Pollination ecology of cultivated and wild raspberry (Rubus idaeus) and the behaviour of visiting insects
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Floral morphology and anthesis were studied in commercial and in wild populations of raspberry (Rubus idaeus). Young flowers offered both nectar and pollen, but medium and old flowers offered nectar only, in diminishing quantity. All the three types had similar ageing patterns and diurnal patterns of nectar secretion, but they differed significantly in the nectar standing crop. Variation in nectar secretion rates within the raspberry cultivars was examined; Glen Moy produced more nectar per flower, and more flowers per meter, than Glen Prosen and wild raspberry flowers. There was significant daily variation in secretion rate, individual flowers in all cultivars showing variable rates of secretion even on the same branch. Time of sampling, effects of insect visitors, flower age and weather conditions all showed significant relationships to nectar availability. The three raspberry types have in common certain insect species as visitors, the most abundant being bumble bees (Bombus lapidarius, B. lucorum, B. terrestris, B. pratorum and B. pascuorum). Apis mellifera, Andrena species and hover flies. Bumble bees were responsible for about 60% of all visits, with honey bees, Andrena and hover flies making up most the remaining visits. Bombus species were more abundant through the particular observation days and through the different seasons, and they were present at almost all times of observations irrespective of climatic conditions in the studied area. The foraging behaviour and activity patterns, pollen loads and pollinating efficiency of the Bombus spp., Apis and Andrena spp. were analysed in relation to plant phenology, anthesis and dehiscence and to climatic variables. All bees had substantial pollen deposited on their bodies during visits, though few specifically collected it. Bombus species were found to strongly select young flowers, especially early in the morning when pollen was most abundant, while Apis and Andrena species visited unselectively. Bumble bees also foraged over substantially longer periods of the day, and in poorer weather, and they visited more flowers per minute than Apis and Andrena species. Bombus species also carried more pollen grains on their bodies than Apis and Andrena species, and deposited more pollen on raspberry stigmas; and because they foraged over longer range, they transferred pollen grains for longer distance than Apis mellifera. The flight directionality of Bombus, Apis and Andrena species among the flowers of Glen Moy and Glen Prosen was analysed. Pollen flow was also studied using fluorescent dyes, in field experiments during 1993 and 1994. Bombus and Apis transferred dye particles (pollen mimics) to different extents in different directions in the field. All the three visitors showed a strong tendency to move in the south-north direction (the direction of the raspberry rows); this would lead to increase in the gene flow within the same row in the presence of pollen carry-over. Pollen was carried up to 60m by Bombus species and 35m by honey bees. The work presented in this thesis provides evidence that (at least in Scotland) bumble bees are likely to be more important as pollinators of raspberries than other visitors. Reason why Bombus may be the preferred insect pollinators in wild and cultivated areas are discussed.
Thesis, PhD Doctor of Philosophy
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