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Spatial Aspects of Bumble Bee (Bombus Spp.: Apidae)

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Spatial Aspects of Bumble Bee (Bombus Spp.: Apidae) SPATIAL ASPECTS OF ,.,.. ,..... , .. - :.;.:.:.:.: ~.:.. BUMBLE BEE (BOMB US SPP.: APIDAE) FORAGING IN FARM LANDSCAPES A Thesis submitted in partial fulfilment of-the requirements for the degree of Master of Applied Science at Lincoln University by M.l. Schaffer Lincoln University 1997 Frontispiece. A large earth bumble bee worker Bombus terrestris (L.) foraging on lucerne flowers Medicago sativa L .. The bee's wings show one of the coloured fluorescent powders used during a mark-reobservation study in this thesis to trace movements by bumble bees in a commercial lucerne seed crop. 111 Abstract of a thesis submitted in partial fulfilment of the requirements for the Degree of M.Appl.Sci. SPATIAL ASPECTS OF BUMBLE BEE (BOMBUS SPP.: APIDAE) FORAGING IN FARM LANDSCAPES by M.J. Schaffer Bumble bees (Bombus spp.: Apidae) are valuable pollinators of many crop and wildflower species. However, in some situations their potential is limited. Evaluation of, and management to improve bumble bee efficacy should include spatial infonnation which is currently limited. Distance and direction determine the success of gene flow via pollen cross-over within and between plant populations at several scales. Studies of movement by bumble bees at large scales in semi-natural and intensively managed habitats are scarce. Few studies of bumble bee dispersal from the nest exist, particularly in relation to crops. At a small scale, directional rather than random movement between flowers has benefits for pollen flow. Results to date of directionality studies at small scales and their interpretation are inconsistent. The purpose of this thesis was to assess distances and directions moved by foraging bumble bees at a range of scales in two contrasting farm habitats in order to predict their pollination potential. A novel method was developed to mark automatically all the occupants of nests of bumble bees B. terrestris (L.) placed around a lucerne seed crop Medicago sativa L. in New Zealand. Reobservation data from eight nests showed that of bumble bees which foraged within the crop, 81 % travelled::;; 50 m and 56% ::;; 20 m from their nest. Results should be interpreted with extreme caution because fewer than 1% of bumble bees marked at nests were reobserved in the crop. Because it was not established where the other 99% of the bumble bees went, foraging areas for nests could not be calculated as anticipated. Theories to explain the non­ specificity of bumble bees to the crop include; resource depletion near nests, competition with honey bees in the crop, or an evolved strategy to disperse in order to minimise nest predation. IV Lucerne flowers contained a significantly lower concentration of sugar in nectar, and significantly fewer pollen grains than did those of purple loosestrife Lythrum salicaria L., a species on which bumble bees appeared to forage in preference lucerne. The higher rewards offered by L. salicaria may have diverted bumble bees from the less-rewarding lucerne crop. In a Norwegian meadow system, all foraging bouts by bumble bees B. lucorum (L.) within a patch of wood cranesbill flowers Geranium sylvaticum L. were random with respect to direction. This result is not consistent with predictions, based on optimal foraging theory, that movement should be directional to enable optimal pollen flow, and to avoid revisitation of just­ emptied flowers by the pollinator. A medium-scale study of several bumble bee species moving between patches of northern wolfsbane Aconitum septentrionale Koelle in Norway revealed considerable loyalty by bumble bees to patches in which they were marked. In a different landscape-scale study (over 5 ha), several bumble bees exhibited a high degree of loyalty to areas in which they were marked (87% were reobserved::; 50 m from marking points). These restricted movement patterns are discussed in terms of potential pollen flow. Of 260 bumble bees marked, only five were recorded crossing between meadows, which could be a result of innate loyalty to small forage areas, an artefact of the sampling technique used, or forest boundaries acting as physical impediments to movement. In the future, spatial data of the type collected in this thesis will aid in the management of bumble bee populations to achieve both commercial and conservation goals. Spatial data can be applied to predict the optimal placement of artificially-reared nests, predict suitable isolation distances for pure seed crops, and aid in the positioning of supplementary forage sources and nest-site refuges. Keywords: Bumble bee, Bombus, foraging behaviour, pollination, movement, spatial approach, scale, distance, direction, foraging area, New Zealand, seed crop, Norway, meadow, pollen, nectar. v CONTENTS Page Abstract iii Contents v List of Tables viii List of Figures viii List of Plates x Chapter 1 General introduction 1 Chapter 2 Literature review and background to the study 5 2.1 .. Introduction . 