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Chapter 18

Taxonomy as a Tool for Conservation of African Stingless Bees and Their Honey Connal Eardley and Peter Kwapong

We dedicate this chapter to Professor Charles Duncan Michener who has been a huge inspiration throughout our career. As ecology embodies taxonomy, the background created by Mich will endure and always be treasured.

18.1

Introduction

In Africa stingless bees are most diverse in the equatorial regions. To the north the Sahara Desert abruptly delimits their distribution. Southwards they become progressively less diverse reaching more or less the Tropic of Capricorn in the interior of the Subcontinent. Their distribution extends farther south along the East coast, and to a lesser extent along the west coast (Eardley 2004). Several species appear confined to the tropical wet forests. Most species, however, occur in both savannah and tropical forests, including the east African coastal forest (Eardley 2004). Two species have been recorded from desert areas, one occurs in the south-western Sahara (Hypotrigona penna Eardley) and there is an unpublished record of Liotrigona from the Richtersveld, South Africa. The African stingless bees are smaller than indigenous African honey bees Apis mellifera L. and their approximately 30 recognised subspecies (Ruttner 1988;

C. Eardley (*) Agricultural Research Council, Private Bag X134, Queenswood, 0121 Pretoria, South Africa School of Biological and Conservation Sciences, University of KwaZulu-Natal, Private Bag X01, Scottsville, Pietermaritzburg 3209, South Africa e-mail: [email protected] P. Kwapong Department of Entomology & Wildlife—International Stingless Bee Centre, School of Biological Sciences, University of Cape Coast, Cape Coast, Ghana 261 P. Vit et al. (eds.), Pot-Honey: A legacy of stingless bees, DOI 10.1007/978-1-4614-4960-7_18, © Springer Science+Business Media New York 2013

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Michener 2007). They also do not produce as much honey as the honey bee does, which has a larger flight range than meliponines. The robbing of honey bee nests by indigenous people has been practiced for millennia in Africa, as documented in rock paintings (Crane 1999; Johannsmeier 2001), and they are still being robbed in Africa (Eardley C, personal observations). Currently meliponiculture is practiced in tropical Africa, but for the most part its history has not been documented and its age is unknown. In Ghana, current stingless beekeeping only recently began as an activity complementary to beekeeping with Apis (Kwapong et al. 2010). It does not appear to have been practiced in southern Africa. However, stingless bee honey, although less in quantity, is highly sought in all of tropical Africa—primarily for its medicinal uses. It fetches higher prices than honey bee honey, and is culturally important. The current value of stingless bees, as pollinators, to biodiversity conservation and agriculture is unknown, but they do visit flowers of many different plants and crops, as seen in the field and often indicated on museum specimen labels. Being social they can possibly be more easily managed than solitary bees and the expansion of meliponiculture to agriculture should be further investigated (Roubik 1995). Vernacular names for stingless bees in South Africa are “mopani” bees or “mocca” bees. In Ghana several of the species are known by their common names: “anihammoa”, “duro kokoo”, “duro tuntum”, “mimina” and “tifuie”.

18.2 Taxonomy of Stingless Bees Prior to Eardley (2004) research articles on stingless bees of the Afrotropical Region were relatively few, and by a handful of authors (Ambougo-Atisso 1990; Darchen 1966, 1969a, b, 1970, 1971a, b, 1972a, b, 1973, 1977a, b, 1981, 1985; Darchen and Louis 1961; Darchen and Pain 1966, Fletcher and Crewe 1981a, b; Kajobe 2006, 2007a, b; Kajobe and Echazarreta 2005; Kajobe and Roubik 2006; Lobreau-Callen et al. 1986, 1990, 1994; Michener 1959; Moure 1961; Moritz and Crewe 1988; Portugal-Araújo 1955a, 1955b, 1956, 1958, 1963; Portugal-Araújo and Kerr 1959; Sakagami et al. 1977), excluding those that described new species. Moure (1961) provides keys for the identification of many African stingless bees, but understanding the small differences between species together with intraspecific variation still prevented confident identification of many species. Consequently a taxonomic revision, based on worker bee morphology, was undertaken (Eardley 2004). Since then a lot of interest has been shown in developing meliponiculture in West Africa (Kwapong et al. 2010). Meliponiculture has been practiced for a long time in East Africa but more recently research into foraging and nesting has been undertaken (Kajobe 2006, 2007a, b; Kajobe and Echazarreta 2005), while little interest has been generated in southern Africa. There has also been interest in documenting meliponines as pollinators and the medicinal use of their honey, but to date there are no substantial data for Africa. Eardley (2004) found that the material available in museums and comparative biological information in the literature were scant, in contrast with the great abundance

