MYCORRHIZAL ASSOCIATIONS: The Web Resource

Section 9. ECTOMYCORRHIZAL FUNGI

A. Introduction

This section provides a brief introduction to ectomycorrhizal fungus diversity, primarily using examples from Australia. More information is available using the links provided below.

Images on this page are by Neale Bougher (Western Australian Department of Environment and Conservation) and Mark Brundrett, unless otherwise stated and cannot be used without permission. Many of Neale Bougher's photos are from the Perth Urban Bushland Fungi Field Book, which contains more information about these species.

Sources of information on Australian Fungi

B. Structural Diversity

Most fungi have fruit-bodies that are clearly visible for a brief period usually in the autumn in temperate regions, or during the wet season in tropical regions, when spores are dispersed. As illustrated in the diagram below, there is a wide taxonomic and structural diversity of visible fungi (called macrofungi or larger fungi). The reproductive structures of larger fungi include epigeous mushrooms, puffballs, coral fungi, crust fungi, etc. and subterranean (hypogeal) fungi called truffles, or truffle-like fungi. Most of these categories contain ectomycorrhizal fungi.

Hover mouse over diagram to see which categories contain ectomycorrhizal fungi (ECMF)

Structural Diversity of Larger Fungi with ECM associations

Many ECM fungi have mushroom-like fruit bodies with gills, tubes, teeth, etc. arising from soil, but there are also many saprophytes in most of these categories.
The majority of subterranean fungi (called truffles or truffle-like) are considered to be ECM associates, but some probably are not.
Fungi fruiting on wood are usually not mycorrhizal, with the exception of some crust fungi.
Additional information is required to identify mycorrhizal fungi (listed in the table below).

C. Preliminary list of Ectomycorrhizal Fungi

The majority of ECM fungi are Basidiomycetes, with some Ascomycetes and a few Zygomycetes, as listed in the Table below. Fungi in this table are organised into clades based on molecular phylogeny with genera confirmed to be symbiotic plant associates indicated. The arrangement of clades in the fungal tree of life is not fully resolved.

