 9. CONCLUSIONS sont indigènes. L’étape suivante a consisté à diviser AND RECOMMENDATIONS les colonies. L’utilisation moderne de ces abeilles va les considérer surtout comme fournisseurs de services pour l’environnement et comme pollinisa- Stingless bee breeding is a significant ac- teurs. Il s’agit d’un nouveau défi et d’une occasion tivity that permits sustainable agricultural de- pour approfondir leur connaissance et leur utilisa- velopment and wildlife conservation, but for tion et pour travailler à leur sauvegarde. many species it requires specific management On trouve la méliponiculture principalement dans techniques and intact natural environments. les pays où cette activité est traditionnelle. Les colonies passent toujours de génération en génération, Villanueva et al. (2005a) mentioned the con- mais leur importance dans la vie moderne diminue. cerns that no new colonies were available for D’autres sources de sucre, comme la canne à sucre Melipona beecheii in the studied area; bees are et le miel d’Apis mellifera, sont meilleur marché. dying from lack of food; colonies have less On n’a pas mis au point de techniques de sélec- and less honey, and colonies are diminishing tion des abeilles sans aiguillon et seules quelques espèces indigènes sont élevées, dont un genre très in size. These are likely to be the same prob- prometteur, Scaptotrigona. Ces abeilles sont égale- lems faced by meliponiculture elsewhere in the ment élevées dans des ruches en argile au Mexique, world. ainsi qu’au Brésil et en Afrique. Le déclin de la mé- Stingless bee beekeeping, at least in some liponiculture au Mexique et la survie de M. beecheii areas, is disappearing. The younger potential suscitent des inquiétudes. La situation au Costa- ff Rica est discutée. Ce pays améliore la méliponicul- beekeepers must join the e ort in maintain- ture par l’éducation à l’environnement et un travail ing the population of stingless bees. Basic avec les petits agriculteurs. Un groupe technique biology studies are needed, mainly concern- issu d’une coopération internationale a été mis en ing the exploitation of natural resources by place afin de fournir un appui scientifique à cette these bees and medium-long-term studies. The activité. Le Brésil possède une tradition apicole avec les abeilles sans aiguillon. Les indigènes uti- new meliponiculture will be organized to ide- lisaient ces abeilles, et cette connaissance a traversé ally provide agriculture and wildlife with the les générations, comme chez les indiens Kayapo ecosystem services essential for pollination. récemment étudiés. La méliponiculture s’améliore Global meliponiculture: challenges and opportunities 289

au Brésil ; les apiculteurs sont généralement re- werden nur einige native Arten gehalten, einschlies- groupés en associations et échangent leurs connais- slich die in unserem Zusammenhang interessante sances pratiques. Le miel est le principal produit, Gattung Scaptotrigona. Ausser in Klotzbeuten wer- il est cher et apprécié comme remède. En Afrique den diese Bienen in Mexiko, Brasilien und Afrika et dans le sud-est asiatique, l’apiculture avec les auch in Tonbeuten gehalten. In der vorliegenden Ar- abeilles sans aiguillon est confrontée à l’apiculture beit zeigen wir, wie die traditionelle Meliponikul- avec les abeilles du genre Apis,plusefficace pour tur in Mexiko abnimmt und wie sich das auf das le commerce. Dans ces régions la connaissance des Überleben vom M. beecheii auswirken kann. Wir abeilles sans aiguillon en est à ses débuts. En Aus- diskutieren ausserdem die Situation in Costa Ri- tralie les services commerciaux de pollinisation des ca, wo die Meliponikultur in Zusammenhang mit cultures commencent à être fournis par des apicul- der Umwelterziehung und Arbeiten mit Kleinbau- teurs qui ont jusqu’à 150 ruches. Le macadamia (qui ern zunehmend Bedeutung gewinnt. In diesem Land fournit la noix) est la culture plus courante à être wurde über eine