**5. Pollination for agricultural productivity and ecological services**

As human population grows, the demand for food is increasing every year. The increase in productivity to improve food security without harming the natural environment, and making the improved productivity sustainable for future generations, is a major challenge [30]. Farmers try to improve the quality of their produce to obtain optimum prices. Pollination is one approach to achieve that goal. Incomplete flowers need pollinators for their fruit sets. Even for self-fertile flowers, cross-pollination is still needed for improved production and better quality of seeds and fruits [31]. Thus, beekeeping not only can improve income but also can increase food security [32].

Stingless bees are candidates for commercial and natural pollination. They are highly diverse and abundant and inhabit the tropical and subtropical parts of the world [33]. In SE Asia there are 68 species from 14 genera [34]. Stingless bees form perennial colonies from which they forage year-round with a variety of body size, nest structure and position, and ecological habitats allowing for selection of the most suitable stingless bee species for a given crop species and crop breeding system [33]. Stingless bees are true generalists that visit a vast array of plants [35]. However, at the individual level, stingless bees tend to specialize in a single flower [23, 36]. Indeed, this combination of traits between generalist and specialist is a characteristic that makes them one of the best contributors to pollination for many crops and wild plants [37].

Native tropical plant crops such as coffee and cacao show a mutually beneficial interaction with stingless bees. The genus *Coffea* (Rubiaceae) is native to tropical and subtropical Africa. The two coffee species, *Coffea arabica* L. and *Coffea robusta*, are grown throughout India and Southeast Asia. Even though *C. arabica* is tetraploid and self-compatible [31], the benefits of cross-pollination are still distinctly shown by producing 30% higher fruit set than autonomous self-pollination [38] and 25% higher fruit weight [39]. During the mass flowering season of coffee, the honeybees and stingless bees are dominant. However, they are often absent in the scattered flowering which occurs frequently in tropical plant blooming behavior. Coffee plantations on the edge of forests are more likely to receive a significant benefit from stingless bees which are more abundant within 600 m of the forest margin than other social bees [40] and other insect pollinators. The study of pollinators in coffee plantations in Sulawesi, Indonesia, found three honey bee species and four stingless bee species as the main pollinating species [40].

For strawberries, it is also reported that the yield is increased using stingless bees. Because the strawberry flower seems not to be attractive to honeybees, stingless bees are the preferred choice for strawberry pollination in greenhouses. Most strawberry cultivars are hermaphrodite and self-fertile, but the anther maturation

**183**

**Figure 11.**

*Southeast Asian Meliponiculture for Sustainable Livelihood*

and stigma receptivity may vary highly in spatial segregation [31] which makes pollination helpful to increase productivity. In Asia, most of the studies of stingless bee pollination have been conducted in Japan [37, 41]. Strawberry pollination with stingless bees has also been seen in Nan, Thailand (O. Duangphakdee, personal communication). In many instances, the fruit and seed crop orchards at the edges of forests noted that natural pollinators are considered in increasing crop yield. Since 1990, SE Asia has suffered deforestation of 33.2 million hectares or 7.6% of the land area [42], which is the highest relative rate of deforestation of any major tropical region. As a consequence, deforestation and forest fragmentation may contribute to declines in crop pollinator populations. Several studies have been conducted to examine the effect of forest proximity on plant pollination ecology. The evaluation of flower visitor diversity, frequency, and fruit set for three crop species has been conducted in mixed fruit orchards of rambutan (*Nephelium lappaceum* L.), durian (*Durio zibethinus* L.), and mango (*Mangifera indica* L.) [43] in southern Thailand. This study compared 10 pairs of orchards that are located at <1 km and >7 km away from the forest edge. Stingless bees were the main visitors for 70.9% of total flower visits on rambutan (**Figure 11**). The distance from forest edge and location of natural stingless bee colony influenced the fruit set of rambutan significantly [43]. Stingless bees were significantly (two times) more frequent on rambutan flowers