Clade No.	Family or Group	Epigeous Larger Fungi (mushrooms, puffballs, etc.)	Truffles (hypogeous, or semi-hypogeous)	Resupinate (crust fungi)
Basidiomycetes
1 (34)	Lyophyllum	Lyophyllum*
2 (37-38)	Tricholomataceae	Leucopaxillus, Tricholoma*
3 (43/44)	Entolomataceae	Entoloma* (not all species)	Rhodogaster, Richoniella
4 (55)	Amanitaceae	Amanita Limacella	Ammarrendia, Torrendia
5 (65)	Hygrophoraceae	Gliophorus, Humidicutis, Hygrophorus*
6 (72)	Hydnangiaceae (Laccaria clade)	Laccaria	Hydnangium, Podohydnangium, Gigaspera*
7 (73)	Cortinariaceae	*Cortinarius, Dermocybe*, Naucoria, Rozites*	Cortinarius, Hymenogaster (in part), Destuntzia, Protoglossum, Quadrispora, Setchelliogaster, Stephanopus, Thaxterogaster*
8 (74)	Phaeocollybia clade in Hymenogastraceae	Phaeocollybia*
9 (95)	Hebeloma clade in Hymenogastraceae	Hebeloma*	*Hymenogaster* (in part)
10 (78/79)	Descolea clade (in Bolbitiaceae?)	Descolea*	*Descomyces, Setchelliogaster**
11 (104)	Inocybaceae	Inocybe*, Auritella	Auritella
12A (Outgroup 1)	Boletes (Boletales) Includes many families	Aureoboletus, Austroboletus, Austropaxillus, Boletellus, Boletochaete, Boletus, Chroogomphus, Fuscoboletinus, Heimielia, Gomphidius, Gyroporus, Leccinum, Paragyrodon, Paxillus, Phlebopus, Phylloporus, Poryphyrellus, Psiloboletinus, Rubinoboletus, Strobilomyces, Suillus, Tylopilus, Xanthoconium, Xerocomus, etc.	*Alpova, Austrogaster, Austrogautieria*, Chamonixia, Gastroboletus, Gastrotylopilus, Gymnogaster, Gymnopaxillus, Horakiella, Hysterogaster, Melanogaster, Mycoamaranthus, Octaviania, Rhizopogon, Royoungia, Sclerogaster, Truncocolumella*, Timgrovea, Wakefieldia, etc.	(Serpula, etc. are saprophytes)
12B (Outgroup 1)	Boletales - Sclerodermatales	Boletoid: Gyroporus, Gasteroid: Pisolithus, Scleroderma, Calostoma*	Astaeus, Horakiella, Scleroderma,*, Velligaster
13 (Outgroup 2)	Russulales	Lactarius, Russula	Arcangeliella, Cystangium, Gymnomyces, Leucogaster, Macowanites, Octaviania, Stephanospora, Zelleromyces	Byssoporia, Albatrellus, Polyporoletus (Stereum, etc. are saprophytes)
14 (Outgroup 3)	Gomphales & Hysterangiales	Bankera, Boletopsis, Clavaridelphus, Gomphus, Hydnum, Hydnellum, Phellodon, Sarcodon, Ramaria** (some NM?)*	Hysterangium clades: Aroramyces, Chondrogaster, Hysterangium* , Mesophelliaceae: Andebbia, Castoreum, Gummiglobus, Malajczukia, Mesophellia, Nothocastoreum, Gallacaceaceae: Austrogautiera, Gallacea, etc., Phallogaster clade: Protrubera, Phallogaster, Trappea Ramaria clade: Gautieria*
15 (Outgroup 4)	Cantharellales	Cantharellula, Cantharellus, Craterellus		Sistotrema?
16AB (Outgroup(s) 5)	Rhizoctonia alliance clades (Sebacinaceae, Ceratobasidiaceae)			Thanatephorus, Sebacina
17 (Outgroup 6)	Thelephorales	Bankera, Boletopsis, Thelephora, Phellodon, Sarcodon		Tomentella, Pseudotomentella, Tomentellopsis*
18 (Outgroup 7)	Clavariaceae	Aphelaria, Clavaria, Clavariadelphus, Clavicorona, Clavulina, Clavulinopsis, Ramariopsis
19 (Outgroup 8)	Atheliales			Amphinema, Byssocorticium, Byssosporia, Piloderma, Tylospora*
Ascomycetes
20 (Ascomycete-1)	Pezizales (inc. Tuberaceae, Helvellaceae, etc.)	Genea, Geopora, Humaria, Hydnotrya, Helvella, Leucangium, Pachyphloeus, Pulvinula, Sarcosphaera, Sphaerosporella, Sphaerozone, Tirmania, Tricharina, Wilcoxina*	Choiromyces, Dingleya, Eremiomyces, Kalaharituber, Labyrinthomyces, Pachyphloeus, Reddellomyces, Tuber, Terfezia, Turmania*	Phialophora* (anamorph)
21 (Ascomycete-2)	Eurotiales (Elaphomycetaceae)		Elaphomyces* Pseudotulostoma*
22 (Ascomycete-3)	Melanommatales			*Cenococcum* (sterile fungus)
Zygomycetes
23 (Zygomycete-1)	Endogonaceae		Densospora, Endogone, Peridiospora, Youngiomyces

