fungi and fire in australian ecosystems: a review of ... · fungi and ﬁre in australian...

Fungi and fire in Australian ecosystems a review of currentknowledge management implications and future directions

Sapphire J M McMullan-FisherAH Tom W MayB Richard M RobinsonCTina L BellD Teresa LebelB Pam CatchesideEF and Alan YorkG

ASchool of Geography and Environmental Studies Private Bag 78 Hobart Tas 7001 AustraliaBRoyal Botanic Gardens Melbourne Private Bag 2000 Birdwood Avenue South Yarra Vic 3141 AustraliaCDepartment of Environment and Conservation Brain Street Manjimup WA 6258 AustraliaDFaculty of Agriculture Food and Natural Resources University of Sydney Sydney NSW 2006 AustraliaEState Herbarium of South Australia PO Box 2732 Kent Town SA 5071 AustraliaF School of Biological Sciences Flinders University PO Box 2100 Adelaide SA 5001 AustraliaGDepartment of Forest and Ecosystem Science University of Melbourne Water Street CreswickVic 3363 Australia

HCorresponding author Email sapphireflyanglercomau

Abstract Fungi are essential components of all ecosystems in roles including symbiotic partners decomposers andnutrient cyclers and as a source of food for vertebrates and invertebrates Fire changes the environment inwhich fungi live byaffecting soil structure nutrient availability organic and inorganic substrates and other biotic components with which fungiinteract particularly mycophagous animals We review the literature on fire and fungi in Australia collating studies thatinclude sites with different time since fire or different fire regimes The studies used a variety of methods for survey andidentification of fungi and focussed ondifferent groups of fungiwith an emphasis on fruit-bodies of epigealmacrofungi and alack of studies on microfungi in soil or plant tissues There was a lack of replication of fire treatment effects in some studiesNevertheless most studies reported some consequence of fire on the fungal community Studies on fire and fungi wereconcentrated in eucalypt forest in south-west and south-easternAustralia andwere lacking for ecosystems such as grasslandsand tropical savannahs The effects of fire on fungi are highly variable and depend on factors such as soil and vegetation typeand variation infire intensity andhistory including the length of time betweenfires There is a post-fireflushof fruit-bodies ofpyrophilous macrofungi but there are also fungi that prefer long unburnt vegetation The few studies that tested the effect offire regimes in relation to the intervals between burns did not yield consistent results The functional roles of fungi inecosystems and the interactions of fire with these functions are explained and discussed Responses of fungi to fire arereviewed for each fungal trophic group and also in relation to interactions between fungi and vertebrates and invertebratesRecommendations aremade to includemonitoring of fungi in large-scalefiremanagement research programs and to integratethe use of morphological and molecular methods of identification Preliminary results suggest that fire mosaics promoteheterogeneity in the fungal communityManagement of substrates could assist in preserving fungal diversity in the absenceofspecific information on fungi

Introduction

Both fire and fungi affect most other biota particularly in fire-prone Australian ecosystems Despite their obvious importancethe two have rarely been considered together Warcup (1981)briefly summarised the interaction between fire and non-vascularplants (including fungi) in Australia For the Jarrah and Karriforests of south-westernAustralia Robinson andBougher (2003)concluded that fire favours some fungi but not others GloballyCairney and Bastias (2007) found that fire generally alters thecommunity structure of fungi in soil with site- or fire-specificeffects which aremore pronouncedwith repeated burning In thisreview we collate and interpret data from published Australianstudies dealingwith the influence offire on fungi in the context of

the ecological roles of fungi in ecosystems and their interactionswith other organisms

The fungal kingdom is extremely diverse with a range ofvital ecological roles and a high degree of interdependency withother organisms Fungi are heterotrophic and employ a varietyof nutritional strategies including saprotrophism parasitism andthe formation ofmutualistic partnerships such asmycorrhizas andlichens (May and Simpson 1997) Saprotrophic or decomposerfungi grow in soil (Bridge and Spooner 2001) or directly in litterand wood (Rayner and Boddy 1988) Due to their ability to breakdown complex compounds such as cellulose and lignin they areparticularly important in the degradation of organic matter Mostplants form mycorrhizas and this partnership is particularly

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important in old highly weathered and nutrient-poor soils(Brundrett 2009) such as those found across the majority ofthe Australian landscape Fungi are essential participants in thecycling of carbon nitrogen phosphorus and other nutrientsin ecosystems (Berg and Laskowski 2006) Consumption offruit-bodies and mycelium of fungi by mycophagous animalsparticularly invertebrates also contributes to nutrient cyclingAnother critical role of fungi in ecosystem function is theirinfluence on soil in relation to ionic exchange particleaggregation carbon content water holding capacity and waterinfiltration (Bastias et al 2006a Claridge et al 2009a)

Fungi are a significant but often overlooked component ofthe Australian biota Some 11 846 described species of fungiare known from Australia of which 3495 form lichens(Chapman 2009) Estimated diversity is at least 10 000 speciesfor macrofungi (about twice as many as are currently known)and as high as 250 000 species for microfungi (Chapman 2009)Not only are there many species awaiting formal descriptionbut the biology and ecology of named species is usually poorlyknown Very few fungi are formally listed on state or nationalconservation schedules (May 1997 2003) Management of fungiin native ecosystems is compromised by the almost total lack ofstaff with expertise in or responsibility for fungi in conservationand management agencies

Fire both planned and unplanned is an integral part ofAustralian ecosystems and impacts on fungi in various waysthrough physical and chemical changes to habitats andsubstrates Heating during fire causes sterilisation of uppersoil layers and loss of nutrients and carbon via volatilisationand combustion of litter and soil organic matter (Certini 2005)Most soil nutrients are concentrated in these upper layers(Jobbaacutegy and Jackson 2001) as are most organisms (Lee andFoster 1991 Dahlberg 2001 Huang et al 2005) For most firesradiation provides enough heat to kill soil organisms includingfungi in the upper soil layer although heat intensity declinesrapidlywith depth (Humphreys and Lambert 1965 Raison et al1985) Heating to 60C is sufficient to kill living tissueparticularly unprotected fungal mycelia (Schenck et al 1975Klopatek et al 1988) and upper layers of soil often reach thistemperature during fires (Humphreys and Lambert 1965Bradstock et al 1992 Pattinson et al 1999) Neverthelessfungi that have evolved in fire-prone Australian environmentshave adaptations to cope with fire such as heat-resistant sporesor other resting structures (Warcup 1981) Fire can also affectfungi by removing and creating substrates for saprotrophs suchas litter and woody debris or by detrimental effects onmutualistic partners such as spore dispersers or mycorrhizalhosts (Dahlberg 2002)