In the case of durian and mango, even though stingless bees have been observed

as flower visitors, the number of fruit sets was significantly influenced by bats (durian) and flies (mango).The distance to the forest did not affect the fruit yield of these two crops [43, 44]. The evidence suggests that the forest is an insect pollinator reservoir. The conservation of natural habitats surrounding a crop orchard is strongly recommended to maintain a population of forest-insect pollinators in

As generalists, stingless bees forage in vast array of plant taxa. A study of pollen foraging and resource partitioning of stingless bees throughout year-round

*A linear regression plot for the number of rambutan fruit sets and insect visitation frequency to rambutan* 

*flowers in a mixed fruit orchard in Southern Thailand [43].*

*DOI: http://dx.doi.org/10.5772/intechopen.90344*

nearer to the forest.

natural habitats [45].

#### *Southeast Asian Meliponiculture for Sustainable Livelihood DOI: http://dx.doi.org/10.5772/intechopen.90344*

*Modern Beekeeping - Bases for Sustainable Production*

markets in other SE Asian countries.

food security [32].

crops and wild plants [37].

of eastern Thailand have the most developed commercial meliponiculture for pollination and honey production at approximately 5000 hives [27, 29]. The selling price of Thai stingless bee honey is 1200–1500 THB (\$37–\$47 USD) per kilogram which 10 times and 3 times higher in price of honey from Thai produced *A. mellifera* and wild *Apis* (*A. florea*, *A. dorsata*, and *A. cerana*), respectively. The propolis and wax cerumen are additional active markets in Thailand with per kilogram returns of 1500–2000 THB (\$47–\$62 USD). In total, [29] evaluated that stingless bee hive products added 5.76 million THB (\$177,500 USD) to the regional economy in 2014. Because of the increase in meliponikeeping, a number of new stingless bee beekeepers in Thailand are increased gradually to support in-country and international

**5. Pollination for agricultural productivity and ecological services**

As human population grows, the demand for food is increasing every year. The increase in productivity to improve food security without harming the natural environment, and making the improved productivity sustainable for future generations, is a major challenge [30]. Farmers try to improve the quality of their produce to obtain optimum prices. Pollination is one approach to achieve that goal. Incomplete flowers need pollinators for their fruit sets. Even for self-fertile flowers, cross-pollination is still needed for improved production and better quality of seeds and fruits [31]. Thus, beekeeping not only can improve income but also can increase

Stingless bees are candidates for commercial and natural pollination. They are highly diverse and abundant and inhabit the tropical and subtropical parts of the world [33]. In SE Asia there are 68 species from 14 genera [34]. Stingless bees form perennial colonies from which they forage year-round with a variety of body size, nest structure and position, and ecological habitats allowing for selection of the most suitable stingless bee species for a given crop species and crop breeding system [33]. Stingless bees are true generalists that visit a vast array of plants [35]. However, at the individual level, stingless bees tend to specialize in a single flower [23, 36]. Indeed, this combination of traits between generalist and specialist is a characteristic that makes them one of the best contributors to pollination for many

Native tropical plant crops such as coffee and cacao show a mutually beneficial interaction with stingless bees. The genus *Coffea* (Rubiaceae) is native to tropical and subtropical Africa. The two coffee species, *Coffea arabica* L. and *Coffea robusta*, are grown throughout India and Southeast Asia. Even though *C. arabica* is tetraploid and self-compatible [31], the benefits of cross-pollination are still distinctly shown by producing 30% higher fruit set than autonomous self-pollination [38] and 25% higher fruit weight [39]. During the mass flowering season of coffee, the honeybees and stingless bees are dominant. However, they are often absent in the scattered flowering which occurs frequently in tropical plant blooming behavior. Coffee plantations on the edge of forests are more likely to receive a significant benefit from stingless bees which are more abundant within 600 m of the forest margin than other social bees [40] and other insect pollinators. The study of pollinators in coffee plantations in Sulawesi, Indonesia, found three honey bee species and four

For strawberries, it is also reported that the yield is increased using stingless bees. Because the strawberry flower seems not to be attractive to honeybees, stingless bees are the preferred choice for strawberry pollination in greenhouses. Most strawberry cultivars are hermaphrodite and self-fertile, but the anther maturation

stingless bee species as the main pollinating species [40].