Clades: Fungal clades are provided in the first column are ordered using the 117 clade classification of the euagarics by Moncalvo et al. 2002, with outgroups numbered to reflect increasing distance from the agarics in that tree.
Orders and Families: Follow Matheny et al. 2006 and Hibbett et al. 2007.
Genera: Follow cited references (some are synonyms).
Classification and Identification References: Arora 1986, 1991, LoBuglio et al. 1996, Binder et al. 1997, Erland & Taylor 1999, Kõljag et al. 2000, Bougher & Lebel 2001, Humpert et al. 2001, Peinter et al. 2001, Moncalvo et al. 2000, 2002, Binder & Bresinsky 2002, Selosse et al. 2002, Tehler et al 2003, Urban et al. 2003, Binder et al. 2005, Douhan & Rizzo 2005, Ferdman et al. 2005, Henkel et al. 2006, Matheny & Bougher 2006, Matheny et al. 2006, Tedersoo et al. 2006, Trocha et al. 2006, Watling 2006, Barroetaveña et al. 2007, Hibbett et al. 2007, Hosaka et al. 2007, Wilson et al. 2007.
Genera in Bold: Confirmed ECM Associations by fungal isolation and re-synthesis under controlled conditions: Warcup 1980, 1985, Kropp & Trappe 1982, Malacjzuk et al. 1982, Molina & Trappe 1982a, Bougher et al. 1985, Godbout & Fortin 1985, Reddell & Milnes 1992, Burgess et al. 1993, Massicotte et al. 94, Thomson et al. 1994, Cripps & Miller 1995, McGee 1996, Kawai 1997, Lu et al. 1998, Reddell et al. 1999, Baxter & Dighton 2001, Yamada et al. 2001ab, Yun & Hall 2004, Brundrett et al. 2005, Henkel et al. 2006.
Genera with an Asterisk(*): Links between fruitbodies and ECM morphotypes established, as summarised by Agerer 2006. Additional information: Norvell 1998, Wilson et al. 2007.
Excluded Taxa: These include:
(i) Catathelasma seems to be ectomycorrhizal, but this is not fully resolved (Hutchison 1992).
(ii) Truffle-like fungi that may be saprobic: Endoptychum, Cribbea, Protubera, Gelopellis, Secotium, Stephanospora, Gastrocybe, Weraroa, Tympanella, etc.
(iii) Ascomycetes that fruit after fire (discomycetes), such as Peziza, Morchella, and Scutellinia, considered to be saprobes (see Fujimura et al. 2005).
(iv) The bolete - Gyrodon (Boletinellus) merulioides is not mycorrhizal (see Section 10).

D. Examples of Ectomycorrhizal Fungi

Examples of larger genera of ECM fungi are provided below, primarily using fungi from Australia and New Zealand. Most of the major groups of ECM fungi are well represented in Australia, including over 700 species that have been named (Bougher 1995, Bougher & Lebel 2001). The majority of these fungi are endemic. However, taxonomic studies on Australian fungi are at an early stage compared to the Northern Hemisphere and new species are continually being discovered (Pascoe 1991, May 2001, Bougher & Lebel 2001). Information on Australian fungi can be obtained from books and the websites listed above.

Laboratory and glasshouse experiments have shown that different fungi vary in their capacity to utilise resources, withstand adverse conditions, etc. For this reason, mycorrhizal fungus diversity is thought to contribute to the resilience of ecosystems, but there is very little comparative data that can be used to test this hypothesis.

2. Tricholomataceae

Fungus: Tricholoma sp. Form: mushroom with gills Habitat: Eucalyptus dominated urban bushland Location: Kensington, Western Australia Photo: Neale Bougher	Fungus: Tricholoma sp. (Matsutake) Form: mushroom with gills (harvested for export) Habitat: conifer dominated temperate forest Location: Yunnan Province, China

3. Entolomataceae
	Fungus: Entoloma moongum Form mushroom with gills Habitat: Eucalyptus marginata dominated urban bushland Location: Kensington, Western Australia Photo: Neale Bougher

4. Amanitaceae

Fungus: Amanita carneiphylla Form: mushroom with gills Habitat: Eucalyptus dominated woodland Location: urban bushland, Perth, Western Australia Photo: Neale Bougher	Amanita sp. Fungus:Form: mushroom with gills Habitat: Eucalyptus marginata dominated mediterranean woodland Location: urban bushland in Perth, Western Australia

