TABLE OF CONTENTS:
LIST OF TABLES
LIST OF ACRONYMS
3. GENERAL ANALYSIS
6. GENERAL RECOMMENDATIONS
LIST OF TABLES:
Table 1: A summary of the races of the Western honey bee, Apis mellifera, their Behavioral and Morphological Description, and Geographical Distribution
Table II: A summary of the species and races of Oriental honey bees, their Behavioral and Morphological Description, and Geographical Distribution
Table 3: A summary of Bee Diseases and Parasites, their Geographical Distribution, Symptoms and Management Measures
Table IV: A summary of Bee Pests and Predators; the Damage they cause and their Geographical Distribution and Management Measures
Table 5: A summary of some pesticides highly toxic to bees (LD50: 0.001-1.99 µg/bee)
Table VI: A summary of some pesticides moderately toxic to bees (LD50: 2.0-10.99 µg/bee)
Table 7: A summary of some pesticides relatively nontoxic to bees
LIST OF ACRONYMS:
Abbildung in dieser Leseprobe nicht enthalten
Bees evolved in specific areas of the world long ago, before they spread to become globally as they are today (Tables 1 and II), according to Kugonza (2009). As they spread, they became adapted to the local ecological conditions of the different areas, changing in morphology and behaviour to fit within the requirements of the ecosystem, giving rise to a wide bee biodiversity of bee species and races we see today. Bees are classified under Animal Kingdom, Phylum Arthropoda. They belong to Class Insecta, which is divided into 29 Orders. Bees belong to the order Hymenoptera, which has three Super families, namely: Apoidea (bees), Formicoidea (ants) and Vespoidea (wasps). There are around 30,000 named species of bees (Apoidea). Apoidea is further divided into several Families, namely Apidae (social bees), colletidae, Andrenidae, Halictidae, Melittidae, Megachilidae, and Anthophoridae. Most of the Families have solitary individuals: each female bee makes her own nest, lays a single egg and provides food for the single larva that develops. However, a high level of social development is shown by the species in Apidae where the individuals live together in a permanent, large colony, headed by a single egg-laying queen (B f D, 2003c). Apidae is composed of four genera: Apis (honeybees), Trigona and Melipona (stingless bees), and Bombus (bumble bees). According to MAAIF (2012a), the genus Apis is comprised of 5 main species of honey bees: Apis dorsata (the giant honey bee); Apis laboriosa (the darker giant honey bee); Apis florea (the little or dwarf honey bee) ; Apis cerana (formerly Apis indica), is the eastern hive honey bee ; and Apis mellifera (western honey bee). These species have evolved and differentiated into more species and races of honey bees. Kugonza (2009) described 9 species of honey bees in the world, which Oldroyd and Wongsiri (2006) grouped under 3 subgenera: Micrapis (Apis florea and A. andreniformis) , Megapis (A. dorsata and A. laboriosa) and Apis (A. cerana, recently recognized as separate races of A. nigrocinta, A. koschevnikovi and A. nuluensis, and A. mellifera) . Dietz (1992), Hussein (2000) and Wikipedia (2012) described over 28 races of A. mellifera alone. Beekeeping started with honey bees (Apis species), a practice called Apiculture, although keeping of stingless bees, belonging to the genera Trigona and Melipona, a practice called Meliponiculture, has recently picked up, as reported by B f D (2003a), Fajardo and Cervancia (2003) and Braga et al., (2009). The main focus of this paper is Apiculture. Evidence of rock paintings on mountain shelters in Spain point that honey bee (Apis) hunting existed as early as 7000 B.C. (Mesolithic times). With time and increasing human population and pressure, the honey bee resource started disappearing. To reduce the hardship and unpredictability of harvesting from wild honey bee colonies, people found ways to increase ownership and management of honey bee colonies kept in hives. The earliest hive probably was a log from a fallen tree in which wild honey bees had nested. Cork and other types of tree bark were then used for hive making and later on, hives were made from planks cut from tree trunks. The first centres of honey bee culture were in the Middle East, in dry and open country (most likely Tropical India and Southeast Asia). The first recognized hives were pottery vessels made during the Neolithic period, from about 5000 B.C. onwards. Woven baskets used as hives came much later on. At this stage protection was provided to the honey bee colony in return for periodic harvests of honey. The idea was to maintain the colony for future harvests instead of destroying it for a one-time harvest. Little was understood as to what was going on inside the hive since events could not be seen. It was not also realized that the large bee was actually a female, the queen bee, which is the mother of all the bees in the hive. It was mistakenly referred to as the “King bee”. The sexes of the workers and drones and the facts of mating between the queen and drones were not known. Neither did man know about bees themselves secreting wax used to build the comb. The relationship between bees, flowers and formation of seeds and fruits was also not known (Kugonza, 2009; and MAAIF, 2012a). Studying the behaviour