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Diversity of Microfungi Preserved in European Palaeogene Amber 12/2017 Senja Laakso Practice Approach to Experimental Governance

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Diversity of Microfungi Preserved in European Palaeogene Amber 12/2017 Senja Laakso Practice Approach to Experimental Governance YEB Recent Publications in this Series ELINA KETTUNEN 8/2017 Mantas Survila Immune Responses to Pathogen Infection in Arabidopsis 9/2017 Jessica M. Coyne Genetics of Longitudinal Growth Data 10/2017 Xing Wan Leuconostoc bacteriocins and Their Application in Genome Editing DISSERTATIONES SCHOLA DOCTORALIS SCIENTIAE CIRCUMIECTALIS, 11/2017 Inka Reijonen ALIMENTARIAE, BIOLOGICAE. UNIVERSITATIS HELSINKIENSIS 4/2018 Chemical Bioavailability of Chromium and Vanadium Species in Soil: Risk Assessment of the Use of Steel Industry Slags as Liming Materials Preserved in European Palaeogene Amber of Microfungi Diversity 12/2017 Senja Laakso Practice Approach to Experimental Governance. Experiences from the Intersection of Everyday Life and Local Experimentation 13/2017 Paulina Deptula A Multifaceted Study of Propionibacterium Freudenreichii, the Food-Grade Producer of Active Vitamin B12 ELINA KETTUNEN 14/2017 Taneli Tirkkonen Porcine mycobacteriosis Caused by Mycobacterium avium subspecies hominissuis Diversity of Microfungi Preserved in European 15/2017 Pär Davidsson Oligogalacturonide Signalling in Plant Innate Immunity Palaeogene Amber 16/2017 Kean-Jin Lim Scots Pine (Pinus sylvestris L.) Heartwood Formation and Wounding Stress: A View from the Transcriptome 17/2017 Marja Rantanen Light and Temperature as Developmental Signals in Woodland Strawberry and Red Raspberry 18/2017 Sara Kovanen Molecular Epidemiology of Campylobacter jejuni in the Genomic Era 19/2017 Johanna Muurinen Antibiotic Resistance in Agroecosystems 20/2017 Johanna Laakso Phosphorus in the Sediment of Agricultural Constructed Wetlands 21/2017 Sadegh Mansouri Plant Biomass-Acting Enzymes Produced by the Ascomycete Fungi Penicillium subrubescens and Aspergillus niger and Their Potential in Biotechnological Applications 22/2017 Anna Salomaa Actors’ Roles and Perceptions on the Opportunities to Increase Nature Conservation Effectiveness: a Study of Interaction Between Knowledge and Policy Process 23/2017 Anniina Le Tortorec Bioluminescence of Toxic Dinoflagellates in the Baltic Sea - from Genes to Models 24/2017 Tanja Paasela The Stilbene Biosynthetic Pathway and Its Regulation in Scots Pine 1/2018 Martta Viljanen Adaptation to Environmental Light Conditions in Mysid Shrimps 2/2018 Sebastián Coloma Ecological and Evolutionary Effects of Cyanophages on Experimental FACULTY OF BIOLOGICAL AND ENVIRONMENTAL SCIENCES Plankton Dynamics DOCTORAL PROGRAMME IN PLANT SCIENCES 3/2018 Delfia Isabel Marcenaro Rodriguez UNIVERSITY OF HELSINKI Seedborne Fungi and Viruses in Bean Crops (Phaseolus vulgaris L.) in Nicaragua and Tanzania 4/2018 Helsinki 2018 ISSN 2342-5423 ISBN 978-951-51-4151-4 Faculty of Biological and Environmental Sciences University of Helsinki Helsinki DIVERSITY OF MICROFUNGI PRESERVED IN EUROPEAN PALAEOGENE AMBER Elina Kettunen ACADEMIC DISSERTATION To be presented for public examination, with the permission of the Faculty of Biological and Environmental Sciences of the University of Helsinki, in lecture room 105 (B6), Viikki B building, on 4th of May 2018, at 12 noon. Helsinki 2018 Supervised by: Prof. Jouko Rikkinen Finnish Museum of Natural History, Luomus, Faculty of Biological and Environmental Sciences, University of Helsinki Reviewed by: Prof. Marc-André Selosse, Muséum National d’Histoire Naturelle, Paris, France Dr. Rafael Felipe Castañeda Ruiz Institute of Fundamental Researches in Tropical Agriculture "Alejandro de Humboldt", Ministerio de la Agricultura, Cuba Follow-up group: Dr. Johannes Enroth Faculty of Biological and Environmental Sciences, University of Helsinki Dr. Tuula Niskanen Royal Botanical Gardens, Kew, London, UK Examined by: Dr. Matthias Gube Georg August University Göttingen, Germany Custos: Prof. Jaakko Hyvönen Finnish Museum of Natural History, Luomus, Faculty of Biological and Environmental Sciences, University of Helsinki Doctoral Programme in Plant Sciences Dissertationes Schola Doctoralis Scientiae Circumiectalis, Alimentariae, Biologicae 4/2018 ISSN 2342-5423 (Print) ISSN 2342-5431 (Online) ISBN 978-951-51-4151-4 (pbk.) ISBN 978-951-51-4152-1 (PDF) Hansaprint Oy Turenki 2018 ABSTRACT Fungi are one of the most diverse groups of organisms, but their fossil record is scarce compared with that of plants and animals. However, many fossils of microfungi have survived as inclusions in amber, which is fossilized resin produced by ancient trees millions of years ago. Some of these fossils were already discovered and described during the 19th century. There are important sources of Palaeogene amber in Europe: the Baltic and Bitterfeld deposits. Baltic amber is about 43–25 Ma old (Eocene), whereas Bitterfeld amber is slightly younger (approx. 