Last Time. Biology of FUNgi. Lecture 24 Community ecology - what rules fungal communities. ! We wrapped up the good, the bad and the ugly.

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1 Biology of FUNgi Lecture 24 Community ecology - what rules fungal communities Last Time! We wrapped up the good, the bad and the ugly.! Systemic fungal infections: blastomycosis, coccidiomycosis, cryptococcosis, and histoplamosis! Opportunistic infections: aspegillosis, candidiasis, zygomycosis plus a great number of air- and soilborne fungi

2 Fungus of the day - Laccaria montana Taxonomy: Phylum (subphylum) Basidiomycota Order - Agaricales Family - Tricholomataceae Common name: salmon mushroom Laccaria montana is another ectomycorrhizal basidiomycete. Species of Laccaria (e.g. Laccaria laccata) are extremely common and colonize young seedlings in the forest nurseries. Many also commonly occur in early successional communities - hence, choice of our shroom du jour. Laccaria montana and a cross section of a mycorrhizal root it mantle, Hartig net and plant vascular stele. Fungus of the day - Laccaria montana Laccaria montana is the most commonly fruiting mycorrhizal fungus in a primary successional glacier forefront. Primary succession occurs after major disturbances. Exemplified by deglaciated areas or areas covered in volcanic eruptions and mine tailings. These areas are void of organic legacies: organic matter, organic nitrogen, and plants and therein associated other organisms. Aerial picture of a glacier forefront, which is dominated by shrub willows that host Laccaria montana.

3 Fungus of the day - Laccaria montana The co-occurrence of ectomycorrhizal fungi and their hosts is essential for plant community establishment. Classic experiments studying pine plantations in areas with no previous pine stands showed that plants could not survive without their fungal symbionts. Look at controls of fungal community assembly Environment Similarly, in primary successional systems, only after arrival of the fungi can the plants grow. Aerial picture of a glacier forefront, which is dominated by shrub willows that host Laccaria montana. Plant community Fungal community

4 Look at controls of fungal community assembly Assembly of fungal communities Filter model components Allochthonous Plant community Environment Fungal community Arrival: Out of site Plant community: Hosts Environment: Tolerances of fungi Competition: Interactions among fungi Local propagule dispersal Autochtonous Autochtonous vegetative colonization

5 Assembly rules Ectomycorrhizal succession: early/ late arrivals. Early successional taxa Late successional taxa Arrival of the fungi Propagules are sparse at early stages of primary succession. MPN studies indicate that fungal are few or absent. Allochthonous Laccaria montana Cortinarius mutabilis estimated number of AM DSE EM * 54.0 ( ) 23. Inocybe lacera Fuscoboletinus ochraceoroseus Distance