of bees inside their nest in dark cavities with small entrances was not possible until the development of the glass-walled observation hive about 200 years ago (Gary, 1992). This advancement helped to understand the lifecycle of bees, bee biology, activities and behaviour of bees inside the hive. However, still, no suitable hive was found until 1891 when Lorenzo Lorraine Langstroth, an American Church minister, discovered the concept of bee space and made the first commercial bee hive, which to-date bears his name. Langstroth is now accepted as the father of modern beekeeping. Today, man is able to exploit the honey bee resource, to a certain degree of sustainability, for commercial pollination services, honey, beeswax, pollen, propolis and bee venom. Clearly from the foregoing explanations, so far, advancements in beekeeping have hinged on man’s understanding of the activities and behaviour of bees. Although, Kenyan Top Bar (KTB) hive, a movable-comb hive technology, has been invented alongside the Langstroth (movable-frame) hive technology to suit tropical African bees, it has not worked well. The bees are still less settled, more nomadic or migratory, annual, more aggressive/defensive and sting prone, with higher absconding and swarming rates, and are therefore more difficult to manage than their temperate counter parts (Paterson, 1999; and Rinderer, 1999). This paper therefore reviews the various honey bee species and races and the characteristic behavior of each species and race, the factors affecting the behavior, and how this knowledge in totality is applied to better manage the honey bee resource.
Honey bee activities and behaviour can be categorized into 2; those displayed under normal conditions and the others displayed under conditions of stress or disturbance. In both cases, the bees are responding to factors or stimuli in their immediate internal and external environments, which are detected by their sensory cells. The bees react to the stimuli in a particular (stereo-typed) way, because their nervous systems are “hard-wired” or “programmed” genetically to react in this manner (Gary, 1992), hence their behaviour. This section, therefore, describes the internal and external stimuli that bees respond to, the pattern of their responses and how the behavioral responses can be manipulated for the benefit of mankind. The various honey bee species and races, their characteristic behaviors and geographical distribution are also described here (Tables 1 and II).
1. The internal factors affecting honey bee activities and behaviour include:-
(i) The caste and sex of the bee: These determine what activities a particular bee will carry out in the colony. For example, the queen is a female with well developed ovaries and the only fertile female in the hive. Its main function is to lay eggs. The only function of the drone bee, which is male, is to mate with the queen. The role of the queen bee and the drone bee is, therefore, to reproduce the species. In addition to the reproductive role, the queen bee controls the activities of all the bees in the colony through pheromone communication. While, worker bees are sterile females and perform a number of activities in the colony including cell/house cleaning; brood, queen and drone feeding; wax secretion and comb building; food transmission and processing; colony defence; undertaking; thermoregulation; foraging; and robbing (Kugonza, 2009; and MAAIF, 2012a). This knowledge is very crucial for the beekeeper to understand which caste is not performing its duty and why, and then design appropriate remedial actions to be taken.
(ii) Stage of development: The activities of worker bees vary with age of development, a phenomenon described by Gary (1992) as “temporal division of labour” or “age polyethism”. For example, within a day or so after emergence, young worker bees begin to feed nectar, diluted honey and pollen to larvae more than 3 days old. During the first 3 days after emergence, they typically clean the hives from which bees have recently emerged. At approximately 6 to 12 days of age, after their brood food (hypopharyngeal) glands are mature enough to secret royal jelly, they begin to feed young larvae less than three days old. Wax secretion starts until the glands are mature enough. Furthermore, stinging behaviour starts when the stinging structure has developed fully. Also, flight is impossible for very young bees because their nervous system and muscles are not fully developed. This aspect of knowledge can help in designing colony multiplication, queen rearing, royal jelly and bee venom production projects.
(iii) Hormones: The hormone system of bees consists of two hormones. The juvenile hormone, in the larva, is responsible for slowing down the rate of development from the stage of pupa to adult. The ecdysone (moulting) hormone is an antagonist to the juvenile hormone and is responsible for quickening the process of development. The ratio between the hormones determines the duration of development (Kugonza, 2009). These hormones may be of use to bee breeders. They may also be of use in bee pest and disease management packaging to delay or quicken the development of brood in relation to the period of occurrence of brood pests or diseases.
