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Options for stingless honey-beekeeping around Udzungwa Mountains National Park, Tanzania, and implications for biodiversity management

Options for stingless honey-beekeeping around Udzungwa Mountains National Park, Tanzania, and... International Journal of Biodiversity Science, Ecosystem Services & Management Vol. 6, Nos. 3–4, September–December 2010, 89–95 Options for stingless honey-beekeeping around Udzungwa Mountains National Park, Tanzania, and implications for biodiversity management a a b Marcelian A. Njau *, Fatina A. Mturi and Pauline M. Mpuya a b Department of Zoology and Wildlife Conservation, University of Dar es Salaam, Dar es Salaam, Tanzania; Ministry of Natural Resources and Tourism, Dar es Salaam, Tanzania Resident stingless honey-bees in the Udzungwa area were investigated for a period of 1 year. Specimens of sting- less honey-bees were captured on flowers and on natural nest entrances. Six local stingless honey-bee species, namely Dactylurina schmidti, Plebeina hildebrandti, Axestotrigona erythra, Meliponula ogouensis, M. lendliana and M. ferruginea, were found in the area, all of which are known to the local people. Honey productivity per colony in experimental hives varied according to species: M. ogouensis average 3.2 L, M. lendliana average 2.7 L, D. schmidti average 1.6 L and P. hilde- brandti average 0.6 L. These findings indicate good potential for beekeeping in the lower altitude area at the foot of the Udzungwa Mountains, where the stingless honey-bee species diversity is greatest. The article discusses some options for intensification of management of the bee resources while maintaining natural biodiversity in the area. Keywords: biodiversity; management; honey; stingless honey-bee species; Udzungwa Introduction queen, several drones and many workers, which are non- The stingless honey-bee has a long history of discovery reproductive females (Van Veen et al. 2004). The colony and description by naturalists and scholars during early size ranges from a few individuals to thousands of colony explorations of tropical regions of the world (Crane 1999), mates. In Africa, where a group of stingless honey-bees but they remain largely under-utilised by man (Kajobe and is considered to have originated (Roubik 1992), they are Roubik 2006). There are several general accounts of sting- found mainly in warmer tropical areas where temperature less honey-bees (Michener 2000; Byarugaba 2004; Eardley and humidity are conducive for their development through- 2004; Cortopassi-Laurino et al. 2006; Kajobe and Roubik out the year. It is estimated that in Africa, there are 32 2006). At present, it is estimated that the number of known stingless honey-bee species, and out of these 19 species in stingless honey-bee species in the world is 50 times more six genera (Eardley 2004) have been identified from speci- that of Apis species (Roubik 2006). With the exception of mens collected mainly from forested areas, of which more the neotropics, especially Latin America, where domesti- than half of the species have been collected from Tanzania. cation of stingless honey-bees is common, the rest of the Stingless honey-bees are not only important pollinators of tropics, particularly Africa, have not been able to effec- flowering plants but are also producers of honey, which tively harness the products of these bees because of a has some medicinal value (Byarugaba 2004). The bene- lack of basic knowledge of their biology and behaviour. fits accrued from using honey from stingless bees have Keeping of stingless honey-bees is known as meliponi- led communities to accord them recognition, especially in culture, and in tropical Africa it is found only in a very terms of aspects of their biology, such as nest construction few countries. In Tanzania, for instance, meliponiculture patterns. could be undertaken countrywide but at present it is lim- Nest construction characteristics, location and archi- ited to forested areas on Mount Kilimanjaro and Meru. tecture are unique to each species and can augment stin- Even where meliponiculture is practiced, as in Costa Rica gless honey-bee species identification (Wille 1983; Roubik (Sommeijer 1999), very few people in Tanzania have suf- 2006). Species of stingless honey-bee sometimes have ficient knowledge to practice it in such a way that the local names based on various aspects of their biology, benefits accrued from stingless beekeeping exceed the cost such as size, appearance of colony entrances and behaviour of investment in the enterprise. Although there is a great (Byarugaba 2004). Local names have shortcomings and are demand for honey produced by stingless honey-bees, not not reliable across the wider scientific community because many people are aware of how to keep these bees or even of language differences. The taxonomy of stingless honey- the many benefits such as crop pollination and conservation bee species, especially in Africa, is currently under revision of biodiversity that can be provided from good manage- and there are still several synonyms that have not been ment of the required bee resources. Stingless honey-bees fully resolved. At present, there are two major honey-bee are eusocial insects and form colonies consisting of a classification schemes (Moure 1971; Michener 2000), and *Corresponding author. Email: manjau@udsm.ac.tz ISSN 2151-3732 print/ISSN 2151-3740 online © 2010 Taylor & Francis DOI: 10.1080/21513732.2010.537699 http://www.infomaworld.com 90 M.A. Njau et al. there have been efforts to reclassify stingless honey-bees Survey of stingless honey-bees using both classical and molecular techniques (Eardley Every study site was subdivided into 100 subplots, each 2004; Rasmussen and Cameron 2007). It is now possi- measuring 1 ha. Sixty of the subplots per site were ran- ble to clearly identify the species of stingless honey-bee domly selected for intensive searches for natural nests. The and this will be useful for purposes of documentation and search was systematically conducted with the help of six domestication. skilled field assistants who also provided the vernacular Domestication of stingless bees for their honey is not names and behaviour of the bees found. The assistants were commonly practiced in Tanzania, even though their honey ex-primary school leavers who had completed 7 years of is highly valued. As in other tropical countries where there formal education and belonged to local beekeeping groups. are stingless bees, in Tanzania it has not been possible They had a thorough knowledge of the study area as well to keep them in ways that would be economically attrac- as of the local stingless bee fauna. Searching for stingless tive to the beekeepers. Most honey from stingless bees in bee nests was from tree to tree in each of the 1-ha subplots; Tanzania is obtained from natural nests, in which extrac- 4 ha were searched per day. The procedure was repeated tion procedures are destructive of both the nest and the in the wet and the dry seasons. Searching for stingless host tree. Domestication of stingless honey-bees in the honey-bees involved thorough inspection of tree trunks for form of meliponiculture as a strategy for conservation indicators of colonies such as nest entrance tubes. Nests of biodiversity would help minimise and eradicate this observed in the upper parts of trees were viewed through problem. The present study was undertaken with the objec- binoculars at a magnification of 8 × 40 mm. Any nest tive of documenting