Fungus: Torrendia grandis Form: truffle-like (semi-hypogeous) Habitat: Eucalyptus dominated mediterranean sclerophyllous forest Location:Western Australia Photo: Neale Bougher	Fungus: Ammarendia oleosa Form: truffle-like (hypogeous) Habitat: Eucalyptus dominated mediterranean sclerophyllous forest Location:Western Australia Photo: Neale Bougher

5. Hygrophoraceae

Fungus: Humidicutis viridimagentea Form: Mushroom with gills Habitat: Eucalyptus and Agonis woodland (probably mycorrhizal) Location: Denmark, Western Australia	Fungus: Hygrocybe sp. (probably not mycorrhizal) Form: Mushroom with gills Habitat: Eucalyptus woodland Location: William Bay, Western Australia Photo: Neale Bougher

6. Hydangiaceae - the Laccaria clade

Fungus: Laccaria sp. Form: mushroom with gills Habitat: Eucalyptus marginata dominated mediterranean sclerophyllous forest Location: Western Australia	Fungus: Hydnangium carneum Form: truffle (hypogeous) Habitat: Eucalyptus dominated mediterranean sclerophyllous forest Location: Western Australia Photo: Neale Bougher

7. Cortinariaceae

Fungus: Cortinarius archeri Form: mushroom with gills Habitat: Eucalyptus marginata dominated mediterranean sclerophyllous forest Location: Western Australia.	Fungus: Cortinarius (Dermocybe) canarium Form: mushroom with gills Habitat: Nothofagus dominated temperate rainforest Location: Near Te Anau, New Zealand

Fungus: Cortinarius persplendidus (Dermocybe splendida) Form: mushroom with gills Habitat: Eucalyptus marginata dominated mediterranean sclerophyllous forest Location: Western Australia.	Fungus: Cortinarius vinaceolamellatus Form: mushroom with gills Habitat: Eucalyptus marginata dominated mediterranean sclerophyllous forest Location: Western Australia
	Fungus: Thaxterogaster purpureum Form truffle-like (partially hypogeous) Habitat: Nothofagus dominated temperate rainforest Location: Near Te Anau, New Zealand Roll-over: collection showing cross section (grid = 1mm)

8. Hebeloma a Clade in the Hymenogastraceae
	Fungus: Hebeloma westraliense Form: mushroom with gills Source: fruiting after inoculaion in pasteurised soil Habitat: Eucalyptus urophylla growing in a pot Location:Western Australia Photo: Neale Bougher

10. Descolea a Clade in the Bolbitiaceae
	Fungus: Descolea (C), with its truffle-like relatives Setchelliogaster (B) and Descomyces (A) Forms: mushroom with gills and truffles Habitat: Eucalyptus marginata dominated mediterranean sclerophyllous forest Location: Western Australia Photo: Neale Bougher

11. Inocybaceae
	Fungus: Inocybe violaceocaulis Form: mushroom with gills Habitat: Eucalyptus dominated mediterranean sclerophyllous forest Location: Western Australia Photo: Neale Bougher

12. Boletales

Fungus: Austroboletus lacunosus Form: mushroom with pores Habitat: Eucalyptus marginata dominated mediterranean sclerophyllous forest Location: Western Australia	Fungus: Austropaxillus sp. Form: mushroom with gills Habitat: Eucalyptus marginata dominated mediterranean sclerophyllous forest Location: Western Australia

Fungus: Calostoma rodwayi Form: stalked puffball (gasteroid) Habitat: Eucalyptus marginata dominated mediterranean sclerophyllous forest Location: Western Australia	Fungus: Pisolithus sp. Form: puffball (gasteroid) Habitat: erupting through bitumen paving under a eucalypt tree in car park Location:The University of Western Australia