(iv) Physical stimuli inside the body of a bee also affect behaviour. For example, the sensing of the stretching of the honey stomach either stimulates or inhibits feeding (Gary, 1992). The knowledge of this fact helps one to determine and plan diurnal feeding requirements of his or her honey bee colonies.
(v) Genetic composition: The genetic composition of the bee is now known to exert a major effect on behaviour. Each bee tends to express different behaviors according to its genetic profile. For example, Rothenbuhler (1968) in Gary (1992) demonstrated that 2 specific genes control the house cleaning behaviour of worker bees. Worker bees endowed with one of the genes uncapped the cells, but did not remove the dead brood. Other workers that contained the other gene did not uncap cells, but removed dead larvae or pupae from cells that had been uncapped. Enough workers with both genes must be present in the colony for effective house cleaning activity. This has a bearing on disease resistance, especially the American foulbrood (Gary, 1992). Genetically controlled behavioral characters of bees have a wider application in selection for desirable qualities and breeding, and biotechnology.
(vi) Bee activities frequently are regulated by internal, physiological “time clocks” that trigger specific behaviors at specific times, especially on the 24 hour cycle rhythm. Bees, therefore, remember the time of day and tend to arrive at nectar and pollen sources at the correct time (Gary, 1992) or to cease foraging activities especially at night when there is complete darkness (Paliwal et al. 2005). This information helps to design appropriate confinement time table for bees during a pesticide spraying programme in an area, as reported by Rashad, et al. (1985) or to schedule harvesting at night.
(vii) The species and race of bees makes them behave differently characteristic of that species or race. Specific management packages can be developed for each species or race based on its activities and behavior.
2. The external factors affecting bee activities and behaviour are:-
(i) Sounds, chemicals or odors, touch, light and magnetic fields are detected by thousands of specialized sensory cells. Nerve impulses from these cells speed along neural pathways of the nervous system and cause the bee to behave in a stereotyped manner when stimulated by the appropriate mixture or “configuration” of stimuli. This fact has led to the discovery of the pheromone communication system in bees, and the development of lures for bee management (Ferguson, 1985), although Gary (1992) deviated from the principle slightly, saying individual bees do not always respond the same way to identical stimuli, owing to their differences in sensitivity.
(ii) Climatic and ecological factors: Bees in regions with generally low temperatures and reliable rainfall pattern, especially in temperate climates, tend to be gentle and perennial, making their management easy. While in the tropical Africa, where there are generally high temperatures, the bees are nomadic or migratory, annual, more defensive and prone to sting, making their management difficult up to today. The available packages for tropical African honey bee management are imported technologies, were developed based on the European honey bee behaviour, and are therefore not satisfactory. It is the very reason for this study to find out more about honey bee behavior, particularly the African honey bees! Bee behavior also varies with specific adaptation to different ecosystems. For example, there are 3 subspecies of Apis mellifera which inhabit the East African ecological zones. A. m. litorea occupies the coastal plain below 500 metres above sea level. Above the coastal escarpment, the major subspecies, A. m. scutellata, is found. This species ranges throughout most of East Africa. However, where mountains cause rain shadows, that rainfall stability has caused the evolution of a third species, A. m. monticola. This subspecies is found between 2,000 and 3,000 metres above sea level. In areas where the ranges of these 3 subspecies meet, hybrid zones exist where the honey bees have intermediate characteristics. Hence the 3 species form a graded, stepwise ecocline along the altitudinal range. A. m. monticola is considered to be more perennial, gentle and a very good honey producer, A. m. scutellata is migratory, prone to abscond and sting, and in comparison, a poorer honey producer. A. m. litorea is similar to A. m. scutellata, except it is more prone to sting and perhaps less migratory due to living in a range with somewhat more predictable rain. East African beekeeping ecological zones with their different subspecies are a useful genetic preserve. A. m. scutellata - A. m. monticola hybrid zone may provide breeding stock that better suits beekeepers and is still adapted to local ecology (Ruttner and Kauhausen, 1985; Rinderer, 1999). Overall, there is a wide bee biodiversity in the world (Tables 1 and II) because of specific adaptation to different climatic and ecological factors.