the species of stingless honey-bee in found, as well as the host tree species, was recorded. Also Udzungwa Mountain National Park and its surrounding recorded were the tree diameter at breast height (dbh) and area, and assessing their potential for domestication in the altitude in meters above sea level. Nests encountered in ter- form of meliponiculture. The presence of stingless bees mite mounds and walls of houses during searching were and the capacity to collect and store honey in the quanti- also recorded. ties observed in other places would stimulate development of beekeeping by the local community, an effort which, if realised, would form an important strategy for long-term Identification of stingless honey-bee species effective management of the biodiversity of the Udzungwa Stingless honey-bees from each study site were collected area and other similar ecosystems. from flowers using a sweep net, and also from natural nests, where they were collected at nest entrances using forceps. Materials and methods The specimens were preserved in 70% ethanol and then taken to the laboratory for identification. Specimens were Study area identified using the key for stingless honey-bee species The study was carried out in Udzungwa Mountains developed by Eardley (2004). National Park and neighbouring areas used by small-scale farmers growing cereals like maize and rice, as well as ◦  ◦ legumes. The park lies at 7 38 S, 36 41 E and occu- Characteristics of colonies, their behaviours and tree pies two regions (Iringa and Morogoro) of Tanzania. A hosts total area of 600 ha, representing the various types of Characteristics of the natural nests, as described by habitat in Udzungwa and comprising three zones delin- Michener (1974) and Wille (1979), were also used to help eated as core (within the park with restricted access to identify the species occurring in the area. The nests were resource utilisation), buffer (with limited resource util- partially opened and their internal structure examined. The isation) and transition (with unrestricted resource util- behaviour of the bees when their nests were approached isation) were selected for sampling and placement of was noted. experimental hives. The delineation of zones was simi- lar to that used for biosphere reserves (UNESCO MAB, Man and the Biosphere Seville strategy for biosphere Productivity of selected stingless honey-bee species reserves) and we used it because it was deemed appro- priate for this study. The sampling sites were located in Larger-bodied stingless honey-bees in the area belong- ◦   ◦   ◦ the core (7 47 31.5 S, 36 54 11.3 E), buffer (7 51 57.9 S, ing to the genera Meliponula, Dactylurina and Plebeina ◦   ◦   ◦ 36 55 7.1 E) and transition (7 45.2 7 S, 36 52 53.7 E) were studied for honey productivity of hives. Two resident zones. Each zone was divided into two study sites, each species of Meliponula and one each from the other genera measuring 100 ha, and 60 ha of each were used for in-depth were used. Twenty experimental nune boxes, designed and study. The sites were located on areas with contrasting alti- constructed based on those used by Nogueira-Neto (1970), tudes, one highland and the other lowland. The whole area per site were hoisted on randomly selected trees at 2 m at the foot of the mountain and below the 300 m a.s.l. con- from the ground for studying hive occupancy and honey tour was regarded as lowlands, while the area above this production. In total, there were 120 hives per each species. was regarded as the highland area. The methods used were Transferring of bee colonies from natural logs to hives was as detailed previously (Njau et al. 2009). undertaken 3 months after collection. Assessment of the International Journal of Biodiversity Science, Ecosystem Services & Management 91 hives was done monthly, and the occupancy determined p < 0.0001, n = 120). The diameter of the entrance tubes using the expression: were also significantly different, with that of M. ogouensis being 0.79 cm, M. lendliana 0.62 cm, P. hildebrandti 0.22 O cm and D. schmidti 0.92 cm. The direction of the entrances HO = × 100, was predominantly east (45%), but some also faced south (O + E) (24%), west (16%) and north (16%). Natural nests of stingless honey-bees were found where HO is hive occupancy, O is the number of occupied in host trees in the core, buffer and transitional zones hives and E is the number of empty hives. Average honey (Table 2). In all, 43,200 trees with a dbh ≥ 20 cm production per hive was determined base on five nune box were searched, and trees occupied by stingless honey- hives per species per hectare in the study area. Honey was bees had a dbh ranging from 46.66 to 450 cm; only P. harvested by tilting the boxes, removing the end disc, pierc- hildebrandti nested on a tree species more than once. ing the honey pots and collecting the flow into a vessel. The More than one nest of P. hildebrandti was observed on volume of the honey was determined using a measuring K. aethiopum. The dbh is important in terms of hosting, cylinder. and the mean dbh of trees hosting natural nests of differ- ent stingless honey-bee species was significantly different (KW = 70.315, p < 0.0001, n = 135), with bigger trees Results proportionally contributing to a higher number amongst Stingless honey-bee species of Udzungwa the hosts (χ = 29.08, p < 0.0001, n = 135). Each stingless There were six distinct species in the Udzungwa honey-bee species had an observed minimum tree diame- area (Table 1). These species were known to the lo- ter requirement for nesting, but no maximum tree diameter. cal communities as Bombwi (Plebeina hildebrandti), The lower dbh limit required for M. ogouensis was 145 Mlenga (Meliponula ogouensis), Vitaparuku (Dactylurina cm, for M. lendliana was 73.77 cm, for P. hildebrandti schmidti), Nganga (Axestotrigona erythra), Limbofu (M. it was 48.66 cm and for D. schmidti it was 155.67 cm. lendliana) and Upula (M. feruginnea). All six species Highly productive stingless honey-bee species used bigger occurred in the lowlands at altitudes between 190 m and trees, with mean dbh of 202.33 cm for M. lendliana and 458 m a.s.l. Two species also occurred at mid-altitudes 205.67 cm for D. schmidti. The dbh of host trees used by M. of 458–753 m a.s.l. and only one species extended its range lendliana and P. hildebrandti was not significantly different to 1500 m a.s.l. from each other. Host minimum tree size dbh was 145 cm for M. ogouensis, 73.66 cm for M. lendliana, 48.66 cm for P. hildebrandti and 77.65 cm for D. schmidti. Trees of these Characteristics of the bee colonies, behaviours and tree sizes were almost depleted in the transitional (farmlands) hosts and buffer zones of Udzungwa, and thus stingless bee- The stingless honey-bee species had distinct nests with fea- keeping in these zones required hoisting hives into trees. tures that could be used to characterise them. The nests The number of colonies of stingless honey-bee in the study –1 –1 of P. hildebrandti had clusters of horizontal cells; nests area ranged from 0.20 ha to 0.43 ha for M. ogouen- –1 –1 of M. ogouensis had brood cells that were cylindrical and sis, 0.20–0.25 ha for M. lendliana, 0.22–0.60 ha for P. –1 layered irregularly; nests of D. schmidti were constructed hildebrandti and 0.5–0.25 ha for D. schmidti. There was high nest density in highland areas relative to lowland areas on tree branches at a great height above the ground; nests during both dry and wet seasons and such numbers indicate of M. lendliana also had