Soil nutrients are altered by fire which in turn impacts theutilisation and genesis of nutrients by fungi and consequentlyother biota Despite a direct loss of nitrogen from soil viavolatilisation nitrogen and phosphorus availability in soilfollowing fire can increase or decrease depending on fireintensity (Raison 1979 Raison et al 1985 Gray and Dighton2006) Changes in nutrient availability subsequently impact onrecovery of vegetation and nutrient cycling (Neary et al 19992005Howell et al 2006) Fire also alters soil fungal communitiesinvolved in decomposition (Neary et al 1999 Boddy 2001)therefore altering carbon nitrogen ratios and mineralisation

rates which ultimately affect plant growth and productivity(Neary et al 1999 2005 Boddy 2001)

The understanding of fire regimes and the effects of fire onAustralian vegetation has developed rapidly from the generallyscience-based endeavour of lsquoFire and the Australian Biotarsquo(Gill et al 1981) to a much more management- consequence-and biodiversity-orientated approach (see Bradstock et al2002 Abbott and Burrows 2003) Recent extensive fires thathave occurred on most continents have prompted recognitionthat the incidence and characteristics of large fires may alter inthe future as a consequence of global climate change (Williamsand Bradstock 2008) Fires whether planned or unplannedindependent of season are patchy and the effects on soil areequally irregular (Bradstock 2008) It is now widely recognisedthat no single optimum fire regime will meet all managementobjectives (Burrows 2008) and diversity of fire regimes amonglandscapes including long unburnt patches is now beingaccepted by land management agencies in adaptive firemanagement programs Fire mosaics resulting from cumulativeheterogeneity (patchiness) from successive fires may contributeto increased local biodiversity (Grove et al 2002 Bradstock2008 Burrows 2008) Although fire management policies aim tomaximise biodiversity the data upon which decisions are basedpredominantly concernvegetation fuel quantities andconditionsand the likely locations of recognised threatened species Thusmost biota especially fungi are not explicitly covered by currentfire management policies

Recently concern has been expressed that management offire for vegetation may not provide the best outcomes forother biota (Clarke 2008 New et al 2010) In turn due to theinterconnectedness of fungi with the biota in general otherorganisms that rely on fungi could also be compromisedby knowledge deficiencies about their fungal partners Thepurpose of this review is to summarise the currentunderstanding of how fire affects fungi in Australianecosystems to identify knowledge gaps and to suggest stepsby which fungi and fire may be better managed After anoverview of the current literature on fungi and fire inAustralia and discussion of fungi that do or do not require fire(and the optimum fire frequency for fungi) we deal with fireand fungi in the framework of fungal trophic groups andsubstrates in order to highlight the functional roles of fungiand to suggest a context that will aid integration of fungi intoecosystem management

Fire and the fungal community in Australian ecosystems

Studies of the effect of fire on fungal communities in Australianecosystems are summarised in Table 1 The 30 field studiesidentified utilised a range of techniques but all compare thefungi present on sites with different times since fire Most studiesincluded recently burnt sites but in some cases (eg Ratkowskyand Gates 2008) the comparison was among sites of known firehistory that had not been burnt for several years to many decadesIn general long unburnt (commonly referred to as control) siteswere incorporated but some studies focussed only on recentlyburnt sites where the fire was less than 10 years ago (Theodorouand Bowen 1982 Warcup 1990 Bellgard et al 1994 Launonenet al 1999 Vernes and Haydon 2001 Vernes et al 2001 2004

Fungi and fire in Australian ecosystems Australian Journal of Botany 71

George 2008 Catcheside et al 2009 Claridge et al 2009aOrsquoBryan et al 2009 Ratkowsky and Gates 2009 Robinson2009) Nine studies (some involving multiple publications)incorporated a comparison of different fire regimes in relationto different frequencies of burning (Hilton et al 1989Bastias et al 2006a 2006b Brundrett et al 1996a 1996bTommerup et al 2000 Glen et al 2001 Vernes and Haydon2001 Vernes et al 2001 2004 Anderson et al 2007Osborn 2007 Campbell et al 2008 Artz et al 2009 OrsquoBryanet al 2009)

Most of the studies listed in Table 1 focussed on particulargroups of fungi Group distinctions were based on taxonomymorphology or trophic assemblage with most studies surveyingfruit-bodies of macrofungi (inclusive of both saprotrophs andectomycorrhizal species) Only one study involved lichens andtwo studies investigated the effect of fire on cryptogamicsoil crusts of which lichens are an important component Fewstudies attempted to identify microfungi and those that did eitherfocussed on a specific group such as the Trichocomaceae (McGeeet al 2006) or included soil microfungi as part of sampling of soilfungi across all taxonomic groups (Bastias et al 2006a) Nostudies dealt with microfungi of aboveground plant tissues suchas the highly diverse suite of leaf-inhabiting fungi of indigenousAustralian plants (May and Simpson 1997 Cheewangkoon et al2009)

The way that fungi were sampled varied considerably acrossthe studies included in Table 1 A few studies determined generalfungal biomass (eg Osborn 2007) or biomass of particulargroups such as truffle-like fungi (Vernes and Haydon 2001)estimated thenumberof ectomycorrhizal root tips (MalajczukandHingston 1981) or utilised hyphal-ingrowth bags to specificallytarget ectomycorrhizal fungi (Bastias et al 2006b) Isolationinto pure culture was utilised to study Trichocomaceae in bark(McGee et al 2006) and fungi in general as a component of thesoil microbial community (Theodorou and Bowen 1982)Isolation into culture is well known to have serious limitationsas a means of assessing the fungal community present in soil orother substrates because most fungi will not grow or areoutcompeted on standard media (Bridge and Spooner 2001)

Most surveys were based on recording fruit-bodies ofepigeous macrofungi (such as mushrooms and coral fungi) orhypogeous macrofungi (sequestrate or truffle-like fungi) Fruit-bodies do indicate active growth of underlying mycelium butparticularly for fleshy macrofungi they are often ephemeral andsporadic in occurrence Thus intensive sampling at times of peakproduction (such as autumn) over several years is required todetect a significant proportion of the species present at a siteFor ectomycorrhizal fungi it is also clear that there is a mismatchbetween the species detected on root tips and those present asfruit-bodies (Gardes and Bruns 1996) For example usingrestriction fragment length polymorphism (RFLP) analysisGlen et al (2001) detected many more species in Jarrah forestsoil than were recorded by fruit-body surveys

Molecular methods show great promise for characterisationof the fungal community in soil and other substrates Directisolation of DNA from the substrate can be utilised in tandemwith community profiling techniques such as denaturing gradientgel electrophoresis (DGGE) or terminal restriction fragmentlength polymorphism (T-RFLP) (Anderson and Cairney 2004)

