Introduction to the biodiversity of fungi

Introduction to the Biodiversity of Fungi Mariusz Tadych Department of Plant Biology and Pathology Rutgers, the State University of New Jersey ICBG Central Asian Program What is biodiversity? Biological diversity or biodiversity means the variability among living organisms from all sources including, inter alia, terrestrial, marine and other aquatic ecosystems and the ecological complexes of which they are part; this includes diversity within species and between species, and the diversity of ecosystems Convention on Biological Diversity, United Nations Conference on Environment and Development, Rio de Janeiro, 1992 Basic terms • Fungus, pl. fungi, (Lat. fungus, original sfungus from Gr. sphongis - a sponge) Fungi are a group of organisms united by their mode of nutrition • Mycology - the study of fungi; mycetology; (Gr. mykes = mushroom + logos = discourse) • Mycologist - one who studies fungi or mycology Phylogenetic position of organisms studied by mycologists* Organisms Fungi Microsporidia Metazoa Kingdom Fungi ?Microsporidia Animalia Protozoa Domain Dictyosteliidae Myxogastridae Lobosa Angiospermae Chlorophyceae Rhodophyta Glaucophyta Plantae Eukaryota Oomycetes Phaeophyceae Ciliophora Apicomplexa Knietoplastida Euglenoidae Straminipila Acrasidae Vahlkampfiidae Protozoa * - Bold text. The tree is unrooted; based on Baldauf et al., 2000 Selected distinguishing characteristics of three kingdoms Character Nutrition Straminipila Autotrophic (photosynthetic and absorptive) Often cellulose; chitin and β-glucans absent Tubular Fungi Heterotrophic (absorptive/osmotrophic) Protozoa Heterotrophic (phagotrophic) or autotrophic (photosynthetic) Absent when trophic; various when present Tubular Cell wall Chitin and β-glucans Mitochondrial cristae Flagellar mastigonemes Flattened Tubular Absent Not tubular Fungi (overview) • Structure – eukaryotic – pleomorphic - dimorphic (Penicillium marneffei, Histoplasma sp.) - polymorphic (Puccinia graminis) Spermogonium Aecia Uredia Telium – Chitin and β-glucans (wall) • • Reproduction – sexual (teleomorph, meiosporic fungi) – asexual (somatic; anamorph, mitosporic fungi) Usually filamentous bodies enclosed by cell walls – hyphae Fungi (overview) Nonmotile (except those in Chytridiomycota) Heterotrophic (without chlorophyll) Temperature (range from 10 to 50°C) pH (range from 3 to 10) Zoospore Habitats preferred: – dark, moist habitats containing organic material • Ecological types of fungi • • • • • Members of the Kingdom Fungi Basidiomycota (rusts, smuts, mushrooms, etc.) Ascomycota (sac fungi, yeasts, Penicillium, etc.) Glomeromycota (arbuscular mycorrhizal fungi) Zygomycota (saprophytes, ectomycorrhizal fungi) Fungi Chytridiomycota (Blastocladiales) Chytridiomycota Chytridiomycota (Monoblepharidales) (Chytridiales, Neocallimastigales Spizellomycetales, Basidiobolus) (zoosporic fungi) Microsporidia (outgroup) Chytridiomycota Basidiomycota Ascomycota Glomeromycota Zygomycota Spizellomycetales Neocallimastigales Chytridiales (outgroup) Chytridiomycota Basidiobolus Monoblepharidales Blastocladiales Synchytrium endobioticum Zygomycota Harpellales Kickxellales Zoopogales Endogonales Rhizopus stolonifer Zygomycota Mucorales Entomophthorales Dimargalitales Ecrinales Asellariales Endogone maritima Chytridiomycota (outgroup) Glomeromycota Glomerales Diversisporales Archeosporales Paraglomerales S. persica Glomus multiforum Glomeromycota Ascomycota, Basidiomycota (outgroup) Acaulospora bireticulata Glomus clarum Glomus deserticola Glomus multiforum Entrophospora baltica Ascomycota Pezizomycotina Saccharomycotina Taphrinomycotina Ascomycota Basidiomycota (outgroup) Podosphaera aucuparia Rhytisma acerinum Sclerotina sp. Taphrina deformans Venturia inaequalis Basidiomycota Hymenomycetes Ustilaginomycetes Uredinomycetes