cylindrical but separate brood that the area is suitable for meliponiculture. cells; and two species (A. erythra and M. ferruginea)were The density (nests per hectare) of natural stingless only caught while foraging, but their nests were not located honey-bee nests is shown in Tables 2 and 3. It was found in the field. The mean length of the entrance tube of that the number of nests in the dry season was higher in M. ogouensis was 5.33 cm, while that of M. lendliana was the highland sites than in the lowland sites for most of 4.76 cm, that of P. hildebrandti was 4.32 cm and that of the species. D. schmidti had a higher number of nests in D. schmidti was 5.46 cm. The forms of the openings var- the highlands than in the lowlands during the wet season. ied significantly among the various species (KW = 32.161, Table 1. Species of stingless honey-bees of Udzungwa. Scientific name Vernacular name Altitude range (meters) Meliponula ogouensis Mlenga 190–753 M. lendliana Limbofu 190–1500 Plebeina hildebrandti Bombwi 190–458 Dactylurina schmidti Vitaparuku 190–458 M. ferruginea Upula 190–458 Axestotrigona erythra Nganga 190–458 92 M.A. Njau et al. Table 2. Host plants of stingless honey-bees occurrence and mean dbh in Udzungwa. No. in No. in Honey-bee Mean dbh No. in buffer transition species Host plant (cm) core zone zone zone Meliponula Eckebergia capensis 202.33 0 0 3 ogouensis Pseudolachnodtylis 175.09 3 8 5 maproneifolia Combretum molle 145.0 17 12 4 Termite mound 8 M. lendliana Eckebergia capensis 196.54 0 6 4 Pseudolachnodtylis 94.66 8 8 3 maproneifolia Combretum molle 82.45 9 4 1 Kigelia aethiopum 73.66 3 2 5 Termite mound 0 Dactylurina Delonix regia 168.67 0 3 6 schmidti Kigelia aethiopum 77.65 6 8 10 Trichlia emertica 205.75 3 6 0 Bombax 205.67 0 0 4 rhodognaphalon Funtumia latifolia 155.67 7 3 0 Sterculia appendiculata 171.68 4 0 0 Plebeina Kigelia aethiopum 54.38 13 15 12 hildebrandti Dalbergia melanoxylon 48.66 0 0 0 Vitex doniana 58.33 7 5 7 Table 3. Number of bee colonies per hectare. Wet season Dry season Honey-bee species Highland Lowland Highland Lowland Meliponula ogouensis 0.62 0.60 0.55 0.50 M. lendliana 0.76 0.71 0.69 0.63 Dactylurina schmidti 0.24 0.30 0.28 0.21 Plebeina hidebrandti 0.86 0.78 0.77 0.71 Also, this species had a higher number of nests in low- to nune hives and did not swarm back. Occupancy for land than in highland areas during the dry season. The P. hildebrandti decreased at first but then estabilised. number of nests per hectare of M. ogouensis, M. lendliana The amount of honey produced by various species of and P. hildebrandti was higher in highland than in lowland stingless honey-bee in the Udzungwa area varied with areas, both during the wet and the dry seasons (Table 3). respect to species (Table 4). The amount of honey produced by M. ogouensis was higher than that of M. lendliana, and D. schmidti produced less honey than M. lendliana Productivity of selected stingless honey-bee species but more than P. hildebrandti. Honey production among Hive occupancy gradually declined, reaching the low- the four species in the Udzungwa area was differed sig- est values between February and April for M. ogouensis nificantly between species (Fr = 314.70; p < 0.0001, and M. lendliana; occupancy for D. schmidti reached n = 120). However, honey produced in the lowlands by zero between August and October; and occupancy for all species showed no significant difference among sites M. lendliana and M. ogouensis increased again between (Fr = 157; p > 0.05, n = 240). When production at differ- May and October. The case was different for P. hildebrandti ent zones was analysed, there was no significant difference as their colonies declined steadily throughout the year. among the species. The number of colonies of M. ogouensis and M. lendliana increased steadily between May and October, indicating Discussion and conclusions new swarms in the hives during the period. Occupancies Distribution of stingless honey-bee species of Udzungwa were not significantly different among the sites (KW = 4.331, p = 0.5028, n = 480), and both M ogouensis The local people of Udzungwa recognised six different and M. lendliana had the same occupancy. Colonies of stingless honey-bee species. It is not surprising that the D. schmidti absconded within the first year of their transfer local people have quite substantial indigenous knowledge International Journal of Biodiversity Science, Ecosystem Services & Management 93 Table 4. Honey production (litres per hive) by stingless honey-bees of Udzungwa. Species Volume in L (mean ± SD) n = 120 Meliponula ogouensis 3.2 ± 0.72 M. lendliana 2.7 ± 0.04 Dactylurina schmidti 1.9 ± 0.49 Plebeina hildebrandti 0.6 ± 0.19 about stingless bees nesting in their local forests, as also survival. A very small proportions of trees was used for found in other studies (Byarugaba 2004; dos Santos and nesting, only 13 out of 600 local tree species hosted sting- Antonini 2008) because they collect honey from the bees. less honey-bee colonies. This is possibly because stingless At present, stingless honey-bee species collected from dif- honey-bees tend to use larger trees for nesting, and there ferent places in Tanzania (Figure 1), include Cleptotrigona are very few such trees. In Udzungwa, stingless honey- cubiceps, D. schmidti, P. hildebrandti, M. ferruginea, bees use live trees with cavities, which are naturally less M. ogouensis, M. lendliana, Hypotrigona gribodoi, H. rus- susceptible to destruction by decomposers such as termites. poli, H. araujoi and Liotrigona bottegoi (Eardley 2004). Considering the fact that the primary functions of a nest are The number of colonies per hectare for each of the four for protection, brood rearing and food storage (Michener more productive species in the Udzungwa area in dif- 1974), it is of significance that stingless honey-bees would ferent seasons differed and the difference observed in find it beneficial to utilise the suitable cavities that provide the first and second half of the year could indicate a the required environment. process of swarming. It would be of great interest to undertake studies on swarming in stingless honey-bees, Productivity of selected stingless honey-bee species particularly in relation to colonisation of suitable cavities. Hive occupancy of stingless honey-bees varied with sea- There are forests patches in Udzungwa under local com- son, and this could be related to variation in humid- munity management where apiaries and meliponiaries have ity, where high air moisture content caused the bees to been established by local beekeeping groups. These forests abscond. For example, there was low occupancy during should be used for education and creation of awareness the wet season from February to April. Hive occupancy of on the need to manage the resources in these areas more D. schmidti decreased to zero between August and October sustainably. during the hot dry season, presumably due to high tem- peratures. High occupancy registered between May and Characteristics of colonies, behaviours and tree hosts October for M. ogouensis and M. lendliana was proba- Stingless honey-bee colony characteristics and bee bly due to swarming, resulting from colony disturbances behaviours, such as location of nesting, are useful in by honey hunters. In general, the number of experimen- species characterisation and may be useful in identifica- tal hives occupied by colonies of stingless honey-bee tion; for example, construction of colonies on tree branches stabilised for a period of only 3 months