Molecular sampling is particularly useful for soil where it candetect macrofungi not producing fruit-bodies at the time ofsampling as well as saprotrophic mycorrhizal and parasiticmicrofungi However as with culturing DNA sampling doesnot distinguish active mycelium from resting stages such asspores or sclerotia (Bridge and Spooner 2001) Sampling ofDNA from soil has been used to study the effects of fire onfungal communities in eucalypt forests in New South Wales andVictoria (Chen and Cairney 2002 Osborn 2007) and in a long-termprescribed burning experiment inEucalyptus pilularis forestat Peachester State Forest in southern Queensland (Bastias et al2006a 2006b Anderson et al 2007 Campbell et al 2008Artz et al 2009)

Assignment of fungi to trophic groups is important forinterpreting ecological roles While DNA primers specific forfungi can be utilised (eg Chen and Cairney 2002) it is notpossible to separate fungal DNA neatly into different trophicgroups by use of specific primers Although the trophic status offungi can vary considerably within higher taxa such as familiesand orders it is highly correlatedwith phylogeny and is generallyuniform within genera (Tedersoo et al 2009b) Thereforemost individual sequences can be assigned to trophic groups(Bastias et al 2006b) as can fungi sampled as fruit-bodiesIn addition the presence of particular trophic groups has beentargeted in soil samples by the use of functional molecularmarkers such as basidiomycete laccase genes putativelyinvolved in decomposition (Artz et al 2009) Other assaystargeted at particular fungal activity were phenol oxidaseactivity (Artz et al 2009) or stable isotope probing to assesscellulolytic activity of soil fungi (Bastias et al 2009)

Identification of fungi to species is one of themost challengingcomponents of fungal ecology and difficulties of identificationare compounded by the large number of species yet to be formallydescribed that are routinely encountered in fungal surveysTraditional morphological identification utilises microscopiccharacters and the differences between species are often subtleIn studies where fruit-bodies were sampled identificationof fungi was mostly on morphological characteristics Forsurveys of epigeal macrofungal fruit-bodies substantialinventories of species were accumulated in some studies suchas the 307 taxa observed by Gates et al (2005) in southernTasmania and the 322 taxa observed by Robinson and Tunsell(2007) in south-west Western Australia All taxa were identifiedto species level but in both studies only about half the taxa wereformally described the rest were assigned lsquotagrsquo or lsquofieldrsquo namesOther studies utilisingmorphological identification also includeda substantial number of lsquotagrsquo names

Molecular methods not only allow isolation of fungi theyalso facilitate identification of fungi Molecular communityprofiling techniques such as DGGE produce banding patternswhere each band is more or less unique at the species levelalleviating the necessity for identification of the fungi concerned(Anderson and Cairney 2004) To identify species the closestmatch for a particular sequencedDNAregion [such as the internaltranscribed spacer (ITS)] canbe comparedwith samples lodged inGenBank The sequencedDNAmay come from samples isolatedas DNA in the first place such as through DGGE and otherprofiling techniques or from conventional samples such as fruit-bodies or ectomycorrhizal roots tips In comparison to the several

72 Australian Journal of Botany S J M McMullan-Fisher et al

hundred species identified from some fruit-body surveys(Gates et al 2005 Robinson and Tunsell 2007) relatively fewtaxa have so far been identified to species level where molecularcommunity profiling has been used because only a selectionof the numerous bands present were selected for sequencing(Bastias et al 2006a)

Where molecular methods were utilised to identify fungifrom soil samples species were rarely able to be determinedAmong the ITS sequences generated byBastias et al (2006b) andChen and Cairney (2002) to identify RFLP types there werefew matches to named species above the 97 level that isindicative of species identity for this region in many groups offungi (Hughes et al 2009) Bastias et al (2006a) were not ableto identify any of 39 selected bands from DGGE profiles tospecies level on the basis of ITS sequences with some bandsmatched only at order or phylum level The inability to identifymany sequences from the studies listed in Table 1 to species iseither because the sequences with high identity in GenBank werefrom unidentified fungi in the first place or because the level ofmatching with sequences of known identity was low implyingthat reliably named sequences of most Australian soil fungi arenot yet available in GenBank

Many studies listed in Table 1 were unreplicated or whenthere was some replication of sites within treatments surveyswere limited to repeated collection of data at different timesafter a fire Replicated sampling tends to occur more withplanned experimental studies (Theodorou and Bowen 1982Warcup 1983 Bellgard et al 1994 Launonen et al 1999McGee et al 2006) compared with descriptive or ecologicalstudies Unreplicated studies can be regarded as indicative ofeffects but need to be backed up bywell replicated investigationsbefore their conclusions can be accepted Due to the variablenature of fire analysis of results is particularly difficult assufficient statistical power is hard to obtain and fire conditionsare hard to replicate The within-site variability may be so greatthat any differences among treatments may be hard to discernOnly nine of the Australian studies used plot-based replicatedsampling and attempted to identify the target fungi to specieslevel and all nine utilised counts of fruit-bodies (Hilton et al1989 Claridge and Barry 2000 Claridge et al 2000a 2000bMcMullan-Fisher et al 2002 Packham et al 2002Robinson andBougher 2003 Claridge and Trappe 2004 Kantvilas and Jarman2006 Robinson et al 2008 Ferguson et al 2009 OrsquoBryan et al2009)

Studies combining fungi and fire in Australia have beenconcentrated in Eucalyptus forests of south-eastern and south-western Australia (Table 1) There is a lack of data for othervegetation types notably grasslands arid areas and woodlandsForests dominated by Eucalyptus marginata (Jarrah) andE diversicolor (Karri) in south-west Western Australia havebeen relatively well studied and fungal communities have beeninvestigated in relation to fire using both morphologicaland molecular techniques (Malajczuk and Hingston 1981Hilton et al 1989 Tommerup et al 2000 Glen et al 2001Robinson and Bougher 2003 Robinson et al 2008) Anothercomparatively well studied ecosystem is wet sclerophyll forestwith E obliqua as a dominant or subdominant tree componentwith investigations of macrofungi (Packham et al 2002 Gateset al 2005 2009 Ratkowsky 2007 Ratkowsky andGates 2009)

lichens (Kantvilas and Jarman 2006) and the soil microbialcommunity including fungi (Theodorou and Bowen 1982)

In addition to the chronosequence studies that compare fungalcommunities on sites of different fire history (Table 1)inventories of fungi from individual sites of known time sincefire could be relevantwhen considering the effects offire on fungiExamples of such inventories come from repeated sampling forfruit-bodies of various groups ofmacrofungi onMtWellington inTasmania which was last burnt in 1967 (Ratkowsky and Gates2002 Gates and Ratkowsky 2004 2005 Trappe et al 2008)Such inventories have the potential to contribute to the pool ofknowledge about fungi in different vegetation types of particularfire histories but at present the information available is too limitedto make meaningful comparisons