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Taxonomy as a Tool for Conservation of African Stingless Bees and Their Honey

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of these bees in the wild. As now recognised, differentiating many stingless bee species and some genera require microscopic or molecular studies (see Rasmussen and Cameron 2010, Chap. 1, in the present book), and cryptic species have been noted (Camargo and Pedro 2007). It is now widely believed that Eardley (2004) underrepresented the true diversity of the taxon (Macharia J, personal communication). Portugal-Araújo and Kerr (1959) discovered Hypotrigona araujoi (Michener) to be a distinct species through observation in a meliponary, and Michener (1959) subsequently discovered subtle differences between it and Hypotrigona gribodoi (Magretti). Darchen (1970, 1981) studied stingless bee biology in West Africa that led to the description of three new species; Meliponula (Axestotrigona) sawadogoi (Darchen), Meliponula (Axestotrigona) richardsi (Darchen) and Meliponula (Axestotrigona) eburnensis (Darchen) whose types have not yet been located. Joseph Macharia found differences in the nest architecture in the species that Eardley documented as Meliponula bocandei (Spinola) (Macharia J, personal communication) suggesting that this taxon is composite. Katherine Krause found size differences in the species that Eardley documented as H. gribodoi (Magretti) which indicate that H. gribodoi comprises more than one species. Further, the fact that the majority of species (10 out of a total of 18 species) occur in distinct habitats, such as tropical forest and dry savannah, suggests a potential greater species diversity than recorded by Eardley (2004) (Table 18.1). The difficulty in separating stingless bees based on morphology necessitates the need for new diagnostic tools. Nest architecture and host plant preferences pose logistical problems in gathering material for taxonomic revisions of genera and would be better suited to studies on differences between identified species rather than being used to recognise different species. The most promising tool for identifying morphologically similar species is evidently DNA barcodes, a method using a short genetic sequence to identify an organism, as suggested by Packer et al. (2009).

18.3

Host Plants and Nests of Stingless Bees

Knowing bee host plant usage is important for understanding pollination as well as the medicinal use of bee honey, as explained among the chapters in the present book, which include studies in Africa, Asia, Australia and the Neotropics. Stingless bees focus their foraging activities on a wide range of food plants. As a group they have been recorded visiting 135 plant genera (Eardley and Urban 2010). A preliminary survey of the data suggests that food plant overlap is greater within bee genera than between the genera. However, the data do not indicate if the bees are collecting pollen or nectar. Until the taxonomy is properly resolved, the degree of host plant specificity will not be understood. In Ghana, stingless bees have been collected from tropical rain forest canopies (Nuttman et al. 2011), crops growing on agricultural landscapes as well as on flowers of vegetables and medicinal plants. The most important native and introduced fruit crops on which stingless bees forage include mango, cashew, avocado, citrus, coconut, oil palm, shea butter tree, passion fruit, pepper and many others.

Table 18.1 African stingless bees and vegetation type in which they occur Ghanaian common Taxon names of bees Cleptotrigona cubiceps (Friese, 1912) Dactylurina schmidti (Stadelmann, 1895) Dactylurina staudingeri (Gribodo, 1893) “tifuie” Hypotrigona araujoi (Michener, 1959) “mimina” “anihammoa” Hypotrigona gribodoi (Magretti, 1884) “mimina” “anihammoa” Hypotrigona penna Eardley, 2004 “mimina” “anihammoa” Hypotrigona ruspolii (Magretti, 1898) “mimina” “anihammoa” Liotrigona bottegoi (Magretti, 1895) Meliponula (Axestotrigona) cameroonensis (Friese, 1990) Meliponula (Axestotrigona) ferruginea (Lepeletier, 1841) “duro tuntum” Meliponula (Meliplebeia) beccarii (Gribodo, 1879) Meliponula (Meliplebeia) griswoldorum Eardley, 2004 Meliponula (Meliplebeia) lendliana (Friese, 1900) Meliponula (Meliplebeia) nebulata (Smith, 1854) Meliponula (Meliplebeia) ogouensis (Vachal, 1903) Meliponula (Meliplebeia) roubiki Eardley, 2004 Meliponula (Meliponula) bocandei (Spinola, 1853) “duro kokoo” Plebeina hildebrandti (Friese, 1900) x x x x x x x x x x x x

x x

x

x x

x x x x x

x x

x x x x

Savannah including dry savannah and desert

Tropical forest including east coast forest

264 C. Eardley and P. Kwapong

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18.4

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Challenges to Stingless Bee Survival