Basidiomycota Ascomycota (outgroup) Tilletia laevis Ustilago trichophora Rust on Bellis perennis Phragmidium tuberculatum Lycoperdon sp. Boletus sp. Biogeography of fungi • Biogeography is a study of the geographical distribution of living organisms (both past and present), their habitats and the reasons for those distributions; biogeography of fungi is called mycogeography • Mycogeographical analysis requires: 1. Understanding of the past – – – – – – Geological changes Climatological changes Biological changes Fossil evidence Trading patterns Human migration 2. Knowledge of species – – – – Distinguishing macro- and microscopic features Current distribution Preferred habitats and climatic conditions Plants or animals associated with fungi Conditions governing the distribution of species Different fungi have different habitats, substrates or hosts requirements • Factors influencing occurrence of fungal species – organic substrate/soil type – moisture/rainfall pattern – temperature – plant community composition – host or partner present Distribution of fungi Fungi because of their osmotrophic mode of nutrition are able to utilize almost an unlimited diversity of nutritional microniches Podaxis pistllaris – widespread fungus in subtropical dry areas of the world Anthracobia sp. – phoenicoid fungi (after the Phoenix - a mythical bird that arose afresh from a fire every 500 years), fungi growing among the ashes of former fires. Species are found only in burnt ground (obligately inhabiting burnt areas) Trichomaris sp. – found on the carapace of crabs Ascobolus sp. – coprophilous fungi, fungi that inhabit or are associate with the dung of animals, including soil contaminated with dung Xeromyces sp. – osmophilic fungi, growing under conditions of high osmotic pressure Neocallimastix frontalis – anaerobic, saprobic fungi occurring in the guts of herbivores (rumen inhabiting chytrids) Knowledge of distribution and abundance of the known fungal species is poor Important questions that need to be answered about fungi • A baseline needs to be developed to measure changes and abundance of species at particular sites in response to natural or human-induced environmental perturbations (e.g., global warming, air, water, and soil pollution as well as forest fragmentation) Because of destruction of ecosystems numerous fungal species will never be described • We need to understand fungal: • diversity • role in nature • potential utility Number of fungi • Fungi are one of the most diverse groups on Earth 1500000 1000000 500000 0 Described 80000 Estimated 1 1500000 At the current rate of description it will take more than 800 years before all taxa are named 1) Hawksworth, 1991 Where are the undescribed fungi? • fungi in tropical forests – – – – – – – – – – fungi of little-studied host plants fungi in special ecological niches mycorrhizal fungi associated with leguminous tropical tree wood decay corticioid and polyporoid fungi endophytes hypogeous fungi lichenicolous fungi fungi associated with insects poorly studied plants nonpathogenic fungi • fungi in unexplored habitats • lost or hidden species – approximately 20 000 already collected fungal species worldwide are awaiting formal description2 2) Hawksworth and Rossman, 1997 Economic value of fungi • Medicinal use: – antibiotics such as penicillin (Penicillium chrysogenum) – immunosuppressants like cyclosporine (Tolypocladium nivenum) – cholesterol-reducing lovastatin, (Monascus ruber, Aspergillus terreus) – antifungal drug/agents like griseofulvin (Penicillium griseofulvum) – Oriental herbal medicines (Cordyceps sinensis) • Recycling organic matter (plant debris) by saprophytic fungi • Mycorrhizal fungi: – associated with roots of 90% of all vascular plants such as arbuscular mycorrhizae in crops (Glomus sp., Gigaspora sp.), and