6 Monoblepharis_hypogyna[AF164334] Hypocrea_rufa[AJ3011] Hypomyces_chrysospermus[M893] Anamylospora_pulcherrima[AF119501] Bulgaria_inquinans[AJ224362] Byssoascus_striatosporus[AB015776] Oidiodendron_tenuissimum[AB015787] Scutellospora_calospora[AJ306443] Coccodinium_bartschii[U77668] Ophiostoma_piliferum[AJ243294] Penicillium_expansum[AF218786] Urnula_craterium[AF4347] Peziza_griseo_rosea[AF133150] Cryptococcus_aquaticus[AB032621] Mrakia_frigida[AB032665] Exophiala_dermatitidis[X79312] Rhodotorula_aurantiaca[AB030354] Arrival of the fungi Where do the fungi come from? To determine fungal community structure in non-vegetated areas DNA was extracted from soil, PCR-amplified and fungi identified. Allochthonous NJ tree of random 18S amplicons. Taphrina Smuts Rusts Laccaria Fusarium_oxysporum[AF141951] Gibberella_pulicaris[AF149875] Myrothecium_leucotrichum[AJ3012] Claviceps_purpurea[AF281178] Fusarium_merismoides[AF141950] Illosporium_carneum[AF289655] Neurospora_crassa[X04971] Sordaria_fimicola[X69851] Chaetomium_elatum[M83257] Monographella_nivalis[AF064049] Seiridium_cardinale[AF346556] Poronia_punctata[AF064052] Xylaria_carpophila[Z49785] Old_33_04[AY168919] * Old_33_19[AY168920] * Old_26_05[AY1689] * Ophiostoma_stenoceras[M85054] Sporothrix_schenckii[M85053] Young_03_18[AY168892] Penicillium_namylowskii[AB028190] Penicillium_freii[AJ005446] Warcupiella_spinulosa[AB002081] Chromocleista_malachitea[D88323] Eupenicillium_javanicum[U21298] Capronia_pilosella[U42473] Cladophialophora_boppii[AJ232946] Young_01_07[AY168885] Ceramothyrium_linnaeae[AF022715] Capronia_dactylotricha[AJ232943] Trapelia_involuta[AF1194] Trapelia_placodioides[AF119500] Baeomyces_rufus[AF085471] Young_02_04[AY168887] * Young_03_06[AY168890] * Leotia_lubrica[L37536] Leotia_viscosa[AF113715] Young_01_04[AY168884] Young_05_13[AY168896] Pseudogymnoascus_roseus[AB015778] Amylocarpus_encephaloides[U45438] Neobulgaria_premnophila[U45445] Young_04_05[AY168893] Alternaria_longissima[AF229511] Paraphaeosphaeria_quadriseptata[AF250826] Helicascus_kanaloanus[AF053729] Helminthosporium_solani[AF120253] Leptosphaeria_bicolor[U04202] Spilocaea_oleaginea[AF338393] Dothidea_hippophaeos[U42475] Stylodothis_puccinioides[AY016353] Old_31_03[AY168917] Wilcoxina_mikolae[U62014] Trichophaea_hybrida[U53390] Scutellinia_scutellata[U53387] Otidea_onotica[AF006308] Selenaspora_guernisacii[AF4667] Young_07_13[AY168902] Strobiloscypha_keliae[AF0063] Urnula_hiemalis[Z49754] Young_01_08[AY168886] Old_31_18[AY168918] Arthrobotrys_robusta[AJ001988] Dactylella_oxyspora[AF6537] Cazia_flexiascus[U42666] Young_07_12[AY168901] Peziza_petersii[AF133152] Old_21_14[AY168908] ** Pulvinula_archeri[U62012] Debaryomyces_castellii[83819] Debaryomyces_hansenii[X58053] Saccharomyces_cerevisae[Z75578] Saccharomycetes Arxula_adeninivorans[AB018123] Taphrina_deformans[U00971] Taphrina_nana[AB000955] Young_03_08[AY168891] Taphrinomycetes + Neolectomycetes Neolecta_vitellina[Z27393] Crucibulum_laeve[AF026624] Hygrophorus_sordidus[AF287834] Laccaria_pumila[AF287838] Young_07_[AY168900] * Athelia_bombacina[M55638] Cystostereum_murraii[AF082850] Young_08_16[AY168904] Panellus_serotinus[AF026590] Calvatia_gigantea[AF026622] Lycoperdon_sp.[AF026619] Entoloma_strictius[AF287832] Pluteus_petasatus[AF026634] Termitomyces_sp.[AB051893] Humidicutis_marginata[AF287833] Tricholoma_myomyces[AF287841] Inocybe_geophylla[AF287835] Old_20_06[AY168907] Old_26_16[AY168912] Boletus_satanas[M94337] Hymenomycetes Paragyrodon_sphaerosporus[M90826] Xerocomus_chrysenteron[M94340] Paxillus_panuoides[AF026628] Tapinella_atrotomentosa[M90824] Old_27_14[AY168916] Gigaspora_albida[Z14009] Gigaspora_margarita[X58726] Gigaspora_gigantea[Z140] Scutellospora_cerradensis[AB041345] Thelephora_sp.[AF026627] Daedalea_quercina[U59067] Old_21_15[AY168909] Russula_compacta[AF026582] Amylostereum_areolatum[AF082845] Laurilia_sulcata[AF334918] Polyporoletus_sublividus[AF287840] Rhizoctonia_solani[D85636] Rhizoctonia_solani[D85643] Thanatephorus_praticola[M920] Marchandiomyces_sp.[AF289662] Young_08_11[AY168903] ** Lecanoromycetes Exobasidium_vaccinii[AJ271382] Sympodiomycopsis_paphiopedili[D14006] Young_02_07[AY168888] Young_03_03[AY168889] Acaulospora_laevis[Y17633] Acaulospora_spinosa[Z14004] Glomeromomycota Glomus_caledonium[Y17653] Glomus_geosporum[AJ245637] Glomus_mosseae[U96141] Glomus_intraradices[AJ301859] Glomus_sinuosum[AJ133706] Dissophora_decumbens[AF157133] Echinosporangium_transversale[AF113424] Mortierella_alpina[AJ271630] Mortierella_wolfii[AF113425] Zygomycota Endogone_pisiformis[X58724] Young_05_16[AY168897] Young_06_04[AY1688]??? Neocallimastix_frontalis[X80341] Spizellomyces_acuminatus[M59759] Chytridiomycota Sordariomycetes Eurotiomycetes Chaethyriomycetes Leotiomycetes + Onygenales Dothideomycetes Pezizomycetes Ustilaginomycetes Camptobasidium_hydrophilum[U75449] Uredinomycetes Young_06_02[AY168898] Arrival of the fungi Allochthonous These studies gave surprising results. Non-vegetated areas contained biotrophic and ectomycorrhizal fungi.