The studies listed in Table 1 investigated widely differentfunctional and morphological groups of fungi and used a varietyof different survey techniques and methods for identification ofthe samples generated Fire history also varied among the studiesSuch diversity makes comparison of results difficult and similardifficulties in comparing studies were encountered by Cairneyand Bastias (2007) in their global review of the effects of fireon forest soil fungi Nevertheless most studies in Australianecosystems reported some consequence of fire on the fungalcommunity and these effects are elaborated on below First wediscuss fungi that occur after fire including considerationof both immediate post-fire (pyrophilous) fungi and alsothose that require long unburnt vegetation as well as theeffects of repeated fire on fungal communities Second we usethe framework of trophic roles and substrates to discuss effects offire on particular trophic groups such as parasites saprotrophsand mycorrhizal fungi and interactions between fire fungi andother organisms

Effects of time since fire

Pyrophilous fungi

A suite of pyrophilous fungi produce fruit-bodies on recentlyburnt sites Many of them are cosmopolitan and most typicallyfruit in the first and second years after fire often in large numbers(Petersen 1970 Warcup 1981 1990 Robinson 2001 Robinsonet al 2008 Claridge et al 2009a) Concomitantly on recentlyburnt sites fruit-bodies are not produced by most fungi typicalof mature vegetation Fire can also stimulate the soil microbialcommunity as indicated by colony counts in cultures of soilsamples In soil from dry sclerophyll woodland in SouthAustralia fungal colony counts decreased immediately afterfire but were significantly higher 2 months after fire in burntcompared with unburnt soil and remained so for at least 7 months(Theodorou and Bowen 1982) In other parts of the worlddepending on fire intensity microbial biomass may eitherdecrease or increase (Cairney and Bastias 2007)

The first and most conspicuous fungi observed after firein Australia are those that produce fruit-bodies fromsubterranean storage organs called sclerotia or pseudosclerotia(McMullan-Fisher et al 2002 George 2008 Robinson et al2008 Robinson 2009) Fruit-bodies of the pyrophilousbasidiomycetes Neolentinus dactyloides Laccocephalummylittae L tumulosum (Fig 1d) L sclerotinium (Fig 1f ) andan unnamed Laccocephalum species (Fig 1e) are common in


Tab

le1

Stud

iesof

fung

alcommun

itiesin

Australianecosystemsusingsitesof

know

nfire

history

Multip

lestud

ieson

thesamesitesaregroupedtogetherF

ireages

with

anasterisk

arethefrequencyforsitesthathadun

dergon

erepeated

burns

Vegetationtype

(dom

inantvegetation)

State

Sites(age

sincefireinyears)

Fun

gigrou

psstud

ied

Sam

plingandidentifi

catio

ntechniqu

esandor

estim

ationof

fungalactiv

ityReference

Fire0ndash12

mon

ths

Fire

gt1ndash10

years

Mature

Trophic

group

Morph

o-logical

group

Sam

ple

collection

techniqu

e

Identifi

catio

ntechnique

(metho

dor

scop

e)

Estim

ateof

fungalactiv

ity(m

etho

d)

Wetscleroph

yllforest(Eo

bliqua

andor

Ed

elegatensis)

Tas

ndashndash

+(25ndash30

gt1

00)

All

EM

FB

MOR(SP)

ndashPackh

ametal(20

02)

Wetscleroph

yllforest(Eo

bliqua)

Tas

++(3)

+(65ndash10

1)L

LFB

MOR(SP)

ndashKan

tvila

san

dJa

rman

(200

6)Wetscleroph

yllforest(Eo

bliqua)

Tas

++

+(73)

All

EM

FB

MOR(SP)

ndashGates

etal(20

09)

Wetscleroph

yllforest(Eo

bliqua)

Tas

++(2ndash3)

All

EM

FB

MOR(SP)

ndashRatkowskyan

dGates

(200

9)Wetscleroph

yllforest(Eo

bliqua)

Tas

ndash+(2ndash3)

+(70)

All

EM

FB

MOR(SP)

ndashGates

etal(20

05)

Wetscleroph

yllforest(Eo

bliqua)

Tas

ndashndash

+(73

10920

0ndash30

0)All

EM

FB

MOR(SP)

ndashRatko

wskyandGates

(200

8)G

ates

etal

(201

0a2

010b)

Wetscleroph

yllforest(Eregnans)

Tas

++(24

7)

+(57)

All

EM

FB

MOR(SP)

ndashMcM

ullan-Fisheretal

(200

2)Wetscleroph

yllforest(Eregnans)

Vic

+(2

mon

ths)

+(14

25mon

ths)

ndashM

(ECM)

SF

SO

MOR(M

G)

BAB

M(ERG)

Launo

nenetal(19

99)

Sclerop

hyllforest(Ed

iversicolor

regrow

thforest)

WA

++

+(17ndash25

)All

EM

FB

MOR(SP)

ndashRob

insonan

dBou

gher

(200

3)R

obinsonetal

(200

8)Sclerop

hyllforest(Em

arginata)

WA

++(6)

+(45)

MSF

SC

MOR(M

G)

RT

Malajczuk

andHingston

(198

1)Sclerop

hyllforest(Em

arginata)

WA

ndash+(2ndash4)

+(28ndash30

)All

EM

FB

MOR(SP)

BM

(WEI)

ndashTom

merup

etal(20

00)

Sclerop

hyllforest(Em

arginata)

WA

ndash+

+All

EM

FB

MOR(SP)

ndashHilton

etal(19

89)

Sclerop

hyllforest(Em

arginata

Corym

biacaloph

ylla)

WA

ndash+(6ndash7)

+(66)

M(ECM)

SQE

MFB

MOR(SP)

MOL(RFLP)

Glenetal(20

01)

Dry

sclerophyllforest(Em

aculata)

SA

++(23

)ndash

All

EM

(Disc)

FB

MOR(SP)

ndashWarcup(199

0)Sclerop

hyllwoo

dland(Ecladocalyx

Eb

axteri)

SA

+ndash

ndashAll

SQE

MFB

MOR(SP)

ndashGeorge(200

8)C

atcheside

(200

9)C

atchesideetal

(200

9)R

obinson

(200

9)Low

open

drysclerophyllwoo

dland

(Eb

axteriE

obliqua)

SA

+(12

7mon

ths)

+(12and

20mon

ths)

ndashS

SF

SOC

UMOR(M

G)

ndashTheod

orou

andBow

en(198

2)Dry

sclerophyllwoo

dland(Erossii

Em

acrorhyncha)

ACT

++(4)

+(21)

All

SQE

MFB

MOR(SP)

ndashTrapp

eetal(20

06)

Woodland(Ea

lbensCallitris

glau

cophylla)riparian

strips

(Eviminalis)grassy

orshrubb

ywoo

dland(Em

ellio

dora)

woo

dland(En

ortonii)gradingto

open

forest(Ed

alrympleana)

NSW

++

ndashAll

EM

FB

MOR(SP)

ndashClaridg

eetal(20

09a)