5 2.2 Bumble bees (Bombus spp.) as pollinators 5 2.2.1 General biology and ecology of bumble bees 5 2.2.2 Bumble bee foraging biology and pollination 8 2.3 The importance of pollination to fannland flora 11 2.3.1 Conservation areas in Europe: "The Cultural Landscape" 12 2.3.2 Commercial crops in New Zealand: Legume seed crops 14 2.4 Previous insect movement studies 17 2.4.1 Movement studies of foraging bees (Hymenoptera) 18 2.4.2 The spatial approach and a review of bumble bee movement studies 19 2.5 The influence of plant features on bumble bee movement 29 2.6 Summary 30 vi Chapter 3 Bombus spp. movement in a semi-natural habitat 31 3.1 Introduction 31 3.2 Site description 32 3.3 Movement by individual bumble bees within patches of flowers 36 3.3.1 Introduction 36 3.3.2 Methods 38 3.3.3 Results 40 3.3.4 Discussion 44 3.4 Movement by bumble bees between patches and between meadows 49 3.4.1 Introduction 49 3.4.2 .. Methods 51 3.4.3 Results 55 3.4.4 Discussion 58 Chapter 4 Bombus terrestris movement in an intensively-managed habitat 67 4.1 Introduction 67 4.2 Methods 72 4.3 Results 78 4.4 Discussion 89 Chapter 5 Pollen and nectar rewards in two plant species visited by Bombus terrestris 98 5.1 Introduction 98 5.2 Methods 103 5.3 Results 108 5.4 Discussion 112 vii Chapter 6 General discussion and conclusions 122 Acknowledgements 130 References 132 Appendicies 149 viii LIST OF TABLES Table Page 3.1 Proportions of bumble bees marked from differently colour-coded meadow areas, and the range of plant species on which they were caught 54 3.2 Breakdown of reobservation data for marked bumble bees including species, caste, colour, and flower species on which bumble bees were observed 57 3.3 Bumble bees which crossed forest boundaries to different meadows during a large-scale mark-reobservation project 58 4. 1 Flight distances for bumble bees from eight nests bordering a flowering lucerne crop 80 5.1 Sugar content of nectar in some common bee-visited flower species (after Beutler 1949) 101 LIST OF FIGURES Figure Page 2.1 Generalised bumble bee life cycle in New Zealand: 1. Presence of brood. 2. Stage of nest development. 3. Periods when each caste is dominant on flowers. (source: Donovan & Macfarlane 1984) 7 3.1 The Norwegian meadow study site in Telemark county, south west Norway. The three meadows and forest between them constituted a total study area of approximately 5 ha. 34 3.2 Grid array over the G. sylvaticum patch in which movement patterns by B. lucorum workers were assessed over the patch as a whole, and between flowers 39 3.3 Frequency distribution of flight directions for each of five foraging bouts by B. lucorum on G. sylvaticum flowers. Vector lengths represent percentage of moves in each compass direction. n= total moves per bout (P > 0.05 in all cases) 42 3.4 Pooled directionality results from several foraging bouts by B. lucorum a) between flowers and b) over a patch of G. sylvaticum . Radar plots represent frequency distributions of the percentage of moves in each direction 43 (P> 0.05 in both cases) 3.5 Frequency distributions of flight distances for pooled foraging bouts by B. lucorum a) between flowers and b) over a patch of G. sylvaticum 43 3.6 Examples of marking codes used on individual bumble bees foraging on patches of A. septentrionale. Each oval represents a dorsal view of a bumble bee thorax divided into quarters. Colour spots are read left to right and assigned values from the table according to their upper or lower position . 52 3.7 Breakdown of numbers and species of bumble bees marked in each of six differently colour coded marking areas within the meadow system, and pooled data for all areas combined . 53 3.8 Frequency distribution of flight distances from 18 reobservations of individually-marked bumble bee between patches of A. septentrionale 55 3.9 Breakdown of bumble bee numbers and species reobserved following marking 56 IX 3.10 Frequency distribution of flight distances for all bumble bee reobserved in six meadow areas in transect walks after marking 57 4.1 Proportional breakdown of all colour-marked bumble bees observed in a lucerne crop (n=197) 79 4.2 Frequency distribution of flight distances for bumble bees observed within a flowering lucerne crop 80 4.3 Three-dimensional mesh plots representing a) bumble bee distribution and b) flower distribution in a lucerne crop, when eight commercially­ reared nests were placed on the crop perimeter. Peaks represent highest concentrations of bees and flowers. Nest colours: 1. Green, 2. Red, 3. Dark yellow, 4. Light orange, 5. Pink, 6. Dark orange, 7. Light yellow, 8. Blue 81 4.4 Bumble bee densities in a lucerne crop with flower densities overlaid as contour lines.
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