7 Arrival of the fungi Propagules were few in early succession Biotrophic and EM fungi common in early successional, nonvegetated samples. Since no or only few plants are present in this area, establishment of propagule bank from allochthonous sources. Allochthonous Host availability Clearly, no fungal associations can occur in absence of appropriate substrate or susceptible host plant. Host: Salix spp. EM fungus: Laccaria sp. Early succession Host specificity Late succession Host: Abies sp. Salix spp. Larix sp. EM fungus: Fuscoboletinus sp.

8 Host availability Host availability Host specific EM fungi appear only later in succession as their host community has established. Host: Salix spp. Early succession Host specificity Late succession Host: Abies sp. Salix spp. Larix sp. Sum up Although the propagule bank may contain a number of different fungi with narrow or broad host ranges, host-fungus compatibility determines which fungi can successfully establish. Host specificity EM fungus: Laccaria sp. EM fungus: Fuscoboletinus sp.

9 Environmental tolerance Environmental tolerance Drastic environmental gradients over the glacier forefront allow determination of fungal tolerances. Early succession Environmental Late succession Local environmental gradients, similarly, give clues to fungal preferences. Canopies provide such local gradient. Environmental Low OM Low N Low microbial biomass Higher OM Higher N Higher microbial biomass Low OM Low N Low microbial biomass Higher OM Higher N Higher microbial biomass

10 Environmental tolerance Environmental tolerance Environmental preferences among fungi determined by sampling of (inter-)canopy areas at various locations along the forefront. Environmental Several environmental preferences detected. Two contrasting examples: 1) Thelephoraceae sp. 1 2) Sordariaceae sp. 2 Environmental Thelophoraceae sp. 1 Sordariaceae sp m 450m 900m 600m 750m Probability of occurrence, p (%) Probability of occurrence, p (%) Distance (m) Distance (m)

11 Environmental tolerance Environmental tolerance 1) Thelephoraceae sp. 1 Early successional and intercanopy areas Adapted to low OM/N conditions Environmental 2) Sordariaceae sp. 2 Late successional and canopy areas Adapted to higher OM/N conditions Late successional species Environmental Thelophoraceae sp. 1 Sordariaceae sp. 2 Thelophoraceae sp. 1 Sordariaceae sp. 2 Probability of occurrence, p (%) Probability of occurrence, p (%) Probability of occurrence, p (%) Probability of occurrence, p (%) Distance (m) Distance (m) Distance (m) Distance (m)

12 Environmental tolerance Biotic Conclusions After selection of fungal species from the propagule bank by the host preference, environmental tolerance such as OM and N (or other correlated with these) select taxa that can persist in the environment. This is a subset of taxa that were selected by hosts from the broader propagule bank. Environmental The components of the biotic include competition among the species that establish with the available hosts and persist in the environmental setting. Only the superior competitors will be able to remain as a persistent component in the community. Biotic

13 Local dispersal In the early stages of succession only few fungi produce. Therefore, the initial colonization is exclusively (?) a result of allochthonous. estimated number of AM DSE EM * 54.0 ( ) Distance 23. Autochthonous Local dispersal Once fungal communities have been established and fruiting occurs, autochthonous sources of important. Locally, vegetative colonization by mycelial connections important factor. Local short-distance dispersal + Local Autochthonous Vegetative colonization Vegetative colonization via mycelial connections

14 Conclusions Local dispersal Summary Allochtonous Autochthonous propagule production contribute increasingly to the propagule bank. Mycelial connections become very important for colonization. Local Autochthonous Vegetative colonization Proposal for a general successional model for fungi - includes allochthonous and autochthonous propagule sources - based on host, environmental and biotic filters Autochtonous Autochtonous vegetative colonization

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