and decreased pro- at great heights above the ground for D. schmidti.The gressively thereafter due to poor insulation. Large-bodied behaviour of nest-building on tree branches has also been species of stingless honey-bee such as the M. ogouensis observed in Trigona dallatorea in Brazil (Michener 1974). and M. lendliana produced more honey relative to small- M. ogouensis nested in cavities close to the ground or even bodied species such as P. hildebrandti. Such productivity in vacated or live nests of termites and ants. M. lendliana could be attributed to the capacity of such species to collect nested in tree cavities located slightly higher but not on the and store honey. There was higher honey production in the ground. Nesting in subterranean areas observed in transi- buffer zone than in either the core or transition zone due to tional areas in Udzungwa was related to the absence of a higher number of colonies and higher occupancy. Honey suitable nesting tree cavities, due to severe deforestation pots were found in clusters segregated from brood cham- by the local communities. bers and pollen. The quantity of honey produced per colony The variations in nesting locations of the various of stingless honey-bee in the study area was high enough to species constitute a strategy to avoid interspecific com- make meliponiculture using nune boxes viable. In northern petition, as well as a strategy for protection, as was the Tanzania, especially in Kilimanjaro, log hives are used for case with D. schmidti, which constructed nests on tree stingless honey-beekeeping, but there is a need to design branches at great heights. Occasionally two or more nests modern box hives for beekeeping of stingless bees in the of the same species shared a cavity, raising the question Udzungwa area. of tolerance among members of different colonies. It is The amount of honey stored in the colonies varied enor- also not clear what were the reasons for nest stratification, mously among species, with some such as M. ogouensis but for stingless honey-bees, defence and diversification and M. lendliana storing as much as 3 L per colony per of resource utilisation could be important strategies for year. Other species like D. schmidti and P. hildebrandti 94 M.A. Njau et al. Figure 1. Map of Tanzania showing distribution of stingless honey-bee species. stored 1.6 L and 0.6 L, respectively. Mean productivity into the Udzungwa area for human needs and as a viable of honey per hive per colony in a season is usually low, additional income-generating activity. This will inevitably for example, Melipona beecheii in Costa Rica produces increase awareness of the importance of conservation of on average only about 2.63 L (Cortopassi-Laurino et al. the local forests and the Udzungwa ecosystem for local 2005). When compared to honey production by the stinging human communities around the game park. Interestingly, it honey-bees, which is usually over 20 L, the small amount was found that stingless honey-bee diversity declined with of honey stored by stingless bees may not be very attractive altitude. Whereas all species could be found in the low- to potential beekeepers. Only some stingless honey-bee altitude area at the foot of the mountains, only one species species store honey in quantities that could be harvested was found on the slopes at high altitude (1500 m a.s.l.). No by humans. However, in the Udzungwa area, local people wonder the occurrence of stingless honey-bees is confined do not usually keep stingless honey-bees but they hunt for to the warmer tropical regions of the world. In Africa, stin- honey, which is used for purposes of worship and medicine. gless honey-bees occur mostly within the tropics (Eardley It would be of much benefit to introduce meliponiculture 2004). International Journal of Biodiversity Science, Ecosystem Services & Management 95 Hunting for honey from stingless bees is currently done Crane E. 1999. The world history of beekeeping and honey hunting. London: Taylor & Francis. through cutting down of host trees. This practice affects the dos Santos GM, Antonini Y. 2008. The traditional knowledge on biodiversity of the area in many ways, such as a vast loss stingless bees (Apidae: Meliponina) used by the Enawene- of genes of the fauna and flora, bees and their associated Nawe tribe in western Brazil. J Ethnobiol Ethnomed. species, and destruction of the bee’s habitat through setting 4:19. fire to farm fields as a method of clearing them for cultiva- Eardley CD. 2004. Taxonomic revision of the African stingless bees (Apoidea: Apidae: Apinae: Meliponini). African Plant tion. Deforestation and use of fire lead to soil erosion and Protection. 10:63–96. other changes to the habitat. Local communities occupy the Kajobe R, Roubik DW. 2006. Honey-making bee colony abun- lower altitude area. In this area, there are presently small dance and predation by apes and humans in a Uganda forest community forests that are managed by these communi- reserve. Biotropica. 38:210–218. ties. Such forests provide additional habitats and resources Michener CD. 1974. The social behavior of the bees: a compar- ative study. Cambridge (MA): Harvard University Press. p. for the bee fauna. The owners of these forests should be 329–346. provided with more awareness of the importance of their Michener CD. 2000. The bees of the world. Baltimore (MD): John forests and be encouraged to plant tree species with flowers Hopkins University Press. that are rich in nectar and pollen for the bees. Moure JS. 1971. Descriçäo de uma espécia de Tetragona do Brasil Ownership of community forests offers the opportu- Central (Hymenoptera–Apidae). Boletim da Universidade Federal do Paraná. Zoologia. 4:47–50. nity to acquire knowledge of the biodiversity of the area. Njau MA, Mpuya P, Mturi FA. 2009. Apiculture potential in pro- There should be a systematic method for studying the tected areas: the case of Udzungwa Mountains National Park, species composition of these forests and to find out ways Tanzania. Int J Biodivers Sci Manag. 5(2):95–101. of increasing such bee species. Information should also Nogueira-Neto P. 1970. Behavioral problems related to the pil- be collected on the various threats to the survival of the lages made by some parasitic stingless bees (Meliponinae: Apidae). In: Aronson LR, Tobach E, Lehrman DS, Rosenblatt biodiversity and management procedures should be insti- JS, editors. Development and evolution of behavior. Essays tuted for wise use of the resources in a manner that will in memory of T.C. Schnerlan. San Francisco (CA): W.H. be sustainable and beneficial to all stakeholders, and espe- Freeman & Co. p. 416–434. cially the local community itself. We therefore recommend Rasmussen C, Cameron SA. 2007. A molecular phylogeny of the the development of eco-villages in these and similar areas Old World stingless bees (Hymenoptera: Apidae: Meliponini) and the non-monophyly of the large genus Trigona.Syst in order to achieve good management of the resident Entomol. 32:26–39. biodiversity. Roubik DW. 1992. Stingless bees: a guide to the Panamanian and Mesoamerican species and their nests (Hymenoptera: Apidae; Meliponinae) In: Quintero D, Aiello A, editors. Acknowledgements Insects of Panama and Mesoamerica Vol. 2. Oxford (UK): Oxford University Press. p. 495–524. We wish to thank the management of Udzungwa Mountains Roubik DW. 2006. Stingless bee nesting biology. Apidologie. National Park and the people living around the park, espe- 37:124–143. cially the beekeeping groups for their assistance and cooperation Sommeijer MJ. 1999. A regional program for training and throughout the study. We also thank WWF for funding the study. research on tropical beekeeping and tropical bees in Costa Rica. Bee World. 80(2):70–79. Van Veen JW, Arce Arce HG, Sommeijer MJ. 2004. Production References of queens and drones in Melipona beecheii (Meliponini) in Byarugaba D. 2004. Stingless bees (Hymenoptera: Apidae) of relation to colony development and resource availability. Proc Bwindi impenetrable forest Uganda and Abayanda indige- Netherlands Entomol Soc. 15:035–039. nous knowledge. Int J Trop Insect Sci. 24(1):117–121. Wille A. 1979. Phylogeny and relationship among the genera and Cortopassi-Laurino M, Imperatriz-Fonseca VL, Roubik DW, subgenera of the stingless bees (Meliponinae) of the world. 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Options for stingless honey-beekeeping around Udzungwa Mountains National Park, Tanzania, and implications for biodiversity management

