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Open ADV Disseration for Submission.Pdf The Pennsylvania State University The Graduate School College of Agricultural Sciences POLLEN NUTRITION, THE FOUNDATION OF BUMBLE BEE FORAGING A Dissertation in Entomology by Anthony Damiano Vaudo © 2016 Anthony Damiano Vaudo Submitted in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy December 2016 The dissertation of Anthony Damiano Vaudo was reviewed and approved* by the following: Christina M. Grozinger Distinguished Professor of Entomology Dissertation Co-Adviser Chair of Committee John F. Tooker Associate Professor of Entomology Dissertation Co-Adviser Harland M. Patch Research Associate of Entomology Special Member David A. Mortensen Professor of Weed and Applied Plant Ecology Heather Hines Assistant Professor of Biology and Entomology Gary W. Felton Professor and Department Head of Entomology *Signatures are on file in the Graduate School ii Abstract Angiosperms and insect pollinators, especially bees, share a rich ecological and evolutionary history in which the radiation of the groups occurred through coevolutionary processes. This is because flowers facilitate reproduction through the transfer of pollen by attracting bees to flowers, and providing bees their entire source of nutrition. Historically, it was believed that bees were innately destined to visit flowers that provided specific or attractive morphologies, colors, or scents, known as pollination syndromes. However, individuals within bee species may visit and collect resources from different plant species during the day, season, and across years. Bee nutrition is partitioned between floral nectar which provides energy (carbohydrates) for foraging bees to collect nutritionally complex pollen (protein, lipids, and micronutrients). Because pollen quality differs between plant species and affects the health and development of bee larvae and adults, we expect that bee species forage to collect the right balance of pollen nutrients from their host-plant species. Therefore, if bees have species-specific nutritional needs, it may explain the foraging patterns among host-plant species observed. In this dissertation, I reveal that bumble bees (Bombus impatiens, Hymenoptera: Apidae) may actively select and balance their pollen diet among host-plant species to meet their optimum nutritional needs of protein and lipids, or protein:lipid ratio. This discovery was confirmed by 1) relating bumble bee pollen foraging preferences among host-plant species to pollen nutritional quality 2) assessing bumble bee preferences for pollen isolated from the host-plant species, 3) assessing bumble bee preferences to single source pollen modified to different protein:lipid ratios and concentrations, 4) assessing bumble bee regulation of synthetic diets modified to different protein:lipid ratios and concentrations, and finally 5) verifying these results in field studies by determining the nutritional value of pollen collected by free foraging colonies in which colonies defended their nutrient intake ratios independent of landscape. This dissertation presents a new theory that bees may be floral generalists, but nutritional specialists, adding a new dimension to previous assumptions of pollination syndromes. The particular floral traits involved in syndromes (i.e. color, morphology, scent) may have evolved as discriminative stimuli that bees associate with reward quality. iii TABLE OF CONTENTS List of Tables ................................................................................................................................. vi List of Figures ............................................................................................................................... vii Preface ............................................................................................................................................ ix References ................................................................................................................................ xiii Acknowledgements ....................................................................................................................... xv Chapter 1. Bee nutrition and floral resource restoration ................................................................. 1 Highlights .................................................................................................................................... 1 Abstract ....................................................................................................................................... 1 Introduction ................................................................................................................................. 2 Bee Nutrition ............................................................................................................................... 2 Floral Resource Nutritional Diversity and Bee Foraging Behavior ............................................ 5 Nectar ...................................................................................................................................... 5 Pollen ...................................................................................................................................... 7 The Importance of Plant Diversity for Bee Health ..................................................................... 9 Applying Bee Nutrition to Floral Resource Habitat Restoration .............................................. 10 References ................................................................................................................................. 15 Chapter 2. Bumble Bees Exhibit Daily Behavioral Patterns in Pollen Foraging .......................... 27 Abstract ..................................................................................................................................... 27 Introduction ............................................................................................................................... 28 Materials and methods .............................................................................................................. 31 Insect and plant species ......................................................................................................... 31 Hoop house ........................................................................................................................... 32 Foraging data collection ........................................................................................................ 33 Floral display ........................................................................................................................ 35 Foraging data metrics ............................................................................................................ 36 Results ....................................................................................................................................... 37 Discussion ................................................................................................................................. 38 The implications of daily foraging patterns for studies of plant-pollinator interactions ...... 38 Foraging patterns reveal interplay between pollen quality and abundance .......................... 41 Conclusion ............................................................................................................................ 44 References ................................................................................................................................. 50 Chapter 3. Macronutrient ratios in pollen shape bumble bee (Bombus impatiens) foraging strategies and floral preferences .................................................................................................................... 54 Abstract ..................................................................................................................................... 54 Significance Statement .............................................................................................................. 55 Introduction ............................................................................................................................... 56 Results ....................................................................................................................................... 59 Host-plant pollen foraging preferences ................................................................................. 59 Isolated pollen feeding preference assay .............................................................................. 60 iv Modified pollen feeding preference assay ............................................................................ 61 Discussion ................................................................................................................................. 63 Materials and Methods .............................................................................................................. 68 Pollen nutritional analysis ..................................................................................................... 68 Assessing host-plant pollen foraging preferences ................................................................. 71 Isolated pollen feeding preference assay .............................................................................. 72 Modified pollen feeding preference assay ............................................................................ 74 References ................................................................................................................................
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