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PHD THESIS Heterobasidion Bref. and Armillaria (Fr.) Staude pathosystems in the Basque Country: Identification, ecology and control. Nebai Mesanza Iturricha PHD THESIS 2017 PHD THESIS Heterobasidion Bref. and Armillaria (Fr.) Staude pathosystems in the Basque Country: Identification, ecology and control. Presented by Nebai Mesanza Iturricha 2017 Under the supervision of Dr. Eugenia Iturritxa and Dr. Cheryl L. Patten Tutor: Dr. Maite Lacuesta (c)2017 NEBAI MESANZA ITURRICHA Front page: Forest, by Araiz Mesanza Iturricha Acknowledgements This work was carried out at Neiker- Tecnalia (Basque Institute for Agricultural Research and Development) and at the Department of Biology at the University of New Brunswick, and it was funded by the Projects RTA: 2013-00048-C03-03 INIA, Healthy Forest: LIFE14 ENV/ES/000179, the Basque Government through a grant from the University and Research Department of the Basque Government, a grant from the New Brunswick Innovation Foundation, and a grant from the European Union 7 th Framework Programme (Marie Curie Action). I am especially grateful to my supervisors Dr. Eugenia Iturritxa and Dr. Cheryl L. Patten for their constant support during this process and for giving me the opportunity to get involved in this project. I would also like to thank Ander Isasmendi and Patxi Sáenz de Urturi for their skillful assistance during the sampling process, and in general to all the people that have shared their knowledge and time with me. My deepest gratitude to Carmen and Vitor, you have been my shelter since I know you. Araiz, you are the best illustrator ever. Thank you very much to you and Erling for the Mediterranean air and the wild boars. I also appreciate the good moments and delicious meals with all my family. I am very thankful to all my friends for being an oasis for my brain. Maialen, I know our situation was not the best but I am very glad we were together in this. Artyom, thank you very much for the music and letting me sing. Urko, let´s go to the swimming pool! Arrate, thank you very much for your perspective. Zurine, I really enjoy our conversations late at night and all the painting. Thank you very much to Heather, Danna, Caroline, Shane, Victoria, TeeJ and Charlotte for making my first Christmas in Canada warm. Finally, I warmly thank Thomas for all his input, fresh ideas, and help; and for all the amazing adventures. List of original articles This thesis is based on the following articles: Mesanza, N., Patten, C. L, Iturritxa, E., 2017. Distribution and characterization of Armillaria complex in Atlantic forest ecosystems of Spain. Forests (Submitted). Mesanza, N., Iturritxa, E., Patten, C.L., 2016. Native rhizobacteria as biocontrol agents of Heterobasidion annosum s.s. and Armillaria mellea infection of Pinus radiata. Biol. Control 101, 8-16. Mesanza, N., Iturritxa, E., 2012. Root and butt rot caused by Heterobasidion annosum in Atlantic coniferous ecosystems of Spain. For. Pathol. 42, 514–520. Summary of contents Summary of contents CHAPTER 1: Introduction and objectives................................................................................ 1 1.1. Armillaria species complex ............................................................................................ 2 1.1.1. Taxonomy ............................................................................................................ 2 1.1.2. Armillaria species and their distribution ............................................................. 3 1.1.3. Host susceptibility ............................................................................................... 5 1.1.4. Infection and life cycle......................................................................................... 6 1.1.5. Symptoms and signs of infection ........................................................................ 8 1.1.6. Damage................................................................................................................ 9 1.2. Heterobasidion species complex .................................................................................... 10 1.2.1. Taxonomy .......................................................................................................... 10 1.2.2. Heterobasidion species and their distribution..................................................... 10 1.2.3. Host susceptibility ............................................................................................. 12 1.2.4. Infection and life cycle....................................................................................... 13 1.2.5. Symptoms and signs of infection ...................................................................... 15 1.2.6. Damage.............................................................................................................. 16 1.3. Area of study ............................................................................................................... 16 1.3.1. Localization and geomorphology of the Basque Country ................................ 16 1.3.2. General climate.................................................................................................. 18 1.3.3. Native forest and tree plantations ..................................................................... 19 1.4. Objectives .................................................................................................................... 22 CHAPTER 2: Distribution and characterization of Heterobasidion and Armillaria complexes in the Basque Country........................................................................................... 23 2.1. Introduction................................................................................................................. 23 2.1.1. Objectives .......................................................................................................... 25 2.2. Material and methods ................................................................................................. 26 2.2.1. Collection of fungal material............................................................................. 26 2.2.2. Identification of spatial distribution patterns ................................................... 26 2.2.3. Description of fungal ecosystems...................................................................... 27 2.2.4. Fungal species identification ............................................................................. 29 i Summary of contents 2.3. Results ......................................................................................................................... 31 2.3.1. Distribution of fungal species ........................................................................... 31 2.3.2. Spatial distribution patterns.............................................................................. 33 2.3.3. Ecosystem characteristics in which fungal species were detected.................... 35 2.3.4. Fungal species identification ............................................................................. 40 2.4. Discussion.................................................................................................................... 42 2.4.1. Conclusions ....................................................................................................... 44 CHAPTER 3: Population diversity of Heterobasidion annosum s.s. and Armillaria complex in selected stands of the Basque Country............................................................................... 45 3.1. Introduction................................................................................................................. 45 3.1.1. Objectives .......................................................................................................... 46 3.2. Materials and Methods................................................................................................ 47 3.2.1. Sample collection............................................................................................... 47 3.2.2. Identification and fungal diversity analysis ..................................................... 48 3.2.3. Host susceptibility ............................................................................................. 50 3.3. Results ......................................................................................................................... 51 3.3.1. Analysis of Armillaria population diversity ...................................................... 51 3.3.2. Analysis of Heterobasidion population diversity................................................ 54 3.3.3. Host susceptibility ............................................................................................. 57 3.4. Discussion.................................................................................................................... 59 3.4.1. Conclusions ....................................................................................................... 60 CHAPTER 4: Native rhizobacteria as biocontrol agents of Heterobasidion annosum s.s. and Armillaria mellea infection of Pinus radiata .................................................................. 61 4.1. Introduction................................................................................................................. 61 4.1.1. Objectives .......................................................................................................... 62 4.2. Materials and methods ...............................................................................................
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