Sclerop

hyllshrubland(Ang

opho

rahispida)

NSW

++(8)

+M

(AM)

SF

SS

MOR(M

G)

ndashBellgardetal(19

94)

Dry

open

sclerophyllforests

(Eg

ummiferaS

yncarpia

glom

uliferaE

squ

amosa

EresiniferaE

haemastoma

Eb

auerana

Allo

casuarina

littoralisA

ngop

hora

hispida)

NSW

+(2

weeks)

+(81

0)+(25)

All

SF

SO

MOL(RFLP

DNA)

ndashChenandCairney

(200

2)

Sclerop

hyllforest(Ep

ilularis)and

mixed

eucalypt

forest(Eo

bliqua

Erub

ida

Erad

iata)

NSWV

ic

ndash+(31

0)

+(gt60

)All

SF

FBS

OMOR(M

G)

MOL(IS

RFLP

T-RFLP)

BM

(ERG)

Osborn(200

7)

Sclerop

hyllforest(Ep

ilularis)

Qld

ndash+(2

4)

+(31ndash34

)SM

(ECM)

SF

SOH

IMOL(D

GGE

RFLPT

-RFLP

DNAS

IR)

BM

(PLFA)

GLP

OIS

Bastia

setal(20

06a

2006b)A

ndersonetal

(200

7)C

ampb

elletal

(200

8)A

rtzetal(200

9)Sclerop

hyllforest(Ep

ilularis)

NSW

+(48h)

ndash+

SMI(Tric)

CU

MOR(SP)

ndashMcG

eeetal(20

06)

TropicalAllo

casuarinaforest

Eucalyptuswoo

dlandandtropical

rainforest

Qld

++(4ndash5)

ndashM

(ECM)

SQ

FBS

PMOR(SP)

BM

(WEI)

Vernesan

dHaydon

(200

1)V

ernesetal

(200

120

04)

Tropicalsavann

asNT

++

+M

(ECM

andAM)

SF

SOS

SMOR(M

G)

BA

Brund

rettetal(19

96a

1996b)

Tem

perategrassland(Themeda

australisP

oasieberiana)

NSW

ndash+(24

8)

+L

CS

FB

MOR(SP)

ndashOrsquoBryan

etal(20

09)

Tussock

grasslandandhummock

sedg

eland

Tas

+(3

mon

ths)

+L

LFB

MOR(SP)

ndashFergu

sonetal(20

09)

Alpinewoo

dland(Eucalyptus

pauciflora

Ed

alrympleana)tall

wetscleroph

yllforest(Eregnans

Efastig

ata)d

rysclerophyllforest

(Em

anniferaE

macrorhyncha)

Vicand

NSW

++(gt1ndash10

)+(11ndash20

21ndash30

gt30

)M

(ECM)

SQ

FB

MOR(SP)

ndashClaridg

ean

dBarry

(200

0)C

laridg

eetal

(200

0a2

000b)

Claridg

ean

dTrapp

e(200

4)Alpineheath

Tas

ndashndash

+(39

56)

All

EM

FB

MOR(SP)

ndashMcM

ullan-Fisheretal

(200

3)Mallee(Callitrispreissiisubsp

verrucosa)

NSW

++(34

7)

+(13

161

835

100

)L

CS

FB

MOR(SP)

ndashEldridg

eandBradstock

(199

4)

Referencesinbo

ldareforstud

iesthathave

replicated

sitesforallageclassesthoseinita

licsareforstud

ieswith

replicated

samples

with

inun

replicated

fireeventsA

bbreviationsT

rophicgroupAll=all

trophicgroups

except

lichensL

=lichensM

=mycorrhizalfungi(ECM

=ectomycorrhizasA

M=arbu

scular

mycorrhizas)andS=saprotroph

sMorph

olog

icalgrou

pCS=cryp

togamicsoilcrusts

EM

=epigeous

macrofungi(Disc=Discomycetes)MI=

microfungi(Tric=Trichocom

aceae)SF=soilfungiand

SQ=sequestratemacrofungiSam

plecollectiontechniqueCU=cultu

redFB=fruit-body

surveyH

I=myceliumfrom

hyph

alingrow

thbagsSC=ectomycorrhizalroottip

sfrom

soilsampleSO=bu

lksoilsampleSP=spores

from

scatsandSS=sporesfrom

soilFun

galidentificatio

ntechniqu

eMOL=identifi

catio

noftaxa

bymolecularmethods

(methodDGGE=denaturing

gradientgelelectrophoresisD

NA=DNAsequencing

andmatchingagainstsequencedatabasesRFLP=restrictionfragment

lengthpolymorphismSIR

=substrateinducedrespirationT-RFLP=term

inalrestrictionfragmentlengthpolymorphism)and

MOR=identifi

catio

noftaxa

from

morph

ology(scopeSP=mostly

identifi

edto

speciesleveltaxaMG=identifi

edtobroadmorph

ogroups)E

stim

ateof

fung

alactiv

ityB

A=Bioassayof

extentof

mycorrhizalcolonisatio

nof

seedlin

gsB

M=fungalbiom

ass(m

etho

dERG=ergo

sterol

analysisPLFA

=phospholipid-derived

fatty

acidsWEI=

weightoffruit-bodies)GL=genetic

diversity

oflaccasegenesIS

=isotopicanalysisPO=phenoloxidaseactiv

ityR

T=countsofectomycorrhizal

root

tips


bothwet anddry eucalypt forests across southernAustraliaThesespecies are well adapted to fire and can develop large fruit-bodiesas rapidly as 2 days after fire (Robinson 2001) Their mycelia

decay logs and buried wood and subterranean sclerotia orpseudosclerotia develop under or adjacent to host logs (Wills1983) It is not known how long it takes for these fungi to

(a) (b)

(c) (d )

(e ) (f )

Fig 1 (a) Anthracobia muelleri a saprotrophic ascomycete (b) Geopyxis carbonaria a mycorrhizal ascomycete and fire moss Funaria hygrometrica(c) Peziza tenacella a common mycorrhizal ascomycete after fire (d) Laccocephalum tumulosum fruit-bodies appear immediately after fire (e) Laccocephalumsp a saprotrophic basidiomycete showing pseudosclerotium ( f ) L sclerotinium showing sclerotia Photographs (a) (b) (e) D Catcheside (c) (d) ( f )R M Robinson


recolonise logs followingfire or how long sclerotia take tomatureto the stage where fruit-body development can occur

The majority of pyrophilous macrofungi produce fruit-bodiesin the first autumn following a spring or summer fire (Robinsonet al 2008) Several basidiomycetes fruit in the first year afterfire including agaricoid (mushroom-like) decomposer fungi inthe genera Coprinus (Fig 2a) Pholiota and Psathyrella (Gateset al 2005 Robinson and Tunsell 2007 Robinson et al 2008Catcheside et al 2009 Claridge et al 2009a) Most pyrophilousfungi however appear to be ascomycetes (Petersen 1970Warcup 1990) of which some are mycorrhizal (Warcup 1990)Species of Anthracobia including A muelleri (Fig 1a) andA melaloma and other ascomycetes produce abundant fruit-bodies over extensive areas and their mycelial mats may beimportant in minimising soil erosion following fires byaggregating soil particles (Claridge et al 2009a)