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International Journal of Biodiversity Science, Ecosystem Services & Management Vol. 6, Nos. 3–4, September–December 2010, 89–95 Options for stingless honey-beekeeping around Udzungwa Mountains National Park, Tanzania, and implications for biodiversity management a a b Marcelian A. Njau *, Fatina A. Mturi and Pauline M. Mpuya a b Department of Zoology and Wildlife Conservation, University of Dar es Salaam, Dar es Salaam, Tanzania; Ministry of Natural Resources and Tourism, Dar es Salaam, Tanzania Resident stingless honey-bees in the Udzungwa area were investigated for a period of 1 year. Specimens of sting- less honey-bees were captured on flowers and on natural nest entrances. Six local stingless honey-bee species, namely Dactylurina schmidti, Plebeina hildebrandti, Axestotrigona erythra, Meliponula ogouensis, M. lendliana and M. ferruginea, were found in the area, all of which are known to the local people. Honey productivity per colony in experimental hives varied according to species: M. ogouensis average 3.2 L, M. lendliana average 2.7 L, D. schmidti average 1.6 L and P. hilde- brandti average 0.6 L. These findings indicate good potential for beekeeping in the lower altitude area at the foot of the Udzungwa Mountains, where the stingless honey-bee species diversity is greatest. The article discusses some options for intensification of management of the bee resources while maintaining natural biodiversity in the area. Keywords: biodiversity; management; honey; stingless honey-bee species; Udzungwa Introduction queen, several drones and many workers, which are non- The stingless honey-bee has a long history of discovery reproductive females (Van Veen et al. 2004). The colony and description by naturalists and scholars during early size ranges from a few individuals to thousands of colony explorations of tropical regions of the world (Crane 1999), mates. In Africa, where a group of stingless honey-bees but they remain largely under-utilised by man (Kajobe and is considered to have originated (Roubik 1992), they are Roubik 2006). There are several general accounts of sting- found mainly in warmer tropical areas where temperature less honey-bees (Michener 2000; Byarugaba 2004; Eardley and humidity are conducive for their development through- 2004; Cortopassi-Laurino et al. 2006; Kajobe and Roubik out the year. It is estimated that in Africa, there are 32 2006). At present, it is estimated that the number of known stingless honey-bee species, and out of these 19 species in stingless honey-bee species in the world is 50 times more six genera (Eardley 2004) have been identified from speci- that of Apis species (Roubik 2006). With the exception of mens collected mainly from forested areas, of which more the neotropics, especially Latin America, where domesti- than half of the species have been collected from Tanzania. cation of stingless honey-bees is common, the rest of the Stingless honey-bees are not only important pollinators of tropics, particularly Africa, have not been able to effec- flowering plants but are also producers of honey, which tively harness the products of these bees because of a has some medicinal value (Byarugaba 2004). The bene- lack of basic knowledge of their biology and behaviour. fits accrued from using honey from stingless bees have Keeping of stingless honey-bees is known as meliponi- led communities to accord them recognition, especially in culture, and in tropical Africa it is found only in a very terms of aspects of their biology, such as nest construction few countries. In Tanzania, for instance, meliponiculture patterns. could be undertaken countrywide but at present it is lim- Nest construction characteristics, location and archi- ited to forested areas on Mount Kilimanjaro and Meru. tecture are unique to each species and can augment stin- Even where meliponiculture is practiced, as in Costa Rica gless honey-bee species identification (Wille 1983; Roubik (Sommeijer 1999), very few people in Tanzania have suf- 2006). Species of stingless honey-bee sometimes have ficient knowledge to practice it in such a way that the local names based on various aspects of their biology, benefits accrued from stingless beekeeping exceed the cost such as size, appearance of colony entrances and behaviour of investment in the enterprise. Although there is a great (Byarugaba 2004). Local names have shortcomings and are demand for honey produced by stingless honey-bees, not not reliable across the wider scientific community because many people are aware of how to keep these bees or even of language differences. The taxonomy of stingless honey- the many benefits such as crop pollination and conservation bee species, especially in Africa, is currently under revision of biodiversity that can be provided from good manage- and there are still several synonyms that have not been ment of the required bee resources. Stingless honey-bees fully resolved. At present, there are two major honey-bee are eusocial insects and form colonies consisting of a classification schemes (Moure 1971; Michener 2000), and *Corresponding author. Email: manjau@udsm.ac.tz ISSN 2151-3732 print/ISSN 2151-3740 online © 2010 Taylor & Francis DOI: 10.1080/21513732.2010.537699 http://www.infomaworld.com 90 M.A. Njau et al. there have been efforts to reclassify stingless honey-bees Survey of stingless honey-bees using both classical and molecular techniques (Eardley Every study site was subdivided into 100 subplots, each 2004; Rasmussen and Cameron 2007). It is now possi- measuring 1 ha. Sixty of the subplots per site were ran- ble to clearly identify the species of stingless honey-bee domly selected for intensive searches for natural nests. The and this will be useful for purposes of documentation and search was systematically conducted with the help of six domestication. skilled field assistants who also provided the vernacular Domestication of stingless bees for their honey is not names and behaviour of the bees found. The assistants were commonly practiced in Tanzania, even though their honey ex-primary school leavers who had completed 7 years of is highly valued. As in other tropical countries where there formal education and belonged to local beekeeping groups. are stingless bees, in Tanzania it has not been possible They had a thorough knowledge of the study area as well to keep them in ways that would be economically attrac- as of the local stingless bee fauna. Searching for stingless tive to the beekeepers. Most honey from stingless bees in bee