Lecture 16 Ascomycota V - Lichenized fungi - Eurotiomycetes - Laboulbeniales Lutzoni et al., 2004, American Journal of Botany Lichen-forming Lichen-forming Sordariomycetes
Nature 21:937-940. 2001 - Nuclear ribosomal DNA phylogeny Euascomycota - Reconstruction of ancestral character-states lichen-forming uncertain Hemiascomycota Archiascomycota Basidiomycota Generalities on Lichens LICHENS = a fungus + a photosymbiont (algae or cyanobacteria) : not a monophyletic group; LICHENS result from SYMBIOSIS between organisms from different Kingdoms. - 15-20,000 recognized lichen species vs.a total of ~30,000 Asco species ==> lichen count for ca. > 40% of of all known Ascomycetes ==> one fifth of all known fungi! - Habitats: primary (eukaryotic) colonizers of harsh / abiotic habitats, including rocks; --- are found in the most extreme environments, e.g., sandstone deserts from hot Namibia to cold Antarctica and up to 7,000 m. high in Mt Everest slopes. Food for thoughts: Heckman et al., Science (2001) : speculate lichens as early colonizers of terrestrial habitats 1.5 byo Kendrick s textbook (2000) statement p.124: it has been suggested that if [early] life were found in another planet, it might well be lichen-like I consider it unlikely, because the existence of lichens depend on the previous existence of a fungus and an algae [on other planets] we should look for Archaebacteria
Generalities on Lichens The wedding partners: --- the fungus = an Ascomycota (very rarely a Basidio:e.g., Multiclavulina); -------- more than one fungus can be associated to the same lichen colony. --- either an algae ------- Trebouxia, in 80% of lichens: unicellular algae, which has never been found free-living, ------- Trentepohlia (a filamentous, also free-living algae). -------. --- or a filamentous cyanobacterium, often Nostoc. These three genera are the photobiont partner in > 90% of lichens Generalities on Lichens Trebouxia Nostoc Arrows show heterocysts, which are specialized cells that fix atmospheric nitrogen. (pics from Kendrick)
Generalities on Lichens Not necessarily lichenized (=captured by a fungus): Trentepohlia, a free living algae mostly found on lightexposedbark and rock. http://www.bioimages.org.uk/html/r133811.htm Generalities on Lichens Lichen thalli have five kinds of macromorphology (See Kendrick, Chapt. 7) (1) crustose -- so closely applied to the substrate that to collect them you often have to take some of the rock or bark they are growing on as well (2) foliose -- having leafy or tongue-like lobes which are not so closely applied to the substrate; (3) fruticose -- with upright or hanging, stalked, shrubby or hair-like thalli (fruticose means bushy, and has nothing to do with fruit); (4) squamulose -- the thallus is made up of small scales (5) leprose -- powdery, essentially made up of soredia-like aggregations, and lacking a cortex. Mixed macromorphology: E.g., in Cladonia the primary thallus is squamulose but there is also a secondary, upright, reproductive fruticose thallus called a podetium.
