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Diversity of Aphyllophoroid Fungi from Taxonomical and Biogeographical Perspectives

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University of Kassel Faculty 10 Mathematics and Natural Sciences Department of Ecology DISSERTATION for obtaining an academic degree “Doktor der Naturwissenschaften (Dr. rer. nat.)” Diversity of aphyllophoroid fungi from taxonomical and biogeographical perspectives by Oleksandr Ordynets Submitted on 07 June 2018 Defended on 23 October 2018 Kassel, 2018 Author’s address Department of Ecology Faculty 10 Mathematics and Natural Sciences University of Kassel Heinrich-Plett-Strasse 40 34132 Kassel Germany Email: [email protected] Supervisor Prof. Dr. Ewald Langer Department of Ecology, University of Kassel, Kassel, Germany Reviewers Prof. Dr. Ewald Langer Department of Ecology, University of Kassel, Kassel, Germany Dr. Christian Printzen Botany and Molecular Evolution Research, Senckenberg Research Institute and Museum of Natural History, Frankfurt am Main, Germany Disputation commission members Prof. Dr. Ewald Langer Department of Ecology, University of Kassel, Kassel, Germany Dr. Christian Printzen Botany and Molecular Evolution Research, Senckenberg Research Institute and Museum of Natural History, Frankfurt am Main, Germany Prof. Dr. Georg Mayer Department of Zoology, University of Kassel, Kassel, Germany Prof. Dr. Raffael Schaffrath Department of Microbiology, University of Kassel, Kassel, Germany TABLE OF CONTENTS ABSTRACT .......................................................................................................................... 3 ABSTRACT IN GERMAN / ZUSAMMENFASSUNG ...................................................... 5 LIST OF ORIGINAL PUBLICATIONS .............................................................................. 7 AUTHORS’ CONTRIBUTION TO PUBLICATIONS ........................................................ 8 INTRODUCTION ............................................................................................................... 10 Fungal perspective on biodiversity knowledge ............................................................... 10 Dimensions of fungal diversity........................................................................................ 11 Is every fungus found everywhere? ................................................................................. 12 Why study aphyllophoroid fungi? ................................................................................... 12 AIMS ................................................................................................................................... 16 MATERIALS AND METHODS ........................................................................................ 17 Data sampling .................................................................................................................. 17 Specimens .................................................................................................................... 17 Sequences .................................................................................................................... 17 Observations ................................................................................................................ 17 Public occurrence data ................................................................................................. 17 Environmental data ...................................................................................................... 18 Morphological study ........................................................................................................ 18 Molecular laboratory work .............................................................................................. 19 DNA sequence-based analyses ........................................................................................ 19 Ecological analyses.......................................................................................................... 20 Gamma diversity and local species richness ............................................................... 20 Beta diversity ............................................................................................................... 20 Multivariate methods ................................................................................................... 21 Data and protocols management...................................................................................... 21 RESULTS AND DISCUSSION .......................................................................................... 23 New species occurrence data (IIV) ............................................................................... 23 Two new Trechispora species with peculiar morphological and ecological traits (II) ... 24 Subulicystidium: twice more species than known before (III) ........................................ 24 Subulicystidium brachysporum and S. meridense are polyphyletic (III) ......................... 26 Nuclear ribosomal DNA sequence variation in Subulicystidium upon a comprehensive taxon sampling (III) ......................................................................................................... 27 Geographical distribution of the newly introduced and revised taxa (II, III) .................. 27 Comprehensive account of the known aphyllophoroid fungal diversity in Europe (IV) 28 Spatial variation of species richness of aphyllophoroid fungi in Europe (IV) ................ 29 1 Spatial variation of community composition of European aphyllophoroid fungi in Europe (IV) ..................................................................................................................... 29 Different environmental drivers do matter for different diversity components (IV) ...... 30 CONCLUSIONS................................................................................................................. 32 ACKNOWLEDGEMENTS ................................................................................................ 34 REFERENCES ................................................................................................................... 36 ERKLÄRUNG GEMÄSS § 8 ABS. 1 SATZ 2 LIT. D AB-PromO .................................. 49 PUBLICATIONS ................................................................................................................ 51 2 ABSTRACT Comprehensive knowledge of biodiversity is a prerequisite for its long-term conservation and sustainable use. Fungi play crucial roles in ecosystems and are among the species-richest organism groups on Earth. However, all aspects of their diversity remain underexplored. In this study, we aimed to fill the gaps in occurrence data and taxonomy as well as in understanding spatial diversity patterns of aphyllophoroid fungi (non-gilled macroscopic Basidiomycota). We digitized and made openly accessible 4,041 records of aphyllophoroid fungi from Central and Eastern Europe and several tropical areas. Numerous specimen records were associated with newly generated nuclear ribosomal DNA sequences (74 of ITS regions and 59 of 28S region) and numerous observations also with photographs depicting diagnostic features of fungal fruitbodies. We described 13 new species of aphyllophoroid fungi based on morphological examination combined with DNA barcoding and phylogenetic Bayesian and Maximum likelihood analyses using ITS and 28S DNA sequences. Two new species belong to the genus Trechispora P. Karst. and eleven to the genus Subulicystidium Parmasto. In the latter, species-level variation in basidiospore size and shape was re-evaluated based on systematic measurements of 2,840 spores from 67 sequenced specimens. An updated identification key to all known species of Subulicystidium was provided. Using DNA-based evidence, in the studied lineage (Trechisporales K.H.Larss.) we showed for the first time a possibility of a transoceanic distribution. Based on a dataset consisting of 14,030 fruitbody occurrences of 1,491 aphyllophoroid fungal species from 39 European areas, we showed that importance of biogeographical regions in determining European aphyllophoroid fungal communities varies for different diversity components. Species richness and nestedness were best explained by European biogeographical regions, whereas overall beta-diversity and species turnover were driven mostly by variation in climate, and nestedness mostly by tree species occupancy. Beta diversity patterns of aphyllophoroid fungi did not differ between southern and northern Europe. Therefore, at the continental scale, aphyllophoroid fungi are less shaped by historical legacies than vascular plant and animal communities. Keywords: Agaricomycetes, beta diversity, biodiversity informatics, biometry, biogeography, distribution, Hydnodontaceae, nestedness, new species, species delimitation, species richness, species turnover, taxon occurrence, taxonomy 3 ABSTRACT IN GERMAN / ZUSAMMENFASSUNG Umfassendes Wissen über die Biodiversität ist eine Voraussetzung für ihre langfristige Erhaltung und nachhaltige Nutzung. Pilze spielen eine wichtige Rolle in Ökosystemen und gehören zu den artenreichsten Organismengruppen auf der Erde. Jedoch sind noch nicht alle Aspekte ihrer Vielfalt vollständig erforscht. In dieser Studie haben wir uns zum Ziel gesetzt, die Lücken in den Verbreitungsdaten und der Taxonomie sowie im Verständnis der räumlichen Diversitätsmuster von aphyllophoroiden Pilzen (makroskopische Nichtblätterpilze der Basidiomycota) zu füllen. Wir haben 4,041 Datensätze von aphyllophoroiden Pilzen aus Mittel- und Osteuropa und verschiedenen tropischen Gebieten digitalisiert und veröffentlicht.
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