nests was from tree to tree in each of the 1-ha subplots; Tanzania is obtained from natural nests, in which extrac- 4 ha were searched per day. The procedure was repeated tion procedures are destructive of both the nest and the in the wet and the dry seasons. Searching for stingless host tree. Domestication of stingless honey-bees in the honey-bees involved thorough inspection of tree trunks for form of meliponiculture as a strategy for conservation indicators of colonies such as nest entrance tubes. Nests of biodiversity would help minimise and eradicate this observed in the upper parts of trees were viewed through problem. The present study was undertaken with the objec- binoculars at a magnification of 8 × 40 mm. Any nest tive of documenting the species of stingless honey-bee in found, as well as the host tree species, was recorded. Also Udzungwa Mountain National Park and its surrounding recorded were the tree diameter at breast height (dbh) and area, and assessing their potential for domestication in the altitude in meters above sea level. Nests encountered in ter- form of meliponiculture. The presence of stingless bees mite mounds and walls of houses during searching were and the capacity to collect and store honey in the quanti- also recorded. ties observed in other places would stimulate development of beekeeping by the local community, an effort which, if realised, would form an important strategy for long-term Identification of stingless honey-bee species effective management of the biodiversity of the Udzungwa Stingless honey-bees from each study site were collected area and other similar ecosystems. from flowers using a sweep net, and also from natural nests, where they were collected at nest entrances using forceps. Materials and methods The specimens were preserved in 70% ethanol and then taken to the laboratory for identification. Specimens were Study area identified using the key for stingless honey-bee species The study was carried out in Udzungwa Mountains developed by Eardley (2004). National Park and neighbouring areas used by small-scale farmers growing cereals like maize and rice, as well as ◦  ◦ legumes. The park lies at 7 38 S, 36 41 E and occu- Characteristics of colonies, their behaviours and tree pies two regions (Iringa and Morogoro) of Tanzania. A hosts total area of 600 ha, representing the various types of Characteristics of the natural nests, as described by habitat in Udzungwa and comprising three zones delin- Michener (1974) and Wille (1979), were also used to help eated as core (within the park with restricted access to identify the species occurring in the area. The nests were resource utilisation), buffer (with limited resource util- partially opened and their internal structure examined. The isation) and transition (with unrestricted resource util- behaviour of the bees when their nests were approached isation) were selected for sampling and placement of was noted. experimental hives. The delineation of zones was simi- lar to that used for biosphere reserves (UNESCO MAB, Man and the Biosphere Seville strategy for biosphere Productivity of selected stingless honey-bee species reserves) and we used it because it was deemed appro- priate for this study. The sampling sites were located in Larger-bodied stingless honey-bees in the area belong- ◦   ◦   ◦ the core (7 47 31.5 S, 36 54 11.3 E), buffer (7 51 57.9 S, ing to the genera Meliponula, Dactylurina and Plebeina ◦   ◦   ◦ 36 55 7.1 E) and transition (7 45.2 7 S, 36 52 53.7 E) were studied for honey productivity of hives. Two resident zones. Each zone was divided into two study sites, each species of Meliponula and one each from the other genera measuring 100 ha, and 60 ha of each were used for in-depth were used. Twenty experimental nune boxes, designed and study. The sites were located on areas with contrasting alti- constructed based on those used by Nogueira-Neto (1970), tudes, one highland and the other lowland. The whole area per site were hoisted on randomly selected trees at 2 m at the foot of the mountain and below the 300 m a.s.l. con- from the ground for studying hive occupancy and honey tour was regarded as lowlands, while the area above this production. In total, there were 120 hives per each species. was regarded as the highland area. The methods used were Transferring of bee colonies from natural logs to hives was as detailed previously (Njau et al. 2009). undertaken 3 months after collection. Assessment of the International Journal of Biodiversity Science, Ecosystem Services & Management 91 hives was done monthly, and the occupancy determined p < 0.0001, n = 120). The diameter of the entrance tubes using the expression: were also significantly different, with that of M. ogouensis being 0.79 cm, M. lendliana 0.62 cm, P. hildebrandti 0.22 O cm and D. schmidti 0.92 cm. The direction of the entrances HO = × 100, was predominantly east (45%), but some also faced south (O + E) (24%), west (16%) and north (16%). Natural nests of stingless honey-bees were found where HO is hive occupancy, O is the number of occupied in host trees in the core, buffer and transitional zones hives and E is the number of empty hives. Average honey (Table 2). In all, 43,200 trees with a dbh ≥ 20 cm production per hive was determined base on five nune box were searched, and trees occupied by stingless honey- hives per species per hectare in the study area. Honey was bees had a dbh ranging from 46.66 to 450 cm; only P. harvested by tilting the boxes, removing the end disc, pierc- hildebrandti nested on a tree species more than once. ing the honey pots and collecting the flow into a vessel. The More than one nest of P. hildebrandti was observed on volume of the honey was determined using a measuring K. aethiopum. The dbh is important in terms of hosting, cylinder. and the mean dbh of trees hosting natural nests of differ- ent stingless honey-bee species was significantly different (KW = 70.315, p < 0.0001, n = 135), with bigger trees Results proportionally contributing to a higher number amongst Stingless honey-bee species of Udzungwa the hosts (χ = 29.08, p < 0.0001, n = 135). Each stingless There were six distinct species in the Udzungwa honey-bee species had an observed minimum tree diame- area (Table 1). These species were known to the lo- ter requirement for nesting, but no maximum tree diameter. cal communities as Bombwi (Plebeina hildebrandti), The lower dbh limit required for M. ogouensis was 145 Mlenga (Meliponula ogouensis), Vitaparuku (Dactylurina cm, for M. lendliana was 73.77 cm, for P. hildebrandti schmidti), Nganga (Axestotrigona erythra), Limbofu (M. it was 48.66 cm and for D. schmidti it was 155.67 cm. lendliana) and Upula (M. feruginnea). All six species Highly productive stingless honey-bee species used bigger occurred in the lowlands at altitudes between 190 m and trees, with mean dbh of 202.33 cm for M. lendliana and 458 m a.s.l. Two species also occurred at mid-altitudes 205.67 cm for D. schmidti. The dbh of host trees used by M. of 458–753 m a.s.l. and only one species extended its range lendliana and P. hildebrandti was not significantly different to 1500 m a.s.l. from each other. Host minimum tree size dbh was 145 cm for M. ogouensis, 73.66 cm for M. lendliana, 48.66 cm for P. hildebrandti and 77.65 cm for D. schmidti. Trees of these Characteristics of the bee colonies, behaviours and tree sizes were almost depleted in the transitional (farmlands) hosts and buffer zones of Udzungwa, and thus stingless bee- The stingless honey-bee species had distinct