A post-fire flush of ascomycetes is a global phenomenon(El-Abyad and Webster 1968a 1968b Petersen 1970 1971Wicklow 1975 Zak 1992 Cairney and Bastias 2007) and severalAustralian studies have reported an abundance of ascomycetefruit-bodies (Fig 1andashc) associated with recently burnt sites(Warcup 1981 1990 Gates et al 2005 Robinson et al 2008Catcheside 2009 Catcheside et al 2009) The causes have beenwidely investigated and include decreased competition fromantagonistic soilborne taxa allowing post-fire germination ofspores heat stimulation of spore germination and tolerance topost-fire soil conditions such as increased pH (El-Abyad andWebster 1968a 1968b Petersen 1970 Wicklow 1975)

A direct and immediate effect of fire is soil heating whichcan potentially kill mycelia and spores of fungi (Pattinson et al1999) However certain fungi are adapted to being heated Forexample some species of Australian ascomycetes that producefruit-bodies immediately after fire have been shown to germinatefrom resting spores in heat-treated soil (Warcup and Baker1963) Similarly various Trichocomaceae (including speciesof Aspergillus Eupenicillium and Penicillium) were recoveredfrom heat-treated and burnt bark from a variety of native plantspecies (McGee et al 2006) as well as from heat-treated soil(Warcup and Baker 1963) Many pyrophilous fungi appear to betotally dependent on fire to stimulate spore germination andmycelial growth but some grow and fruit readily on both burntand unburnt sites (Robinson et al 2008)

Fungi that require long unburnt vegetation

In the northern hemisphere there are several fungi many ofwhich are included in Rarity Endangerment and Distributionlists whose fruit-bodies are only associated with large welldecayed logs in long undisturbed sites (Ing 1993 Berg et al1994 Odor et al 2006) These fungi are probably not themselvessensitive to fire per se but are associated with substrates ormicroclimate that are most common in areas which have notbeen disturbed for a long time In dry eucalypt forests ofAustralia large well decayed logs are more likely to beconsumed by fire than undecayed logs (Hollis et al 2008) InVictoria fruit-bodies of Hypocreopsis amplectens (Fig 2c d)have been found predominantly in long unburnt over-maturestands of Heath tea-tree Leptospermum myrsinoides (Johnstonet al 2007) The rare lichen Roccellinastrum flavescens is found

exclusively on leaves ofArthrotaxis cupressoides a tree found inlong unburnt areas in Central Tasmania (Kantvilas 1990 2000)Some species in the ectomycorrhizal genus Russula were foundmore frequently in long unburnt Jarrah forests than in sites with aprescribed burn frequency of 6ndash8 years (Hilton et al 1989 Glenet al 1998 2001)

Cool temperate rainforests in Tasmania and Victoria supportlarge and diverse communities of macrofungi (Fuhrer andRobinson 1992 T W May pers obs) and lichens (Kantvilasand Jarman 1993 Kantvilas 2000 Morley and Gibson 2004)Many are restricted to rainforest either because they areectomycorrhizal fungi strictly associated with the dominantNothofagus cunninghamii (Fuhrer and Robinson 1992) orbecause they apparently prefer the cooler and moistermicroclimate generated by closed canopies Thus many fungiappear to rely on the lack of fire that allows rainforest to flourishIn wet eucalypt forest mature (100ndash200 years) and over-mature(200ndash400 years) stands appear to be important for maintainingbiodiversity particularly wood-decay fungi that rely on largewell rotted logs (Wardlaw et al 2009) Fruit-body surveys inmature longunburnt (~80years)Eucalyptus obliqua forest on theFleurieu Peninsula in South Australia yielded an unusually highdiversity of macrofungi (Catcheside and Catcheside 2008) InE obliqua forest in Tasmania macrofungal species assemblagesdiffered between mature and 25ndash30-year-old regrowth forestregenerated following fire (Packham et al 2002) and betweenvery long unburnt plots and those burnt in 1934 and 1898 (Gateset al 2010a 2010b) In the latter study the distinctiveness of thelitter assemblage on sites of different age since fire was not sopronounced as for the soil and wood fungi Overall these studiessuggest that long unburnt wet eucalypt forests are importanthabitat for the fruiting ofmacrofungi Thedegree towhich speciesthat preferentially produce fruit-bodies in long unburnt forest arepresent in recently burnt forest as vegetative mycelium or restingstages is unknown

Effects of repeated fire

Despite the increasing use of prescribed burning acrossAustraliathere is limited information on how fungi are affected by repeatedburning in Australian ecosystems The effects of repeated fire onfungi are difficult to test experimentally because the post-firesuccession may extend over several or more years Macrofungalcommunities in some eucalypt forests differ each year after firefor at least 5ndash7 years (McMullan-Fisher et al 2002 Robinsonet al 2008) and little is known about the consequences ofinterrupting this succession before communities recover totheir pre-fire composition Ideally sites of different firehistory including long unburnt controls need to be burntsimultaneously and succession followed on each site to assesswhether successional communities differ between treatments(Wittkuhn et al 2010) Sites with different repeated fireregimes will have different species compositions at the end oftheir respective cycles but the key question is are the changespermanent

Results from the few studies on fungi and fire regimes are notconsistent In mixed eucalypt forests in Victoria repeated low-intensity prescribed fire of 3- and 10-year cycles at differentseasons (spring or autumn) had little effect on the richness and


diversity of fungal communities as measured by T-RFLPprofiling nor on diversity of fruit-body morphotypes andtrophic groups (Osborn 2007) In E pilularis forests in New

South Wales ergosterol concentrations (a chemical measure ofliving fungal biomass) in the topsoil (0ndash5 cm) from frequentlyburnt (every 3 years) sites were on average 35 lower than in

(a) (b)

(d )(c )

(e ) (f)