internationale Zusammenarbeit ei- pollinisée et la production de miel et de cérumen ne Expertengruppe aufgebaut, die diese Aktivitä- est en augmentation. ten wissenschaftlich unterstützt. Brasilien hat ei- Partout où la méliponiculture se développe, les gou- ne Tradition in der Haltung Stachelloser Bienen. lets d’étranglement sont : (i) comment conserver le Eingeborenenstämme nutzen diese Bienen und ge- miel des abeilles sans aiguillon, (ii) comment les ben, wie kürzlich für die Kayapo-Indianer gezeigt, élever en grandes quantités, (iii) comment faire des dieses Wissen über Generationen hinweg weiter. traitements insecticides et maintenir les abeilles, Die Meliponikultur in Brasilien ist in einer Auf- (iv) comment utiliser leurs services et préserver schwungsphase und Imker sind in der Regel in Im- leurs populations et (v) comment fournir une in- kergemeinschaften assoziiert und tauschen ihr prak- formation de qualité et une formation à tous les ni- tisches Wissen aus. Der Honig Stachelloser Bie- veaux. nen ist teuer und wird gerne auch als Medizin ver- abreicht. Afrika und Südostasien sind Regionen in denen die Haltung Stachelloser Bienen mit den in méliponiculture / abeille sans aiguillon / élevage / / / / / wirtschaftlicher Hinsicht wichtigeren Honigbienen pollinisation miel cire Apidae Meliponini konfrontiert sein sollte, und in diesen Regionen steckt die Kenntnis Stachelloser Bienen noch in den Kinderschuhen. In Australien werden Stachellose Zusammenfassung – Globale Meliponikultur: Bienen jetzt von Imkern mit bis zu 150 Völkern zur Herausforderungen und Chancen. Stachellose Bestäubung kommerzieller Früchte eingesetzt, vor allem Makadamia. Zudem ist die Produktion von Bienen sind soziale Bienen, die in den Tropen und ff Subtropen der gesamten Welt verbreitet sind. Die Honig und Cerumen im Anstieg begri en. Bedeutung ihrer Produkte, vor allem Honig (als An allen Orten, an denen die Meliponikultur in Aus- Süssstoff und Medizin) ist besonders in Südameri- breitung ist, liegen die Engpässe ihrer Entwicklung ka deutlich. Schriftdokumente, vor allem die Maya darin, wie der Honig Stachelloser Bienen konser- Codices, zeigen die zentrale Rolle Stachelloser Bie- viert werden kann, wie Völker in grosser Anzahl nen im religiösen und kulturellen Bereich. Melipo- gezüchtet, wie Pestizide appliziert und die Bienen na beecheii war die erste Bienenart, die in grossen gehalten werden können, wie ihre Dienste genutzt Völkerzahlen gehalten wurde. Es wird von Melipo- und Populationen konserviert werden können und narien mit hunderten von Kolonien in Baumklötzen wie qualifizierte Information und Schulung auf al- berichtet. Die Haltung von Bienen in Klotzbeuten len Ebenen verbreitet und betrieben werden kann. war der erste Schritt zu ihrer landwirtschaftlichen Nutzung in vielen Teilen der Welt, in denen Stachel- lose Bienen vorkommen. Der nächste Schritt war Meliponikultur / Stachellose Bienen / Bienen- die Teilung der Kolonien. Die Nutzung im moder- zucht / Honig / Wachs / Bestäuber / Apidae / Me- nen Sinn zielt vor allem auf die Rolle dieser Bienen liponini als Lieferanten von Ökosystemdiensten, wie z.B. Bestäubung. Dies stellt eine neue Herausforderung dar und liefert auch Chancen für ihr Studium, ihre Nutzung und Konservierung. Meliponikultur findet man heute vor allem in den REFERENCES Ländern in denen diese Aktivität eine Tradition hat. Arce A.H.G., van Veen J.W., Sommeijer M.J., Ramírez Mexiko war das erste Land. Kolonien werden hier A.J.F. (1994) Aspectos técnicos y culturales immer noch über Generationen hinweg weiterge- de la crianza de abejas sin aguijón (Apidae: geben, aber ihre Bedeutung im modernen Leben Meliponinae) en Costa Rica, in: Veen Van nimmt ab. Andere Zuckerquellen wie Rohrzucker J.W., Arce H.G., Ortiz R.A. (Eds.), Apicultura: und Apis mellifera Honig sind im allgemeinen ko- Estrategias para el manejo adecuado de las col- stengünstiger. 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