Generalities on Lichens fruticose Crustose / foliose http://www.msu.edu/course/bot/423/plntlist8mosslichen.html 3) Asexual (vegetative) reproduction: - soredia: the upper surface of the thallus ruptures, exposing a powdery mass of propagules called soredia, which are small groups of algal cells entangled in fungal hyphae. - isidia: small, finger-like or branched structures called isidia, which grow up from the thallus, then break off. 4) Sexual reproduction: ascus ---> ascospores Generalities on Lichens 5) Lichens produce a wide array of unique secondary metabolites that have been used for systematics (chemotaxonomy); - many lichens have antibiotic properties and have been used in traditional medicine around the world. - extremely slow growth rate; best measured from old gravestones! - extreme longevity; some lichen colonies are thought to be > 4,000 year old - extreme sensibility to air pollution mostly acid rain and carbon dioxide; many cities are virtually lichen-free! Can be used for biocontrol of air-quality. See for instance: http://www.ac.wwu.edu/~fredr/lichen_table.htm
Two small groups of lichens in the Sordariomycetes clade - Lichinomycetes - Arthoniomycetidae (= Arthoniomycetes) Lutzoni et al., 2004, American Journal of Botany Lichen-forming Lichen-forming Sordariomycetes
Sordariomycetes clade:lichinomycetes - derived from Dothideomycetidae - mostly crustose; bitunicate ascus - photobiont = generally cyanobacteria - Examples: Peltula tortuosa Anema nummularium http://www.uni-kl.de/fb-biologie/botanik/sys_tax_flechten.htm Sordariomycetes clade: Arthoniomycetidae - photobiont = Trebouxia - mostly crustose; elongate apothecia with bitunicate ascus - Examples: Arthonia glaucomaria Roccella fusiformis http://www.bildungsservice.at/faecher/geo/flechten%20und%20lichenicole %20Pilze/flechten_und_lichenicole_pilze2.htm http://www.alm-neu.de/jersey/thumbnails/3/roccella%20fuciformis.htm
Lecanoroycetidae, Acarosporoycetidae, and Ostropomycetidae (three predominantly lichen-forming clades = Lecanoromycetes ) - includes the vast majority of lichens - sexual stage of the fungus = mostly Inoperculate Discomycetes (but also some perithecium) - many conflicts between traditional and molecular classification Visit www.lichen.com/ (consider this website to be a mandatory reading) Some interesting or common Lecanoromycetes Letharia vulpina (the Wolf Moss ) - grows mostly on old or dead trees in dry, low temperature old growth coniferous forests; also on old wooden fenceposts. - known only from North America and Europe. - highly sensitive to human disturbance, especially the eradication of old growth forests. ----- It is now reportedly extinct in Finland!. - Toxic; widely used in both Native American and early European civilizations: ---- Scandinavians used to mix with meat to kill wolves, hence its common name, the Wolf Moss. ---- The Achomawi in Northern California used it to make poison arrowheads. ---- Water extracts also used as a medicine for internal problems and in stronger solution to wash external sores and wounds. ---- Also employed as a yellow dye for staining baskets and fibers. http://bio.research.ucsc.edu/people/goff/letharia.htm
Some interesting or common Lecanoromycetes Letharia vulpina (the Wolf Moss ) http://bio.research.ucsc.edu/people/goff/letharia.htm Some interesting or common Lecanoromycetes Xanthoparmelia chlorochroa - a "vagrant" lichen, not attached to any substrate and blown around by wind; - source of dye for Navajo weavers. http://www.lichen.com/
Some interesting or common Lecanoromycetes Usnea species - long use in traditional Chinese and European medicine, also by natives in the Pacific Islands and New Zealand. - contain usnic acid; recently used as antibiotic in salves and deodorants; also used in contemporary homeopathic medicine. http://www.perspective.com/nature/fungi/ Some interesting or common Lecanoromycetes Cladonia species Peltigera species
Eurotiomycetes Two clades: - Chaetothyriomycetidae ----- Capronia, Verrucaria, Pyrenula, Dermatocarpon, Chaetothyriales - Eurotiomycetidae - ascomata = cleistothecium (sometimes lacking) - asci --- scattered within the cleistothecium (i.e. not forming an hymenium) --- thin walled, often ellipsoid, inoperculate, evanescent --- many conidial forms (anamorphs) known Major Orders: Ascosphaerales, Eurotiales and Onygenales Eurotiomycetes Ascosphaerales Genus Ascosphaera - associated with bees: --- feed on pollen and nectar(saprobic); --- some species pathogens on larvae. - ascospores become united in cysts From Alexopoulos