24 Ma, Oligocene). The aim of this thesis was to increase our knowledge of the microfungi preserved in the Baltic and Bitterfeld ambers. The material studied included both historic collections and previously unstudied amber specimens. The approach led to several advances in the field of palaeomycology. The systematic affinities of the microfungi described by Robert Caspary and Richard Klebs over a century ago were reassessed. None of these historical specimens belong to the extant fungal genera they were originally assigned to. Amended descriptions were provided for the historical specimens, and several new types of fungi were described from novel amber specimens. These included the first fossils of lichen-associated filamentous fungi. The results demonstrate that relatively few fossil microfungi in amber can be identified accurately enough to be used as minimum age constraints in dating phylogenetic trees of different fungal lineages. All the fossil fungi studied grew either on or in the immediate vicinity of resin-producing trees, which made them likely candidates for preservation in amber. 3 ACKNOWLEDGEMENTS Jouko Rikkinen introduced the world of amber to me when I was his Master's student. Jouko, thank you for all the encouragement, inspiration and support you have given to me as the supervisor of this PhD thesis. The work on amber linked me with Alexander Schmidt and his research group in the University of Göttingen. Alexander, thank you for all your help and support, this thesis would not have been possible without you. I would also like to thank other amber researchers in Göttingen, big thanks to Leyla, Eva and Christina, it was great to do research with you. Thanks to my other co-authors for their co-operation. I am very grateful to all private collectors, whose amber specimens were used in this study. Many thanks to Eija, Maarit, Åsa and Risto for their help and support. To my PhD follow-up group, Johannes Enroth and Tuula Niskanen, thank you for your advice and guidance. Thanks to Hanna, Seppo, Panu, Lotta, Kaisa and Jenna for good discussions and support, especially in the early phases of my thesis research. Marc-André Selosse and Rafael Felipe Castañeda Ruiz reviewed the thesis, warm thanks for their helpful comments and suggestions. Lastly I would like to thank my family and all my friends for their support during these years, thank you for your encouragement and advice. This work was funded by the Jenny and Antti Wihuri Foundation and the Otto A. Malm Foundation. 4 CONTENTS Abstract .............................................................................................................. 3 Acknowledgements ......................................................................................... 4 Contents ............................................................................................................. 5 List of original publications .......................................................................... 6 Summary ............................................................................................................ 8 1. Introduction ............................................................................................. 8 Amber .......................................................................................................... 8 Baltic and Bitterfeld ambers ........................................................... 10 Study and challenges of amber ....................................................... 12 Fungi and their fossil history ..................................................................... 13 Fungi in amber ................................................................................. 15 Hyphomycetes .................................................................................. 17 2. Aim of the thesis .................................................................................... 18 3. Material and methods .......................................................................... 18 Material ...................................................................................................... 18 Methods..................................................................................................... 20 4. Main results ............................................................................................ 21 5. Discussion and conclusions ............................................................... 22 6. References .............................................................................................. 25 I The enigmatic hyphomycete Torula sensu Caspary revisited II Caspary’s fungi from Baltic amber: historic specimens and new evidence III Lichen-associated fungi from Paleogene amber IV Diversity of lichen-associated filamentous fungi in European Paleogene amber 5 LIST OF ORIGINAL PUBLICATIONS This thesis is
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