Fungus: Austrogautieria manjimupana Form: truffle Habitat: Eucalyptus marginata dominated mediterranean sclerophyllous forest Location: Western Australia Photo: Neale Bougher	Fungus: Rhizopogon rubescens Form: truffle (semi-hypogeous) Habitat: Pinus radiata plantation Location: Western Australia Photo: Neale Bougher

13. Russulaceae

Fungus: Lactarius deliciosus Form: mushroom with gills Habitat: Edible fungus collected in the wild and sold in a market Location: Yunnan Province, China	Fungus: Lactarius eucalypti Form: mushroom with gills Habitat: Eucalyptus marginata dominated mediterranean sclerophyllous forest Location: Western Australia

Fungus: Russula persanguinea Form: mushroom with gills Habitat: Eucalyptus marginata dominated mediterranean sclerophyllous forest Location: Western Australia	Fungus: Zelleromyces sp. Form: truffle (hypogeous) Habitat: Eucalyptus marginata dominated mediterranean sclerophyllous forest Location: Western Australia
	Fungus: Cystangium sessile Form: truffle (hypogeous) Habitat: Eucalyptus marginata dominated mediterranean sclerophyllous forest Location: Western Australia Photo: Neale Bougher

14. Gomphales

Fungus: Ramaria ochraceosalmonicolor Form: coral fungus Habitat: Eucalyptus marginata dominated mediterranean sclerophyllous forest Location: Western Australia	Fungus: Ramaria abitina Form: coral fungus Habitat: conifer dominated boreal forest Location: Ontario, Canada

Fungus: Ramaria versatilis Form: coral fungus Habitat: Eucalyptus marginata dominated mediterranean sclerophyllous forest Location: Western Australia	Fungus: Gomphus floccosus Form: gomphoid (with folds) Habitat: mixed conifer and deciduous forest Location: Quebec, Canada


Fungus: Hydnum repandum sp. (roll-over shows close-up of teeth under cap) Form: tooth fungus (mushroom-like) Habitat: Eucalyptus dominated mediterranean sclerophyllous forest Location:Augusta, Western Australia	Fungus: Phellodon sp. Form: tooth fungus (mushroom-like) Habitat: Eucalyptus dominated mediterranean sclerophyllous forest Location:Western Australia Photo: Neale Bougher

Fungus: Hysterangium sp. Form: truffle (hypogeous) Habitat: Eucalyptus dominated mediterranean sclerophyllous forest Location: Western Australia Photo: Neale Bougher	Fungus: Mesophellia brevispora Form: truffle (hypogeous) Habitat: Eucalyptus conferruminata and Taxandria sp. mallee woodland Location: Bald Island, Western Australia Photo: Neale Bougher

15. Cantarelloid clade

Fungus: Cantharellus sp. Form: chanterelle (mushroom-like) Habitat: mixed Eucalyptus and Agonis forest Location: Augusta, Western Australia	Fungus: Craterellus cornicopioides Horn of Plenty Form: chanterelle-like (no gills) Habitat: mixed conifer and deciduous forest Location: Quebec, Canada

17. Thelephoraceae
	Fungus: Thelephora sp. Form: coral-like (form higly variable) Habitat: opportunist in pots with Melaleuca sp. Location: Kings Park, Western Australia Roll-over: microscopic view of spores

	Fungus: Tomentella sp. Form: crust fungus on wood (resupinate) Habitat: Eucalyptus dominated mediterranean sclerophyllous forest Location:Western Australia Photo: Neale Bougher
	Mycorrhiza: associated with this crust fungus

18. Clavariaceae
	Fungus: Clavaria amoena Form: coral fungus (suspected to be mycorrhizal) Habitat: Eucalyptus dominated woodland Location: William Bay, Western Australia Photo: Neale Bougher