Fig 2 (a) Coprinus angulatus a saprotrophic basidiomycete fruiting in the first year after fire (b) Pycnoporus coccineus a common saprotroph on fallen andstanding timber (c d) Hypocreopsis amplectens growing on fallen branches of senescing Leptospermum in long unburnt woodland (e) Daldinia sp a semi-parasitic ascomycete fruiting on Hakea in the second year after fire ( f ) Mesophellia trabalis a truffle-like ectomycorrhizal fungus excavated from soil at arecently burnt site this species also fruits on unburnt sites Photographs (a) (b) (e) D Catcheside (c) (d ) T W May ( f ) R M Robinson


soil from long unburnt (45 years) sites but because ofconsiderable within-treatment variability the differences werenot significant (Osborn 2007) In E marginata forest in south-westernAustralia in terms of fruit-body yield a long unburnt sitehad fewer decomposer and more mycorrhizal fungi than a sitewhich was burnt every 10 years (Tommerup et al 2000) and longunburnt sites also had more ectomycorrhzal root tips (Glen et al1999) In the same forest type in the only study where alltreatments (including controls) were sampled at the same time(4 and 5 years) since fire the composition of macrofungalcommunities on sites with repeated fire at short (5 yrs)intervals differed from those on sites that had long (10 years)intervals between fire but neither differed significantly fromcommunities on sites with moderate (6ndash9-year) fire intervals(Wittkuhn et al 2010)

In E pilularis forest at Peachester State Forest in Queenslandseveral studies have utilised molecular characterisation ofsoilborne fungal communities and determination ofbasidiomycete enzyme activity and cellulose utilisation withrespect to 2- and 4-yearly repeated fire regimes and unburnt(since 1972) controls (Bastias et al 2006a 2006b 2009Anderson et al 2007 Campbell et al 2008 Artz et al 2009)While species richness was similar across the treatmentscommunity composition was altered by repeated burning andthe effectwas greatest in the biennially burnt treatment (Andersonet al 2007) Similarly the communityof ectomycorrhizal fungi insamples from the upper 10 cm of soil from burnt plots differedfrom long unburnt plots but at the 10ndash20-cm depth there was nodifference (Bastias et al 2006a) Biomass of both fungi andbacteria were reduced by 50 on plots burnt at 2-year intervalscompared with the 4-year interval and long unburnt plots(Campbell et al 2008) and there were fewer active cellulolyticfungi in the biennially burnt plots (Bastias et al 2009) Phenoloxidase activity due to saprotrophic fungi also decreased but notsignificantly with increased fire frequency but the community ofbasidiomycete laccase genes from frequently burnt plots wasmore diverse and more even than that from the long unburntplots (Artz et al 2009) Observations from lsquogreen islandsrsquowithinsites burnt every two years were included by Bastias et al(2006a 2006b) to demonstrate that the most recent fire on the2-year sites was patchy and of low intensity and thus theshort-term effect of fire was similar on both 2- and 4-year sitesand was not confounding the long-term effects However aswith most other studies discussed in this section the post-firefungal succession was not followed throughout its course onall treatments (including unburnt controls) Ideally intensivedocumentation of fungal communities after fire should becomplemented by longer-term studies of succession across alltreatments

For biological soil crusts which include lichens responses tofire frequency varywith vegetation type Inmallee woodlandfirefrequencies of less than 10 years alter soil crusts to favour algal-dominated crusts and lichen composition was highest when timesince fire was between 13 and 35 years (Eldridge and Bradstock1994) In contrast in temperate grassland lichen cover was foundto be greatest on sites burnt every second year (OrsquoBryan et al2009)

Fire frequency affects vegetation composition through effectson propagules For plants species that rely on seed production for

post-fire recruitment will be eliminated when the timebetween fires is shorter than the time required for recruitmentmaturation and replacement of the seed store before the next fire(Ooi et al 2006 Yates et al 2008Watson et al 2009) Recoveryof fungal biomass after fire requires either regeneration ofexisting communities from surviving spores or mycelia orrecolonisation from surrounding unburnt areas (Bruns 1995)Indirect effects of fire on soil such as loss of nutrients througherosion and leaching change in water repellence greaterabsorption of heat by blackened soil surfaces and loss of theshading cover of vegetation (Gochenaur 1981) will also affectrecolonisation by soil fungi

Little is known about the dynamics of the lsquobankrsquo or store offungal propagules which includes spores hyphal fragmentsand resting structure such as sclerotia Fungal propagules suchas spores may survive independently of hosts Data on thelongevity of ascomycete spores in soil are available (Warcupand Baker 1963 El-Abyad and Webster 1968a Warcup 1981)and research on spore banks inNorthAmerican conifer-grasslandcommunities suggests that spores of the truffle-like Rhizopogoncan persist for up to 20ndash30 years (Kjoller and Bruns 2003 Izzoet al 2006) Some fungimay also survivewithin or in associationwith host tissue such as in mycorrhizas Hyphae that remainassociated with roots may survive better than those that are not(Pattinson et al 1999) Propagules at greater depths will be lessaffected by heating from fire but the relative colonising potentialof propagules at different depths is unknown Some macrofungidevelop underground storage organs that allow fruiting after firebutwe know little about the time they take to develop andmatureor how long they persist before production of fruit-bodies istriggered by fire (Robinson et al 2008)

Other factors that could influence recolonisation by fungi afterfire include the size of disturbance ndash a large burnt patchmaymakerecolonisation from spores more difficult compared with a smallpatch The age of the vegetation (and the corresponding stage ofsuccession in the fungal community) around a burnt site may alsobe critical as could be the intensity with which a patch burns Inaddition recolonisation by fungi that require vectors such asmammals may be affected by changes in the abundance andmovements of the vectors All these factors remain to be testedexperimentally

The effect of fire on different trophic and substrategroups of fungi

The variation in physiology and ecological roles among fungiand the range of substrates and mutualistic partners mean thatdifferent trophic and substrate or host groups merit separateconsideration when considering the effects of fire on fungiThis is because fire may have quite different effects on thegenesis and consumption of substrates for fungi on the onehand and on the growth and survival of mutualistic partners onthe other hand Consideration of the functional significance toother biota of fire-mediated changes to fungal communities willalso be facilitated by discriminating the different trophicfunctions of fungi (Cairney and Bastias 2007)

Saprotrophic fungi

Saprotrophic fungi decompose dead organic matter includingdeadwood on live trees coarsewoody debris (CWD) and litter on


the forest floor as well as some materials of animal originThe process of decay and decomposition of wood and othersubstrates is facilitated by the succession of specialist fungiSome saprotrophic fungi are host specific while others canutilise a wide range of hosts In addition many are substratespecific and prefer wood of a particular size or stage of decay(Boddy 1984 Spooner 1987)

On living standing trees high intensity fires cause scars thatact as entry points for decay-causing and pathogenic fungi(Parmenter 1977 Abbott and Loneragan 1983 McCaw 1983)Some decay fungi are important in the development of habitatfor other organisms including invertebrates and small reptilesand for generating nesting hollows for birds and animals (Kileand Johnson 2000 Hopkins et al 2005) Alternatively trees maybecome stressed or weakened making them more susceptibleto attack from root and canker pathogens (Parmenter 1977) orthey may be killed creating new habitat for wood decay fungi(Penttilauml and Kotiranta 1996) either as standing dead trees orfallen log and branch material In Finland 65 of polypore andcorticioid fungi recorded on dead standing spruce trees before ahigh intensity fire were not found after the fire with speciescolonising dead trees in an advanced stage of decay being themost affected However 46 of species recorded post-fire werenot recorded before the fire because the disturbance reducedcompetition or provided new substrates in the form of freshlyfallen trees and branches for the establishment of early colonisers(Penttilauml and Kotiranta 1996)

