Ectomycorrhizal fungi in Amazonian tropical forests in Colombia Aída Marcela Vasco-Palacios Cover design: Clara Mendoza. Layout Sayonara Plata, Fotos: Aída M. Vasco-Palacios. PhD thesis Utrecht University, Utrecht, NL (2016) [email protected] With references and summaries in Dutch and Spanish The research described in this thesis was conducted at the CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands, and The Laboratory of Taxonomy and Ecology of Fungi, Antioquia University, Medellín, Colombia. Copyright © 2016 by A. Vasco-Palacios, Fundación Biodiversa Colombia. All rights reserved. Printed by Panamericana Formas e Impresos S.A., Colombia ISBN 978-958-46-9373-0 This research was supported by the Netherlands Scholarships Programs (NFP-NUFFIC 2010- 2015), the Faculty for the Future - Schlumberger Foundation (FFTF, Grant to Aida Vasco 2011- 2013), the International Science Foundation (IFS Grant D/5052-1, 401 2011-2-13f), Utrecht Ectomycorrhizal fungi in Amazonian tropical forests in Colombia Ectomycorrhiza schimmels in tropische Amazone bossen in Colombia (met een samenvatting in het Nederlands) Proefschrift ter verkrijging van de graad van doctor aan de Universiteit Utrecht op gezag van de rector magnificus, prof. dr. G.J. van der Zwaan, ingevolge het besluit van het college voor promoties in het openbaar te verdedigen op maandag 26 september 2016 des middags te 6.00 uur door Aída Marcela Vasco-Palacios geboren op 13 januari te Bogotá, Colombia Promotoren: Prof. dr. H.A.B. Wösten Prof. dr. T. Boekhout To Gloria Galeano that will always be present in my heart and to my lovely family and friends Contents CHAPTER 1 General Introduction 9 SECTION 1 DIVERSITY CHAPTER 2 Diversity of ectomycorrhizal fungi from white-sand 21 forests in the Colombian Amazonia CHAPTER 3 Ectomycorrhizal fungal communities associated 43 with the dipterocarp Pseudomonotes tropenbosii in Colombian Amazonia CHAPTER 4 Forest type and soil chemistry are important drivers 73 that structure the highly diverse fungal communities in lowland tropical forests in Colombia SECTION 2 TAXONOMIC NOVELTIES CHAPTER 5 Austroboletus amazonicus sp. nov. and Fistulinella 103 campinaranae var. scrobiculata, two common boletes from a forest dominated by the dipterocarp Pseudomonotes tropenbosii in Colombian Amazonia CHAPTER 6 Synopsis of Coltricia and Coltriciella in the 121 Neotropics, new species from Amazonian ecosystems in Colombia CHAPTER 7 Sarcodon in the Neotropics: four new species from 141 Colombia and a key to selected species CHAPTER 8 General Discussion and Summary 161 References 169 Samenvatting 189 Resumen 192 Acknowledgements 198 Curriculum vitae 200 List of Publications 201 CHAPTER 1 General Introduction - Chapter 1 - GENERAL INTRODUCTION The Amazonian region encompasses an area of about 8 million km2 that is located in the northern part of South America. It is home to the largest forests of the planet representing nearly 50 % of the global tropical rainforest area (Hoorn & Wesselingh 2010). The area includes two main ecoregions, the Amazon basin that holds the drainage basin for the Amazon River and its many tributaries, and the Guiana Shield (Eva et al. 2005). The Amazon basin can be found in Bolivia, Brazil, Colombia, Ecuador, Guyana, Peru, Surinam, and Venezuela, while the Guiana Shield is located in Guyana, Surinam, French Guyana, and some parts of Venezuela, Colombia, and Brazil (Hoorn & Wesselingh 2010). The Amazon region holds a great diversity of terrestrial and aquatic organisms and plays a significant role in regulating the global climate, the hydrological cycle, and the carbon and nitrogen cycles (Hoorn & Wesselingh 2010; Phillips et al. 2009). The actual deforestation rates in the area are extremely high, ranging from 25,000 to 50,000 km2 per year (Fearnside & Laurance 2003; Malhi et al. 2008; Shukla et al. 1990). This may result in the total disappearance of this tropical rainforest in a relatively short time (Malhi et al. 2008; Shukla et al. 1990). The overview of the biodiversity of Amazonia remains to be completed and biological processes underlying this diversity are poorly documented. While 90-95 % of plants, mammals, and bird species have been described, the vast majority of the other organisms, such as bacteria, insects, and fungi are virtually unknown (Behling et al. 2010). Understanding Biodiversity in Amazonian Forests Processes such as long-term landscape evolution and climate change have been identified as important drivers of speciation and extinction, thus shaping the current patterns of biodiversity in Amazonian forests (Hoorn et al. 2010). Four important historical events have strongly contributed to the current pattern of diversity in the Amazonia region: 1. a long period of isolation, 2. the Andean uplift, 3. the closing of the Panama isthmus, and 4. changes in the global climate (Burnham & Graham 1999; Hoorn et al. 2010). The different geology from East to West in combination with differences in rainfall resulted in a gradient of soil fertility, which also contributed to the pattern of biodiversity (Duque et al. 2002; Quesada et al. 2009; Sombroek 2000; ter Steege et al. 2006, 2010). Growth of plant species is restricted by a subset of environmental conditions that determines plant diversity (Fine et al. 2006). In particular, soil characteristics, such as water holding capacity, drainage, degree of flooding, soil structure, and nutrient contents affect plant growth, mortality, and levels of diversity (Duivenvoorden & Lips 1993; Ferry et al. 2010; Quesada et al. 2009). Northwestern Amazonia is typified by its relatively rich soils and has an exceptional high richness of plant diversity per unit area (Duivenvoorden & Duque 2010; Gentry 1988; ter Steege et al. 2006, 2010). In contrast, soils of Northeast Amazonia are relatively poor and this area presents a high dominance of Fabaceae (Henkel et al. 2012; Jimenez et al. 2009; McGuire 2007; ter Steege 2010). At a smaller scale, diversity and composition of plants are driven by factors such as local disturbance, predation, and soil nutrients (Augspurger 1983; Connell 1971, 1978; Duivenvoorden & Duque 2010; Fine et al. 2004; Janzen 1970; Stropp et al. 2009). Recent studies also 12
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