**182**

and stigma receptivity may vary highly in spatial segregation [31] which makes pollination helpful to increase productivity. In Asia, most of the studies of stingless bee pollination have been conducted in Japan [37, 41]. Strawberry pollination with stingless bees has also been seen in Nan, Thailand (O. Duangphakdee, personal communication). In many instances, the fruit and seed crop orchards at the edges of forests noted that natural pollinators are considered in increasing crop yield. Since 1990, SE Asia has suffered deforestation of 33.2 million hectares or 7.6% of the land area [42], which is the highest relative rate of deforestation of any major tropical region. As a consequence, deforestation and forest fragmentation may contribute to declines in crop pollinator populations. Several studies have been conducted to examine the effect of forest proximity on plant pollination ecology. The evaluation of flower visitor diversity, frequency, and fruit set for three crop species has been conducted in mixed fruit orchards of rambutan (*Nephelium lappaceum* L.), durian (*Durio zibethinus* L.), and mango (*Mangifera indica* L.) [43] in southern Thailand. This study compared 10 pairs of orchards that are located at <1 km and >7 km away from the forest edge. Stingless bees were the main visitors for 70.9% of total flower visits on rambutan (**Figure 11**). The distance from forest edge and location of natural stingless bee colony influenced the fruit set of rambutan significantly [43]. Stingless bees were significantly (two times) more frequent on rambutan flowers nearer to the forest.

In the case of durian and mango, even though stingless bees have been observed as flower visitors, the number of fruit sets was significantly influenced by bats (durian) and flies (mango).The distance to the forest did not affect the fruit yield of these two crops [43, 44]. The evidence suggests that the forest is an insect pollinator reservoir. The conservation of natural habitats surrounding a crop orchard is strongly recommended to maintain a population of forest-insect pollinators in natural habitats [45].

As generalists, stingless bees forage in vast array of plant taxa. A study of pollen foraging and resource partitioning of stingless bees throughout year-round

#### **Figure 11.**

*A linear regression plot for the number of rambutan fruit sets and insect visitation frequency to rambutan flowers in a mixed fruit orchard in Southern Thailand [43].*

#### **Figure 12.**

*Pollen resource partitioning of stingless bees: (a) pollen-type richness of samples between different colonies of stingless bees (C =* T. collina *(four colonies); MA =* T. melina*; MCA =* T. melanocephala*), (b) dissimilarity of pollen samples within the monospecific* collina*-aggregation over time, (c) dissimilarity of pollen samples within the mixed aggregation, and (d) flowering activity as a function of time in the habitat [46]. Dissimilarity is calculated as (1-Sørensen-index); Sørensen-index* is index *of similarity base on the equation of Sorensen [47].*

**185**

items on the international market are suggested.

**6. Conclusion**

*Southeast Asian Meliponiculture for Sustainable Livelihood*

showing their highly adaptive trait in pollen foraging.

adaptive species that are able to contribute to eco-services in SE Asia.

flowering dynamic in northern Borneo rainforest [46] unveils some interesting results. They compared pollen foraging within one monospecific (three colonies of *Trigona collina*) and one mixed nesting aggregation (one colony of *T. collina*, and one colony of each of the close relatives *T. melina* and *T. melanocephala*) in lowland tropical rain forest in Sabah, Malaysia. The results suggest that stingless bees, *Trigona collina*, show specificity of pollen source judged by the pollen similarity among the same species in the same aggregation sites. Within the two aggregations of *T. collina*, the similarity of pollen samples showed a strikingly different pattern over time, and there was no similarity at all between colonies in the mixed nest aggregation (**Figure 12**). Nevertheless, the resource partitioning occurs in their geographic location, with those other species of *T. melanocephala* and *T. melina*