20. Pezizales (Ascomycetes)
	Fungus: Labyrinthomyces sp. Form: truffle (hypogeous) Scale: Bar = 1 cm Habitat: Eucalyptus dominated mediterranean sclerophyllous forest Location: Western Australia Photo: Neale Bougher
	Fungus: Hydnoplicata convoluta Form: semi-hypogeous, truffle-like cup fungus (suspected to be mycorrhizal) Habitat: Eucalyptus dominated mediterranean sclerophyllous forest Location: Western Australia Photo: Neale Bougher
	Fungus: Tuber melanospora Form: hypogeous, French Black Truffle Habitat: cultivated in plantation (1 kg) Location: Manjimup, Western Australia Roll-over: ectomycorrhizal roots Photos: Nick Malajczuk The Wine & Truffle Co.

22. Melanommatales (Ascomycetes)
	Fungus: Cenococcum sp. Form: sterile mycelia (arrows) and black ECM (S) Habitat: Eucalyptus globulus plantation Location: Western Australia

23. Endogonaceae (Zygomycetes)
	Fungus: Endogone pisiformis Form: Truffle-like (epigeous on sphagnum moss) Habitat: coniferous forest Location: Ontario, Canada

E. Ectomycorrhizal Morphotypes

Characteristic features of ECM root tips used to identify associated fungi (Agerer 1986, Ingleby et al. 1990, Haug & Pritsch 1992) are briefly summarised in a table below. Some examples of roots colonised by known fungi in glasshouse or sterile culture experiments are also provided. Comprehensive information on mycorrhizal morphotypes is provided on the Deemy website.

A. Ectomycorrhiza of Pinus radiata inoculated with Suillus brevipes with dichotomously branched roots (arrows).	B. Ectomycorrhiza of Pinus radiata inoculated with Amanita mascaria
C. Pinus radiata and Boletus edulis mycorrhiza with dichotomously branched roots (arrows).	D. Eucalyptus maculata and Astraeus pteridis ectomycorrhiza with numerous mycelial strands (star).
E. Synthesised association of Tylopilus sp. and E. urophylla seedlings grown in the glasshouse. Note black ECM roots (arrow) and abundant mycelium (star).	F. Synthesised association of Redellomyces sp. and E. grandis seedlings grown in the glasshouse. Note shinny brown ECM roots (arrows).
G. Synthesised ECM associations of Pisolithus sp. and E. urophylla seedlings grown in the glasshouse. Note yellow-orange roots (arrows) and mycelial strands (asterisk).	H. Synthesised ECM associations of Amanita sp. and E. urophylla with yellow roots (arrows) and a dense hyphal covering (star).
Source: Figure 4.7 in Brundrett et al. 1996

tuberculate ECM roots of Eucalyptus (17KB)

Synthesised ECM associations of Thanatephorus gardneri (Rhizoctonia alliance) and Melaleuca sp. grown in the glasshouse. Note relatively thin reddish ECM roots and thick black mycelia.

Tuberculate ECM of Eucalyptus pilularis collected in Queensland. Numerous ECM roots (arrows) are encased within an outer rind (R).

R = peridium,
Arrows = ECM roots

Table of diagnostic features of ectomycorrhizal roots that may be associated with particular ectomycorrhizal fungi.

Mycorrhizal development

root branching patterns and density
root thickening
reduction in root elongation

External hyphae

presence, abundance, distribution
arrangement - single, strands, rhizomorphs
structure - colour, wall, thickness, clamp connections, crystals
sclerotia

Mantle

thickness
colour, changes with bruising, or age
hyphal organization in surface and in deeper layers

loose or compact hyphae or pseudo-parenchyma, prosenchyma
hyphae - thickness, wall structure, clamp connections, septae

cystidia, crystals, exudates
reactions to chemicals (Melzer's reagent, KOH, etc.)
staining reactions
odour
fluorescence

Hartig net

presence, thickness
hyphal organisation
hyphal structure

Index