ectomycorrhizae in most woody plants (Lactarius sp., Laccaria sp.) • Edible fungi: – edible mushrooms, cultivated or wild (Boletus edulis, Morchella sp., Tuber sp., Lentinula edodes, Agaricus bisporus) vesicles arbuscule Glomus aggregatum Glomus sp. Glomus sp. Boletus sp. Saprophytic fungi Economic value of fungi (cont.) • Production of food in agriculture and the food-processing industry – Candida krusei, Geotrichum sp., Penicillium camemberti, Saccharomyces cerevisiae • Use in biological control of insects, nematodes, pathogenic fungi, and weeds A. quisqualis – Ampelomyces quisqualis, Cordyceps sinensis, Nematophthora gynophila, Verticillium chlamydosporium, Phlebia gigantea, Trichoderma sp., Fusarium sp. • Use in ecology as indicators of vegetational or atmospherical changes – Lichens (association of a fungus and an alga) Eudarluca caricis • Source of natural dyes – Macromycetes (Boletus spp., Cortinarius spp., Hygrocybe spp.) – Lichens (Roccella spp. Ochrolechia tartarea, Parmelia saxatilis) • Source of commercially important enzymes and natural products – Trichoderma reesei, Aspergillus sp., Mucor sp., Rhizopus sp. Economic value of fungi (cont.) • Cause of human and animal mycoses • Fungal spores causing allergic responses – Alternaria sp., Aspergillus sp., Candida albicans, Cladosporium sp., Penicillium sp. Cladosporium sp. – Candida albicans, Histoplasma capsulatum, Rhizopus arrhizus, Microsporum canis, Penicillium marneffei. • Plant pathogenic fungi – Phytophthora sp., Puccinia sp., Fusarium sp. (about 80% of plant diseases are caused by fungi) • Fungi causing food spoilage • Fungi producing toxic substances (mycotoxins) Microsphaera sp. – Aspergillus sp., Fusarium sp., Penicillium sp., Rhizopus sp. – Ochratoxins – Aspergillus ochraceus, Penicillium viridicatum (on cereal grains) – Aflatoxins – Aspergillus flavus, A. parasiticus (on various nuts and grains) Reasons for conservation of fungi Biodiversity of fungi throughout the world needs to be conserved • Ecological roles of fungi - vital importance in ecosystem maintenance - saprophytic fungi as natural recyclers - mycorrhizal fungi as important symbionts with plants • Effect of fungi on humans and human-related activities 650 Macrofungi were Reported as Extinct / Missing or Threatened with Extinction in European Red Data Lists3 3) Lizon, 1993 Fungal Conservation Groups • International Union for Conservation of Nature and Natural Resources (IUCN) • International Mycological Association • European Council for the Conservation of Fungi • European Mycological Association • Mycological Society of America • British Mycological Society • Forest Mycology Team • The Endangered Species Unit (ESU) of Australian National Parks and Wildlife Service Legislation for Fungal Protection • Red Data Lists of Fungi (26 European countries) • List of Fungi Protected by Law (Poland) • Projects involving the production of databases on fungal distribution (Canada, USA) • The Priority Flora List of the Department of Conservation and Land Management (Australia) • New Zealand Threat Classification System Lists • Committee for Fungi of the IUCN-Species Survival Commission The diversity of fungi is immense, and their importance, variety, and numbers mandate their inclusion in conservation and biodiversity projects Photos, from left top to right bottom: (1) Robert Puschendorf, Anthony G. Miller, John E. Randall, Jack Jeffrey Photography, Dai G. Herbert, Troy Inman, (2) Rebecca Cairns-Wicks, Tony Palliser, Andrew G. Duthie, John S. Donaldson, Suzanna León-Yánez, Christoph Scheidegger, (3) Anna Lushchekina, Marlon Machado, Malcolm Pym Photography, Shedd Aquarium (photo by Edward G. Lines, Jr.), Farah Ishtiaq, Ross Alford, (4) Bill Konstant, Wendy Strahm, Sun International Resorts, Inc., Craig Hilton-Taylor, Michael Franzen, Tony Palliser, Dada Gottelii