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Drivers of the Diversity and Distribution of Wild Bees in a Species Poor Region

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Drivers of the Diversity and Distribution of Wild Bees in a Species Poor Region Norwegian University of Life Sciences Faculty of Environmental Science and Technology Department of Ecology and Natural Resource Management Philosophiae Doctor (PhD) Thesis 2016:23 Drivers of the diversity and distribution of wild bees in a species poor region Pådriverne bak diversitet og utbredelse av villbier i en artsfattig region Markus Arne Kjær Sydenham Drivers of the diversity and distribution of wild bees in a species poor region Pådriverne bak diversitet og utbredelse av villbier i en artsfattig region Philosophiae Doctor (PhD) Thesis Markus Arne Kjær Sydenham Department of Ecology and Natural Resource Management Faculty of Environmental Science and Technology Norwegian University of Life Sciences Ås 2016 Thesis number 2016:23 ISSN 1894-6402 ISBN 978-82-575-1352-8 Ph.D. supervisors Professor Stein Ragnar Moe – main supervisor, Dr. Katrine Eldegard, Professor Ørjan Totland, Department of Ecology and Natural Resource Management, Norwegian University of Life Sciences, NO-1432 Ås, Norway PhD evaluation committee Professor Jens Mogens Olesen, Department of Bioscience – Genetics, Ecology and Evolution, Århus University, DK-8000 Århus, Denmark Professor John-Arvid Grytnes, Department of Biology, University of Bergen, NO-5020 Bergen, Norway Committee administrator: Professor Tone Birkemoe, Department of Ecology and Natural Resource Management, Norwegian University of Life Sciences, NO-1432 Ås, Norway ii Acknowledgements I would like to thank my supervisors for their guidance and mentoring during the past three years. My main supervisor, Prof. Stein R. Moe for giving me the freedom to juggle different research ideas but at the same time being there to show me how to contextualize and develop them into testable hypotheses. Thank you also for always having your door open and so generously revising, discussing and following the production of the manuscripts that have led to this thesis. Prof. Ørjan Totland, I am grateful to you for introducing me to the world of pollinators and that of the people who study them. Thank you also for all your ideas, comments and suggestions that greatly improved the manuscripts in this thesis. Dr. Katrine Eldegard, thank you for being so generous with your time and for guiding and supporting me while developing research ideas. Thank you also for introducing me to the world of statistical modelling and for always being available to provide improvements of manuscripts and ideas as well as a supportive pat on the shoulder. I am grateful to Statnett for funding the fieldwork and species identification that made this thesis possible. Studying at the Dept. of Ecology and Natural Resource Management (INA) has been a fantastic experience, much thanks to the great working environment there. I would like to thank everyone in the tech-admin group (Annie, Gunnar, Grethe, Kari T., Mette, Ole Wiggo, Stig D. and Ågot) for your patience and for making things run so smoothly. I would especially like to thank Tone G. for making long days in the entomology lab a lot more enjoyable. I would like to thank the co-authors of the manuscripts that led to this thesis. Collaborating with you has been a great source of motivation. Lise D. Häusler for your hard work in the field and your work in the lab. Diana N. Stanescu-Yadav for your work in the lab. Dr. Stein J. Hegland and Dr. Anders Nielsen for your contributions and thorough revisions of our manuscript and for the great times at SCAPE. Prof. Simon G. Potts, Prof. Michael Kuhlmann and Mr. Stuart P.M. Roberts for generously contributing traits data for the Norwegian bees and for all your constructive revisions and insightful comments on our manuscript. I would like to thank all editors and anonymous reviewers whose strenuous efforts greatly improved this thesis. Mike and Sue Edwards, Stuart P.M. Roberts, David Baldock and everyone else at the Bees Wasps and Ants Recording Society (BWARS), thank you for providing a welcoming environment for everyone interested in learning about Aculeates. Thank you Mike and Sue for iii all your hospitality and generosity and for introducing me to bee taxonomy. On that note, I would also like to thank Prof. Emer. Eline and Sigmund Hågvar for talking me into getting started with bees in 2010. I would also like to thank Frode Ødegaard at the Norwegian Institute for Nature Research for taking your time to verify problematic bee specimens and Lars O. Hansen at the Natural History Museum in Oslo for granting me access to the bee collections. To all my friends, present and former colleagues at INA, you have made long days seem too short. Prof. Kari K. and Prof. Douglas Sheil, working with you to get the Ecology Research Group up and