nests with fea- keeping in these zones required hoisting hives into trees. tures that could be used to characterise them. The nests The number of colonies of stingless honey-bee in the study –1 –1 of P. hildebrandti had clusters of horizontal cells; nests area ranged from 0.20 ha to 0.43 ha for M. ogouen- –1 –1 of M. ogouensis had brood cells that were cylindrical and sis, 0.20–0.25 ha for M. lendliana, 0.22–0.60 ha for P. –1 layered irregularly; nests of D. schmidti were constructed hildebrandti and 0.5–0.25 ha for D. schmidti. There was high nest density in highland areas relative to lowland areas on tree branches at a great height above the ground; nests during both dry and wet seasons and such numbers indicate of M. lendliana also had cylindrical but separate brood that the area is suitable for meliponiculture. cells; and two species (A. erythra and M. ferruginea)were The density (nests per hectare) of natural stingless only caught while foraging, but their nests were not located honey-bee nests is shown in Tables 2 and 3. It was found in the field. The mean length of the entrance tube of that the number of nests in the dry season was higher in M. ogouensis was 5.33 cm, while that of M. lendliana was the highland sites than in the lowland sites for most of 4.76 cm, that of P. hildebrandti was 4.32 cm and that of the species. D. schmidti had a higher number of nests in D. schmidti was 5.46 cm. The forms of the openings var- the highlands than in the lowlands during the wet season. ied significantly among the various species (KW = 32.161, Table 1. Species of stingless honey-bees of Udzungwa. Scientific name Vernacular name Altitude range (meters) Meliponula ogouensis Mlenga 190–753 M. lendliana Limbofu 190–1500 Plebeina hildebrandti Bombwi 190–458 Dactylurina schmidti Vitaparuku 190–458 M. ferruginea Upula 190–458 Axestotrigona erythra Nganga 190–458 92 M.A. Njau et al. Table 2. Host plants of stingless honey-bees occurrence and mean dbh in Udzungwa. No. in No. in Honey-bee Mean dbh No. in buffer transition species Host plant (cm) core zone zone zone Meliponula Eckebergia capensis 202.33 0 0 3 ogouensis Pseudolachnodtylis 175.09 3 8 5 maproneifolia Combretum molle 145.0 17 12 4 Termite mound 8 M. lendliana Eckebergia capensis 196.54 0 6 4 Pseudolachnodtylis 94.66 8 8 3 maproneifolia Combretum molle 82.45 9 4 1 Kigelia aethiopum 73.66 3 2 5 Termite mound 0 Dactylurina Delonix regia 168.67 0 3 6 schmidti Kigelia aethiopum 77.65 6 8 10 Trichlia emertica 205.75 3 6 0 Bombax 205.67 0 0 4 rhodognaphalon Funtumia latifolia 155.67 7 3 0 Sterculia appendiculata 171.68 4 0 0 Plebeina Kigelia aethiopum 54.38 13 15 12 hildebrandti Dalbergia melanoxylon 48.66 0 0 0 Vitex doniana 58.33 7 5 7 Table 3. Number of bee colonies per hectare. Wet season Dry season Honey-bee species Highland Lowland Highland Lowland Meliponula ogouensis 0.62 0.60 0.55 0.50 M. lendliana 0.76 0.71 0.69 0.63 Dactylurina schmidti 0.24 0.30 0.28 0.21 Plebeina hidebrandti 0.86 0.78 0.77 0.71 Also, this species had a higher number of nests in low- to nune hives and did not swarm back. Occupancy for land than in highland areas during the dry season. The P. hildebrandti decreased at first but then estabilised. number of nests per hectare of M. ogouensis, M. lendliana The amount of honey produced by various species of and P. hildebrandti was higher in highland than in lowland stingless honey-bee in the Udzungwa area varied with areas, both during the wet and the dry seasons (Table 3). respect to species (Table 4). The amount of honey produced by M. ogouensis was higher than that of M. lendliana, and D. schmidti produced less honey than M. lendliana Productivity of selected stingless honey-bee species but more than P. hildebrandti. Honey production among Hive occupancy gradually declined, reaching the low- the four species in the Udzungwa area was differed sig- est values between February and April for M. ogouensis nificantly between species (Fr = 314.70; p < 0.0001, and M. lendliana; occupancy for D. schmidti reached n = 120). However, honey produced in the lowlands by zero between August and October; and occupancy for all species showed no significant difference among sites M. lendliana and M. ogouensis increased again between (Fr = 157; p > 0.05, n = 240). When production at differ- May and October. The case was different for P. hildebrandti ent zones was analysed, there was no significant difference as their colonies declined steadily throughout the year. among the species. The number of colonies of M. ogouensis and M. lendliana increased steadily between May and October, indicating Discussion and conclusions new swarms in the hives during the period. Occupancies Distribution of stingless honey-bee species of Udzungwa were not significantly different among the sites (KW = 4.331, p = 0.5028, n = 480), and both M ogouensis The local people of Udzungwa recognised six different and M. lendliana had the same occupancy. Colonies of stingless honey-bee species. It is not surprising that the D. schmidti absconded within the first year of their transfer local people have quite substantial indigenous knowledge International Journal of Biodiversity Science, Ecosystem Services & Management 93 Table 4. Honey production (litres per hive) by stingless honey-bees of Udzungwa. Species Volume in L (mean ± SD) n = 120 Meliponula ogouensis 3.2 ± 0.72 M. lendliana 2.7 ± 0.04 Dactylurina schmidti 1.9 ± 0.49 Plebeina hildebrandti 0.6 ± 0.19 about stingless bees nesting in their local forests, as also survival. A very small proportions of trees was used for found in other studies (Byarugaba 2004; dos Santos and nesting, only 13 out of 600 local tree species hosted sting- Antonini 2008) because they collect honey from the bees. less honey-bee colonies. This is possibly because stingless At present, stingless honey-bee species collected from dif- honey-bees tend to use larger trees for nesting, and there ferent places in Tanzania (Figure 1), include Cleptotrigona are very few such trees. In Udzungwa, stingless honey- cubiceps, D. schmidti, P. hildebrandti, M. ferruginea, bees use live trees with cavities, which are naturally less M. ogouensis, M. lendliana, Hypotrigona gribodoi, H. rus- susceptible to destruction by decomposers such as termites. poli, H. araujoi and Liotrigona bottegoi (Eardley 2004). Considering the fact that the primary functions of a nest are The number of colonies per hectare for each of the four for protection, brood rearing and food storage (Michener more productive species in the Udzungwa area in dif- 1974), it is of significance that stingless honey-bees would ferent seasons differed and the difference observed in find it beneficial to utilise the suitable cavities that provide the first and second half of the year could indicate a the required environment. process of swarming. It would be of great interest to undertake studies on swarming in stingless honey-bees, Productivity of selected stingless honey-bee species particularly in relation to colonisation of suitable cavities. Hive occupancy of stingless honey-bees varied with sea- There are forests patches in Udzungwa under local com- son, and this could be related to variation in humid- munity management where apiaries and meliponiaries have ity, where high air moisture content caused the bees to been established by local beekeeping groups. These forests abscond. For example, there was low occupancy during should be used for education and creation of awareness the wet season from February to April. Hive occupancy of on the need to