Stingless bees also show interspecific differences in foraging behavior such as the speed of detecting new food sources. Observations at an artificial feeder revealed that *T. melanocephala* arrived at honey baits quicker than *T. melina*, whereas *T. collina* was reluctant to visit the feeding site [48]. Agriculture in SE Asia is frequently multicultivar system. Multiple plant species are cultivated together. For pollination of several flower phenotypes in a mixed plantation, stingless bees are certainly good choices. The diverse crops in those mixed orchards provide a high-quality foraging habitat for pollinators [40]. Much evidence showed that stingless bees are highly

Nowadays, meliponiculture (stingless beekeeping) in SE Asia is significantly increased. The promotion of stingless beekeeping as an additional activity for rural villages, together with high stingless bee species diversity [34], stimulates a revival of this activity. For instance, in Thailand, at least six species (*Tetragonula pagdeni*, *T. laeviceps*, *T. fuscobalteata*, *Lepidotrigona terminate*, *Heterotrigona itama*, and *Geniotrigona thoracica*) are commonly managed for commercial pollination services and honey production. Of these, *T. pagdeni* and *G. thoracica* can be the most easily transferred natural colonies to artificial wooden hive boxes. In addition, there are not only short time to colony recovery after dividing colony (4–8 weeks after dividing) but also show high yield of colony production—honey and pollen [29]. Stingless beekeepers increase colony number of these two stingless bee species for both colonies selling and producing honey in short time periods. Colony of *T. pagdeni* and *G. thoracica* can be sole in price of 800–1500 THB (\$25–\$47 USD) and 4000–5000 THB (\$125–\$157 USD) per colony, respectively. Therefore, *T. pagdeni* and *G. thoracica* are more suitable to promote and develop for meliponiculture in SE Asia. However, the comparisons of honey and propolis yields from common domestic species of stingless bee of SE Asia are highly suggested. The evaluation of status of a potential industry with the stingless bees with regard to honey production and yield, its commercialization, and management should be also taken into account. Unlike *A. mellifera*, meliponiculture in SE Asia is particularly suggested to the small-scale beekeeping with regard to the flora providing source from multi-cultivar systems that are commonly found in this region. The competitive situation of the prices of honey and other products between *Apis* and Meliponini is also a further issue to be determined. The species-based problems and solution that stingless beekeepers faced should be standardized precisely [49–51]. Finally, the management and production scenario should be developed to collectively improve a substantial quantity and quality of stingless bee products as significant competitive

*DOI: http://dx.doi.org/10.5772/intechopen.90344*

*Southeast Asian Meliponiculture for Sustainable Livelihood DOI: http://dx.doi.org/10.5772/intechopen.90344*

flowering dynamic in northern Borneo rainforest [46] unveils some interesting results. They compared pollen foraging within one monospecific (three colonies of *Trigona collina*) and one mixed nesting aggregation (one colony of *T. collina*, and one colony of each of the close relatives *T. melina* and *T. melanocephala*) in lowland tropical rain forest in Sabah, Malaysia. The results suggest that stingless bees, *Trigona collina*, show specificity of pollen source judged by the pollen similarity among the same species in the same aggregation sites. Within the two aggregations of *T. collina*, the similarity of pollen samples showed a strikingly different pattern over time, and there was no similarity at all between colonies in the mixed nest aggregation (**Figure 12**). Nevertheless, the resource partitioning occurs in their geographic location, with those other species of *T. melanocephala* and *T. melina* showing their highly adaptive trait in pollen foraging.

Stingless bees also show interspecific differences in foraging behavior such as the speed of detecting new food sources. Observations at an artificial feeder revealed that *T. melanocephala* arrived at honey baits quicker than *T. melina*, whereas *T. collina* was reluctant to visit the feeding site [48]. Agriculture in SE Asia is frequently multicultivar system. Multiple plant species are cultivated together. For pollination of several flower phenotypes in a mixed plantation, stingless bees are certainly good choices. The diverse crops in those mixed orchards provide a high-quality foraging habitat for pollinators [40]. Much evidence showed that stingless bees are highly adaptive species that are able to contribute to eco-services in SE Asia.