running has been great fun. Prof. Vidar Selås, Dr. Mari H. Lie and Dr. Ronny Steen for the great trips/field courses in Jotunheimen. Prof. Michael Ohlson and Dr. Richard Bischof for taking me on as a teaching assistant, I am grateful for that experience. Mari, thank you for fun trips in the field and to Denmark and Scotland. My good friend and office mate Christian, thank you for your friendship and for reminding me that Kattegat is never far away. Rannveig, John (best-dressed Dane on the slopes), Massimo (for your company in the Oslo café’s offices), Hanne, Anne, Carolyn, Walid (for keeping the A-line represented at INA), Sam, Ronny, Jens, Erik, Fredrick and Richard for making the spots in between a lot more fun. Many people outside my studies have also contributed with their support and friendship. Adrian, thank you for your friendship, assistance in the field and our bee safari to DK. Embret for our skiing trips and nights out in Oslo. Silje, Sigrid, Trygve, Eivor, Harald, Hanna, Håkon, Bull, Helle, Markus, Mari, Brita, Luca, Eirik El Gaucho and Lise Brunbjørn your friendship has been a source of energy during the past three years. Tox, Nikolai, Humle, Troels and André thank you for still reserving a special place in DK for me even after 10 years. Thank you Mum, Dad, Thomas, David, Rikke, Alma and Otto for always being supportive of my studies (especially Alma) and for all your visits to Norway. Thank you also to everyone in my Norwegian family for all your support and keeping a place for me here in Norway. Finally, I want to thank Celin. You have stood by me in field work; plucking bees out of soup bowls in the middle of the night, installing window traps in two metres of snow and put up with me ranting on about bees 24/7. Most importantly, you have provided a loving home for me to return to late in the evening, I am so grateful for having you by my side. Markus, Oslo, 2016 iv Contents Acknowledgements...............................................................................................................................iii Contents..................................................................................................................................................v Listofpapers........................................................................................................................................vii Summary................................................................................................................................................ix Introduction............................................................................................................................................1 Figure1.IllustrationofdifferingthreatenedstatusforbeesinNorwayandEurope.............3 Table1.ComparisonoftheEuropeanandNorwegianbeefauna............................................4 Objectives...........................................................................................................................................6 Figure2.Illustrationofenvironmentalconditionsinpowerlineclearings.............................6 Mainhypothesesposedinthisthesis:..............................................................................................7 A:Thebeephylogenypredictscommunitylevelnicheoverlapandbiogeography(PaperI)....7 B:Hierarchicallystructuredecologicalfiltersdrivebeediversity(PaperII)...............................7 C:ThediversityofbeesdependonnonͲtrophicinteractionswithnonͲbeetaxa(PaperIII).....7 D:Effectsofhabitatmanagementonbeediversitydependonecologicalfilters(PaperIV).....7 E:Bothstochasticanddeterministicprocessesdrivecommunityassembly(PaperV)..............7 Materialsandmethods..........................................................................................................................9 Studysystem(PaperI).......................................................................................................................9 Datasetandexplanatoryvariables................................................................................................9 Statisticalanalyses.......................................................................................................................10 Table2.OverviewofhypothesesandstatisticaltestsappliedinPaperI.............................10 Studysystem(Papers II-V).............................................................................................................11 Figure3.ThelocationofstudysitesincludedinPapersII-V.................................................12 FunctionaltraitsincludedinPapers II-V........................................................................................13 Table3.ListofthefunctionaltraitsappliedinpapersII-Vandtheirdescription................13 MethodsspecifictoPaperII...........................................................................................................14
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