manage the resources in these areas more D. schmidti decreased to zero between August and October sustainably. during the hot dry season, presumably due to high tem- peratures. High occupancy registered between May and Characteristics of colonies, behaviours and tree hosts October for M. ogouensis and M. lendliana was proba- Stingless honey-bee colony characteristics and bee bly due to swarming, resulting from colony disturbances behaviours, such as location of nesting, are useful in by honey hunters. In general, the number of experimen- species characterisation and may be useful in identifica- tal hives occupied by colonies of stingless honey-bee tion; for example, construction of colonies on tree branches stabilised for a period of only 3 months and decreased pro- at great heights above the ground for D. schmidti.The gressively thereafter due to poor insulation. Large-bodied behaviour of nest-building on tree branches has also been species of stingless honey-bee such as the M. ogouensis observed in Trigona dallatorea in Brazil (Michener 1974). and M. lendliana produced more honey relative to small- M. ogouensis nested in cavities close to the ground or even bodied species such as P. hildebrandti. Such productivity in vacated or live nests of termites and ants. M. lendliana could be attributed to the capacity of such species to collect nested in tree cavities located slightly higher but not on the and store honey. There was higher honey production in the ground. Nesting in subterranean areas observed in transi- buffer zone than in either the core or transition zone due to tional areas in Udzungwa was related to the absence of a higher number of colonies and higher occupancy. Honey suitable nesting tree cavities, due to severe deforestation pots were found in clusters segregated from brood cham- by the local communities. bers and pollen. The quantity of honey produced per colony The variations in nesting locations of the various of stingless honey-bee in the study area was high enough to species constitute a strategy to avoid interspecific com- make meliponiculture using nune boxes viable. In northern petition, as well as a strategy for protection, as was the Tanzania, especially in Kilimanjaro, log hives are used for case with D. schmidti, which constructed nests on tree stingless honey-beekeeping, but there is a need to design branches at great heights. Occasionally two or more nests modern box hives for beekeeping of stingless bees in the of the same species shared a cavity, raising the question Udzungwa area. of tolerance among members of different colonies. It is The amount of honey stored in the colonies varied enor- also not clear what were the reasons for nest stratification, mously among species, with some such as M. ogouensis but for stingless honey-bees, defence and diversification and M. lendliana storing as much as 3 L per colony per of resource utilisation could be important strategies for year. Other species like D. schmidti and P. hildebrandti 94 M.A. Njau et al. Figure 1. Map of Tanzania showing distribution of stingless honey-bee species. stored 1.6 L and 0.6 L, respectively. Mean productivity into the Udzungwa area for human needs and as a viable of honey per hive per colony in a season is usually low, additional income-generating activity. This will inevitably for example, Melipona beecheii in Costa Rica produces increase awareness of the importance of conservation of on average only about 2.63 L (Cortopassi-Laurino et al. the local forests and the Udzungwa ecosystem for local 2005). When compared to honey production by the stinging human communities around the game park. Interestingly, it honey-bees, which is usually over 20 L, the small amount was found that stingless honey-bee diversity declined with of honey stored by stingless bees may not be very attractive altitude. Whereas all species could be found in the low- to potential beekeepers. Only some stingless honey-bee altitude area at the foot of the mountains, only one species species store honey in quantities that could be harvested was found on the slopes at high altitude (1500 m a.s.l.). No by humans. However, in the Udzungwa area, local people wonder the occurrence of stingless honey-bees is confined do not usually keep stingless honey-bees but they hunt for to the warmer tropical regions of the world. In Africa, stin- honey, which is used for purposes of worship and medicine. gless honey-bees occur mostly within the tropics (Eardley It would be of much benefit to introduce meliponiculture 2004). 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Zoologia. 4:47–50. nity to acquire knowledge of the biodiversity of the area. Njau MA, Mpuya P, Mturi FA. 2009. Apiculture potential in pro- There should be a systematic method for studying the tected areas: the case of Udzungwa Mountains National Park, species composition of these forests and to find out ways Tanzania. Int J Biodivers Sci Manag. 5(2):95–101. of increasing such bee species. Information should also Nogueira-Neto P. 1970. Behavioral problems related to the pil- be collected on the various threats to the survival of the lages made by some parasitic stingless bees (Meliponinae: Apidae). In: Aronson LR, Tobach E, Lehrman DS, Rosenblatt biodiversity and management procedures should be insti- JS, editors. Development and evolution of behavior. Essays tuted for wise use of the resources in a manner that will in memory of T.C. Schnerlan. San Francisco (CA): W.H. be sustainable and beneficial to all stakeholders, and espe- Freeman & Co. p. 416–434. cially the local community itself. We therefore recommend Rasmussen C, Cameron SA. 2007. A molecular phylogeny of the the development of eco-villages in these and similar areas Old World stingless bees (Hymenoptera: Apidae: Meliponini) and the non-monophyly of the large genus Trigona.Syst in order to achieve good management of the resident Entomol. 32:26–39. biodiversity. Roubik DW. 1992. Stingless bees: a guide to the Panamanian and Mesoamerican species and their nests (Hymenoptera: Apidae; Meliponinae) In: Quintero D, Aiello A, editors. Acknowledgements Insects of Panama and Mesoamerica Vol. 2. Oxford (UK): Oxford University Press. p. 495–524. We wish to thank the management of Udzungwa Mountains Roubik DW. 2006. Stingless bee nesting biology. Apidologie. National Park and the people living around the park, espe- 37:124–143. cially the beekeeping groups for their assistance and cooperation Sommeijer MJ. 1999. A regional program for training and throughout the study. We also thank WWF for funding the study. research on tropical beekeeping and tropical bees in Costa Rica. Bee World. 80(2):70–79. Van Veen JW, Arce Arce HG, Sommeijer MJ. 2004. Production References of queens and drones in Melipona beecheii (Meliponini) in Byarugaba D. 2004. Stingless bees (Hymenoptera: Apidae) of relation to colony development and resource availability. Proc Bwindi impenetrable forest Uganda and Abayanda indige- Netherlands Entomol Soc. 15:035–039. nous knowledge. Int J Trop Insect Sci. 24(1):117–121. Wille A. 1979. Phylogeny and relationship among the genera and Cortopassi-Laurino M, Imperatriz-Fonseca VL, Roubik DW, subgenera of the stingless bees (Meliponinae) of the world. 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Journal

International Journal of Biodiversity Science, Ecosystem Services & Management – Taylor & Francis

Published: Dec 1, 2010

Keywords: biodiversity; management; honey; stingless honey-bee species; Udzungwa

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