Eurotiomycetes Ascosphaerales Genus Eremascus - lacks ascomata (secondary loss?), otherwise typically ascomycota (no yeast phase known as in Archaeascomycota or Hemiascomycota) - basically, multiple, independent formation of asci from the mycelial mat, - saprophytic From Alexopoulos Eurotiales Eurotiomycetes - Many genera known as teleomorph of Penicillium and Aspergillus spp., and other phialidic anamorphs. (Note that neither Penicillium or Aspergillus are monophyletic ) - Major genera: Talaromyces, Eupenicillium, Eurotium, Emericella Talaromyces anamorphs: Penicillium, Paecilomyces... Pics from Malloch s web site Eurotium anamorphs: Aspergillus,... Eupenicillium anamorphs: Penicillium,... http://www.univ-brest.fr/esmisab/sitesc/ Myco/fiches/Eochra.htm
Penicillium Talaromyces From Alexopoulos Eurotiomycetes Eurotiales: Emericella Emericella nidulans A. Colony on Oatmeal agar, 14 days at 25 C. B. Macrograph of the conidial heads and ascoma. C-D. Hülle cells: C. 700x; D. 1200x. E. Conidiophore and conidia 1700x. F. Ascospores 1600x. Note the Aspergillus anamorph http://www.iums.org/icpaemenid.htm
Eurotiomycetes Onygenales - High diversity in cleistothecium types. - species with the ability to degrade keratin (a major component of skin, hairs, nails, horns, feathers etc ) are all found in this order ===> associations with animals (saprobic or pathogens); pathogenic forms generally the anamorph (sometimes yeast-like), which may have an optimal growth temperature at 37C! Some important genera of human / animals pathogens: - Ajellomyces, Coccidioides, Arthroderma Arthroderma species Conidial anamorphs Microsporum, Trichophyton - saprobic in soil, ringworm on skin Eurotiomycetes Onygenales - High diversity in cleistothecium types - Taxonomic diversity From Alexopoulos
Eurotiomycetes Onygenales Ajellomyces capsulatum Yeast anamorph = Histoplasma capsulatum - lung infection; in immunodepressed people the disease can spread to other body parts including bones <===> More info from http://botit.botany.wisc.edu/toms_fungi/jan2000.html cleistothecium Eurotiomycetes Onygenales Ajellomyces dermatitidis Yeast anamorph Blastomyces dermatitidis (mutinucleate, generally 2-5 nuclei) - skin disease& lesion in humans, dogs, cats etc.; widespread in Africa More info from http://botit.botany.wisc.edu/toms_fungi/jan2001.html <===> Blastomyces dermatitidis Ajellomyces dermatitidis: simple cleistothecium
Eurotiomycetes Onygenales An anamorph Coccidioides immitis: teleomorph not known - systemic human pathogen; also grow in soil - generally starts from lung infection; can transfer to other body parts; allergic reactions (e.g. reddening of skin) - dimorphism mycelium <==> asexual endospores (better adapted for circulation in the blood) <===> More info from http://botit.botany.wisc.edu/toms_fungi/jan2002.html Laboulbeniales / Laboulbeniomycetes - All the ca 1700 described morphospecies are invariably found attached to the exoskeleton of insects, or occasionally, millipedes and mites. - Poorly studied group; evolutionary relationships to other Ascomycota still unclear;. - apparently, don't produce anamorphs; - extreme species and site-specificity: many species are only found in one part of their insect host, for example, on a particular limb, or restricted to either males or females. Fig. on the right: Development of Stigmatomyces baerii on houseflies: An ascospore (A) becomes attached to the animal, germinates, and sends a foot into the exoskeleton to absorb nutrients. Although haustoria may penetrate as far as the epidermal cells, there is never any real invasion of host tissues. The ascospore develops a median septum, and the upper cell becomes differentiated into a male organ, with several phialide-like cells (B) that produce spermatia. The lower cell then develops an ascogonium with a trichogyne, which is fertilized by the spermatia (C). Several asci then develop from the ascogonium, and eventually deliquesce. The mature ascoma is spine-like, projecting from the exterior of the host, and can be seen with a hand-lens (D). Other genera exhibit the same basic features (Hesperomyces - E and Corethromyces - F). From Kendrick