Table of contents Chapter General introduction 7 Chapter Ancestral predisposition toward a domesticated 19 lifestyle in the termite-cultivated fungus Termitomyces Chapter Genetic population structure of the agaric 45 Blastosporella zonata reveals cryptic species and different roles for sexual and asexual spores in dispersal Chapter Four new genera and six new species of 59 Lyophylloid agarics (Agaricales, Basidiomycota) from three different continents Chapter Species and speciation in the termite-cultivated 79 fungus Termitomyces Chapter High diversity and low host-specificity of 99 Termitomyces symbionts cultivated by Microtermes spp. indicate frequent symbiont exchange Chapter Termitomyces cryptogamus sp. nov. associated 113 with Macrotermes natalensis in Africa Chapter General discussion 129 References 156 Summary 172 Acknowledgements 176 PE&RC training and education statement 180
General introduction
SymbiosisThe word symbiosis is derived from the Greek words 'syn' and 'bios' meaning "living together". In biology it is defined as a close, long-term interaction between two different organisms. Such an interaction can be commensal when one party benefits with a neutral effect on the other party (so no benefit or harm), parasitic, or mutualistic. An example of a commensal interaction are remora fish which attach themselves to sharks and eat food leftovers while the shark does not gain any benefits nor negative effects. Parasitism is usually associated with bacterial, fungal, or viral diseases and ectoparasites live and feed on the host.Examples of this are fleas, lice, ticks or, in the case of plants, root parasites such as broomrapes (Orobanchaceae). In a mutualistic symbiosis both parties receive a benefit. An important example of mutualistic symbiosis is mycorrhiza, a mutualistic symbiosis between plant roots and mycorrhizal fungi. The plant provides the fungus with sugars in exchange for minerals. Another similar example are nitrogen fixing Rhizobium bacteria in the root nodules of legumes, which fix nitrogen from the air in exchange for sugars.The main topic of my thesis is insect fungiculture; a mutualistic symbiosis where insects provide a protective growth environment for fungi, in exchange for nutritional benefits. Mutualistic symbioses can be directly linked to major evolutionary innovations (Szathmáry and Maynard-Smith 1995). Lynn Margulis proposed the idea that symbiosis in some cases, over a long time-span could lead to symbiogenesis (Sagan 1967). Symbiogenesis, also known as the endosymbiosis theory, is often used to explain how eukaryotes have evolved. Evidence for this are organelles such as the mitochondrion and the chloroplasts in plants which are thought to have evolved from a free-living prokaryotic ancestor.