Kwapong et al. (2010) discuss some of the challenges stingless bees face in Ghana in their booklet on their management and utilisation. Conservation of stingless bees is threatened by loss of habitat from logging, bush fires and wild honey hunting, pests and predators. As most stingless bees are arboreal, when trees are cut the colonies are lost. Bush fires which constantly sweep through tropical forest during dry season burn up trees or meliponary rustic hives harbouring stingless bee colonies. Quite a number of rural communities are aware of stingless bee nests. When harvesting honey they often burn the bees and thereby destroy the colonies. The most important obstacles facing domesticated colonies of stingless bees are predators and pests, notably the small hive beetle Aethina tumida Murray (Coleoptera: Nitidulidae) whose larvae destroy entire colonies. Hive beetle adults live in close association with both honey bees and stingless bees. If hive beetles get an opportunity to oviposit in a colony the eggs hatch and the larvae quickly destroy the colony or cause the bees to abandon the nest. Other predators such as lizards, ants and spiders also threaten stingless bee colonies.

18.5

Justification for Further Taxonomic Research

The species name is the main tool to access the existing information on biology. If the taxonomy is inadequate, accurate biological information cannot be disseminated. The increasing demands of the human population result in the need for more food. Many foods result from pollination, and therefore pollinators need to be properly studied (Roubik 1995). As agriculture intensifies, pollination management will become more important. Increased agriculture and urban sprawl will most likely also place more pressure on the natural environment, resulting in a greater need to conserve biodiversity and the habitat of these organisms. Social bees have an advantage in pollination management in that many individuals live in a colony and they can be moved more easily than solitary bees, but similarly the loss of a colony results in the loss of many pollinators. The ability to move pollinators also introduces the risk of moving them to areas where they do not naturally occur. Moving honey bees in South Africa has had some disastrous consequences, such as the production of the pseudoclone (Neumann and Hepburn 2002) which is a social parasite of Apis scutellata Lepeletier.

18.6

Conclusions

Through personal observations the authors’ impressions are that in East Africa the importance of stingless bees in traditional medicine is well appreciated and widely used by traditional healers. Here meliponiculture is practiced, but the detailed uses for the honey appear to be trade secrets. In other parts of Africa stingless bee honey

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appears to be less widely used for medical purposes, and if meliponiculture exists it is uncommon. Its wide use for food is mainly through nest robbing. Meliponiculture for agriculture is limited and very recent. Consequently, little is published on stingless bees in Africa. Nevertheless it appears from the limited studies that have recently taken place that stingless bees are an invaluable resource in Africa for biodiversity conservation, agriculture and medicine. A number of scientists throughout the continent are showing an interest in studying these bees and in the future their biology and honey should become better documented. There is clearly a need for an updated taxonomic revision of the African stingless bees, following the recent advance made by Eardley (2004). This need is justified by their apparent importance as pollinators for agriculture and biodiversity conservation. DNA barcoding could be introduced as a complementary tool for separating stingless bee taxa and facilitate the recognition of those morphological characters that are useful in separating species. A study should be undertaken that systematically surveys the stingless bees of Africa to maximise the likelihood of discovering the entire fauna and to document their biogeography. Where possible, host plants and nest architecture should also be documented, which provides a tool for identifying bees in the field. The data should be stored, using relational database technology, in such a way that they will be useful for research including biogeographic analyses, phylogeny and pollination ecology. Finally, before the honey, its composition and uses can be studied, the taxonomy of all living species needs further consideration. Acknowledgements Dr. Janine Kelly and Dr. Claus Rasmussen are thanked for critical reading of the manuscript. Editorial support was kindly provided by P. Vit, S.R.M. Pedro and D.W. Roubik.

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