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Thesis.Pdf (3.979Mb) FACULTY OF BIOSCIENCES, FISHERIES AND ECONOMICS DEPARTMENT OF ARCTIC AND MARINE BIOLOGY Cyclically outbreaking geometrid moths in sub-arctic mountain birch forest: the organization and impacts of their interactions with animal communities — Ole Petter Laksforsmo Vindstad A dissertation for the degree of Philosophiae Doctor – October 2014 Cyclically outbreaking geometrid moths in sub-arctic mountain birch forest: the organization and impacts of their interactions with animal communities Ole Petter Laksforsmo Vindstad A dissertation for the degree of Philosophiae Doctor University of Tromsø – The arctic university of Norway Faculty of Biosciences, Fisheries and Economics Department of Arctic and Marine Biology Autumn 2014 1 Dedicated to everyone who has helped me along the way 2 Supervisors Professor Rolf Anker Ims1 Senior researcher Jane Uhd Jepsen2 1 Department of Arctic and Marine Biology, University of Tromsø, Tromsø, Norway 2 Norwegian Institute for Nature Research, Fram Centre, Tromsø, Norway Cover photos Front cover – Larvae of Epirrita autumnata feeding on mountain birch during a moth outbreak in northern Norway. Photo: Moritz Klinghardt Study I – Portrait of Agrypon flaveolatum. One of the most important larval parasitoid species in study I. Photo: Ole Petter Laksforsmo Vindstad Study II – Carcass of an Operophtera brumata larva, standing over the cocoon of its killer, the parasitoid group Protapanteles anchisiades/P. immunis/Cotesia salebrosa. Photo: Ole Petter Laksforsmo Vindstad Study III – Larva of the parasitoid group Phobocampe sp./Sinophorus crassifemur emerging from Agriopis aurantiaria host larva. Photo: Tino Schott Study IV – An area of healthy mountain birch forest, representative for the undamaged sampling sites in study IV and V. Photo: Jakob Iglhaut Study V – An area of mountain birch forest that has been heavily damaged by a moth outbreak, representative for the damaged sampling sites in study IV and V. The stems without foliage have all been killed by the outbreak. Photo: Jakob Iglhaut 3 Contents List of original papers ............................................................................................................. 6 Preface ...................................................................................................................................... 7 Abstract .................................................................................................................................... 8 1. Introduction ....................................................................................................................... 10 1.1. Introduction to insect outbreaks and population cycles .............................................................. 10 1.1.1. Insect outbreaks and population cycles as disturbance factors in forest ecosystems ......... 10 1.1.2. General trends in the research on insect outbreaks and population cycles ........................ 10 1.2. The ecological consequences of insect outbreaks and cycles: extensive but understudied ........ 12 1.2.1. Potential consequences of insect outbreaks for forest ecosystems ...................................... 12 1.2.2. Potential consequences of insect population fluctuations for natural enemy communities 14 1.3. The role of natural enemies in insect outbreaks and cycles: the need to consider enemy community organization .................................................................................................................... 15 1.3.1. General trends in the research on OCI-enemy interactions................................................ 15 1.3.2. Diversity-functionality relationships in enemy communities............................................... 15 1.3.3. OCI range expansions and enemy diversity-functionality relationships ............................. 16 1.3.4. Phenological organization of enemy communities .............................................................. 17 1.4. The study system: cyclically outbreaking geometrid moths in North-Fennoscandian mountain birch forest ......................................................................................................................................... 18 1.4.1. Introduction to the Fennoscandian birch-moth system ....................................................... 18 1.4.2. Key topics and results in the research on moth population cycles and outbreaks .............. 20 1.4.3. Ecological consequences of moth outbreaks ....................................................................... 20 1.4.4. Knowledge gap: indirect impacts of moth outbreaks on saproxylic beetles and passerine birds ............................................................................................................................................... 21 1.4.5. Research on the role of trophic interactions in moth population dynamics ........................ 22 1.4.6. Research on parasitoids as drivers of moth population cycles ........................................... 23 1.4.7. Limitations to the research on parasitoids in the Fennoscandian birch-moth system ........ 24 1.4.8. Knowledge gap: the causes and consequences of spatial variation in larval parasitoid guild structure ............................................................................................................................... 24 1.4.9. Knowledge gap: the phenological organization of larval parasitoid guilds ....................... 27 1.4.10. Knowledge gap: larval parasitism of invasive geometrid species .................................... 28 1.5. Specific questions addressed by the PhD project ....................................................................... 29 2. Materials and methods ...................................................................................................... 31 2.1. Study regions .............................................................................................................................. 31 2.2. Study species .............................................................................................................................. 31 2.2.1. Geometrid moths ................................................................................................................. 31 2.2.2. Larval parasitoids ...............................................................................................................32 2.3. Study design ............................................................................................................................... 32 2.3.1. Study I .................................................................................................................................. 32 2.3.2. Study II ................................................................................................................................ 34 2.3.3. Study III ............................................................................................................................... 35 2.3.4. Study IV ............................................................................................................................... 35 2.3.5. Study V................................................................................................................................. 36 4 2.4. Statistical analyses ...................................................................................................................... 36 2.4.1. Analytical philosophy and general considerations ............................................................. 36 2.4.2. Study I .................................................................................................................................. 37 2.4.3. Study II ................................................................................................................................ 37 2.4.4. Study III ............................................................................................................................... 38 2.4.5. Study IV ............................................................................................................................... 38 2.4.6. Study V................................................................................................................................. 39 3. Results and discussion ....................................................................................................... 39 3.1 Study I ......................................................................................................................................... 39 3.1.1. Spatiotemporal dynamics of larval parasitoid guilds ......................................................... 39 3.1.2 Relationships between larval parasitoid guild structure and total larval parasitism rates . 41 3.2 Study II ........................................................................................................................................ 42 3.2.1. Phenological organization of the larval parasitoid guild of O. brumata ............................ 42 3.2.2. Phenological mismatch between larvae and parasitoids at high altitudes? ....................... 46 3.2.3. Methodological implications for studies of larval parasitism ............................................. 46 3.3 Study III ....................................................................................................................................... 47 3.3.1. Accumulation of larval parasitoid species on invasive birch forest geometrids ................. 47 3.3.2. Parasitism rates and
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