Bumble Bees and Their Parasites Across European Communities: Sphaerularia Bombi in Native and Non-Native Hosts
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Bumble bees and their parasites across European communities: Sphaerularia bombi in native and non-native hosts Thesis submitted for the degree of Doctor of Philosophy (PhD) University of London Royal Holloway, University of London Egham Surrey TW20 0EX Catherine Mary Jones September 2013 1 Declaration page I, Catherine Mary Jones, declare that this thesis and the work presented in it is entirely my own. Where I have consulted the work of others, it is always clearly stated. Catherine Mary Jones 2 Acknowledgments First and foremost, I would like to thank my supervisor, Mark Brown, for selecting me as his first PhD student at Royal Holloway, University of London. Without his support and encouragement, the four years of my research would not have been possible. I would also like to thank my advisor, Vincent Jansen, and the technical and administration staff in the School of Biological Sciences, especially Tracey Jeffries, Rob Prouse and Sue Baldwin. I am grateful to the BBSRC for the DTG to fund my research and the YES and Vitae courses that I attended. My thanks also go to Joe Colgan for sharing his knowledge about Irish bumble bees and nematodes, to Roland Loosli, Paul and Regula Schmid-Hempel for helping with my fieldwork in Zurich, to Joe Colgan, Jim Carolan, and Nicola Marples for helping with my fieldwork in Dublin, and to Heather Rumble for looking after my bees in the lab while I was away. I would like to thank all the members of the Brown Lab at Royal Holloway, for their support in the Bee Room and the Ecology Lab, especially Matthias Fürst and Gemma Baron - and for organising so many fabulous social events. My thanks also go to Matthias Fürst, for showing me how to extract DNA from tissue samples and prepare it to send for sequencing, to Frances Medaney, for always being willing to help in the Molecular lab and to Selena Brace for spending days (weeks?) helping me to edit, align and analyse my DNA sequences and building the phylogenetic tree. My thanks and love to Christine and Eric Jones for reading my first manuscript and asking questions about my research, and to Oscar, Percy, Daisy and Millie (who is no longer with us) for providing support as only a dog or cats can. Finally, many thanks, and lots of love, to Lindsay Pilling, for financial support, for practical advice and always being there to give me encouragement and a hug. 3 Abstract My research investigated the host-parasite relationships between Bombus species and their generalist parasites, considering both the host community and the parasite community. This research was based in Europe and thus the focus was on European Bombus spp. particularly B. terrestris and B. hypnorum, and their parasites species, particularly the nematode Sphaerularia bombi. Bumble bees are important pollinators and are in global decline. Thus investigating their parasites, one of the factors that may be driving the decline, is vital both for the continued provision of this ecosystem service and for the conservation of bumble bees. Initially I investigated the generalist endoparasites, particularly S. bombi, across three European populations of B. terrestris, collected in England, Switzerland and Ireland. I found that parasite prevalence differed across the European populations sampled e.g. the prevalence of S. bombi in B. terrestris queens is highest in Ireland. I also provide details of the bee husbandry and dissection methods used throughout my research. I focused on England to examine the prevalence and impact of parasite communities found in the non-native species, B. hypnorum, and in five native Bombus host species. I also estimated the genetic diversity of the non-native B. hypnorum, from both the invaded (i.e. England) and native range (i.e. continental Europe), and that of two native Bombus species, B. terrestris and B. lucorum. The invasive B. hypnorum had higher parasite prevalence and lower functional genetic diversity than native species. Although parasites had a higher impact on the invader’s fitness than on native species, parasites and low genetic diversity have not prevented the rapid invasion of the UK by B. hypnorum. Having observed that infected B. hypnorum queens do not appear to deposit S. bombi parasite larvae in their faeces, I quantified parasite reproduction in infected hosts to examine the competence of the non-native B. hypnorum as a host for this generalist parasite. I found that S. bombi larvae are not deposited in the faeces of infected B. hypnorum queens suggesting that B. hypnorum is not a competent host for S. bombi. The host–parasite relationship between S. bombi and this non-native 4 bumble bee may alter the relationships between S. bombi and congeneric native host species. Using standard molecular techniques, I investigated the phylogeography of S. bombi across Europe and asked whether the S. bombi parasites, found in non- native B. hypnorum in England, originate from England or from Continental Europe: Did the non-native B. hypnorum acquire parasites in the UK or were they co- introduced with the invading host? I found that the S. bombi population did not appear to be structured across the European native and non-native hosts sampled and therefore I was unable to establish whether these parasites were acquired or introduced. Finally, I discuss what I have discovered during my research, how has this work added to the current knowledge on the subject and which areas warrant further investigation. 5 Table of Contents Table of Contents ....................................................................................................... 6 Table of Figures ....................................................................................................... 11 Table of Tables ......................................................................................................... 12 Table of Plates ......................................................................................................... 13 Chapter 1. General introduction ......................................................................... 14 Introduction ........................................................................................................... 14 Species introductions and biological invasions ..................................................... 14 Parasites ............................................................................................................... 15 The role of parasites in biological invasions ......................................................... 15 Parasite spill-over and parasite spill-back ......................................................... 16 Parasite dilution ................................................................................................. 16 Reservoir host ................................................................................................... 17 Host competence .............................................................................................. 17 Enemy release hypothesis (ERH) ..................................................................... 17 Evolution of increased competitive ability (EICA) .............................................. 18 Local adaptation ................................................................................................... 18 Generation time ................................................................................................. 19 Migration rates .................................................................................................. 19 Virulence and transmission routes .................................................................... 19 Population size and reproduction ...................................................................... 20 Study system ........................................................................................................ 20 Host system: Bumble bees ................................................................................... 20 Global bumble bee declines .............................................................................. 20 Bumble bee lifecycle ......................................................................................... 21 Bumble bee global distribution .......................................................................... 21 Haplodiploidy ..................................................................................................... 21 Commercial bumble bees ................................................................................. 22 6 British bumble bees ........................................................................................... 22 B. hypnorum ...................................................................................................... 23 Bee husbandry in the laboratory ....................................................................... 24 Parasite systems .................................................................................................. 26 Sphaerularia bombi ........................................................................................... 26 Apicystis bombi ................................................................................................. 27 Crithidia bombi .................................................................................................. 27 Nosema bombi .................................................................................................. 28 Locustacarus buchneri .....................................................................................