Consideration of CWD in forest management has recentlybeen recognised as important because of its role in long-termnutrient cycling maintenance of biodiversity carbon storagemoisture retention tree health forest structure and habitat forfauna and fungi (Grove et al 2002) Saprotrophic fungi areimportant in CWD decomposition (Grove and Meggs 2003Mackensen et al 2003) which involves a succession of fungalspecies (Dix andWebster 1995Boddy2001)Many saprotrophicfungi that colonise CWD have specific associations with sizewood type decomposition stage or moisture level (Johnston2001 McMullan-Fisher et al 2002 Grove and Meggs 2003Berg and Laskowski 2006 Wardlaw et al 2009 Gates et al2010b)

Timber harvesting silviculture practices and firewoodcollection impact on the amount and quality of CWD which inturn affects species and communities dependent on large diameterlogs including saproxylic (wood-inhabiting) invertebrates andfungi (Huston 1996 Jonsson et al 2005 Wardlaw et al 2009)Fire can reduce the total amount of CWD and alter itscharacteristics (Robinson and Bougher 2003 Hollis et al2008) Alternatively fire can increase heterogeneity of CWDby altering decay processes changing microclimates and bygenerally increasing the diversity of species which contributeto the woody debris pool (Grove et al 2002) Fire is a key cyclicdisturbance that kills trees thus contributing to the production ofCWD in forests

Intensefiresmay reduce the amountofCWDon the forestfloorand impact significantly on species of fungi that colonise anddecay dead wood (Penttilauml and Kotiranta 1996 Robinson et al2008) In regrowth Karri forest in Western Australia the numberof species fruiting on wood was shown to be significantly lowerfor 3 years following an intense wildfire but after 5 years species

richness had increased and was higher than on comparableunburnt sites (Robinson et al 2008) Other studies haveshown similarly decreased species richness of fruit-bodies ofwood-inhabiting macrofungi on more recently (2ndash5 years) burntsites (McMullan-Fisher et al 2002 Gates et al 2005)

Litter is composed of dead leaves and fine woody materialfrom plants The time since last fire is particularly relevant whenstudying fungal communities as litter is generally consumed byfire Short-term loss of litter-decomposing macrofungi afterfire has been reported for several Australian ecosystems(McMullan-Fisher et al 2002 Robinson et al 2008) andelsewhere (Berg and Laskowski 2006) In E marginata andE diversicolor forests in Western Australia fruit-bodies oflitter-dwelling fungi such as Mycena and Marasmius appear assoon as 2 years after fire while other genera such as Entolomadid not fruit until an organic layer had formed below the litterlayer (Robinson et al 2008) Litter-dependent fungi inE regnansforest in Victoria did not fruit until after the canopy hadclosed 7 years after fire (McMullan-Fisher et al 2002) Suchdifferences will be related to the rate of accumulation of litterafter fire which depends on fire intensity and varies significantlyacross forest types and ages (Ashton 1975 McCaw et al 19962002 Department of Sustainability and Environment 2003)Production of fruit-bodies is also influenced by microclimateseason canopy development tree health and other factors Firesreset or interrupt the succession of litterndashdecomposing fungi andstudies from a range of eucalypt forests (McMullan-Fisher et al2002 Gates et al 2005 Robinson et al 2008) suggest that it maytake 5ndash10 years for litter-dwelling macrofungal communitiesto recover and produce fruit-bodies present in the pre-firecommunity

Litter saprotrophs may be influenced indirectly by fire due torestricted associationswith plant hostswhose presence is reducedor enhanced by fire For example onMtWellington in Tasmaniafruit-bodies of an unidentified discomycete were restricted tolitter produced by Orites acicularis a plant whose cover wassignificantly more abundant on sites burnt 56 years previouslycomparedwith sites burnt 39 years previously (McMullan-Fisheret al 2003) Conversely there may well be fungi restricted toplants that are favoured by fire

All above-mentioned studies in this section utilised samplingof fruit-bodies from wood or litter An alternate approach is tomeasure decomposition activity through substrate or gene assaysBastias et al (2009) amended soilwith 13C labelled cellulose andcompared soil fungal communities that did and did not utilisecellulose by stable isotope probing in combination withDGGE In a comparison of long unburnt and regularly burnt(every 2 years) sites in E pilularis forest they conclude that thediversity of cellulolytic fungi was reduced by repeated burningOn the same sites Artz et al (2009) found that repeated burningcaused a significant shift in the basidiomycete laccase-encodinggene pool Laccases are involved in the breakdown of woodalthough they may also be produced by some ectomycorrhizalfungi

Parasitic and endophytic fungi

Parasitic fungi most of which are microfungi are highly diversebecause they are often host-specific at the species genus or family


level Despite this high diversity little is known of the ecology ofnative parastiic fungi especially in relation tofire There is a post-fire (3ndash5 years) increase in fruiting of semi-parasitic ascomycetessuch as Daldinia spp (Fig 2e) and Hypoxylon spp (Gates et al2005 Robinson et al 2008) These fungi live within the woodof healthy understorey trees and shrubs and rapidly formmacroscopically visible fruit-bodies after the host plant hasbeen weakened or killed by fire (Robinson et al 2008) Forthe numerous parasitic microfungi that cause cankers and leaf-spots on native plants in Australia there is no informationavailable about the effect of fire Many healthy plants containendophytic fungi some of which may become parasites orsaprotrophs when the plants become weakened by stress ordie Again information on the response of such species to fireis lacking

Information on interactions between parasitic fungi and firein Australian ecosystems is mostly limited to studies aboutcontrol of two high profile root pathogens In an attempt tocontrol dieback disease caused by Cinnamon fungusPhytophthora cinnamomii (a fungoid member of theChromista) high intensity fire was successfully trialled inJarrah forest in Western Australia (Shea et al 1979) Thespread of the pathogen was reduced through promotion of aresistant leguminous understorey High intensity fire may alsobe detrimental to the root pathogen Armillaria luteobubalinaas fire has the potential to destroy the outer sapwood ofstumps and to burn tree buttress and lateral roots on which thefungus would normally survive (Kile 1980 1981) Ironically theincreased accumulation of fuel caused by such parasitic fungimay promote fire (Robinson and Bougher 2003)

Mycorrhizal fungi

Mycorrhizal fungi associate with the fine roots and occasionallywith other underground structures of plants and facilitateexchange of nutrients between plant and fungus (van derHeijden and Sanders 2002 Brundrett 2004 Cairney 2005)They can also help to protect plants against some pathogens(Zak 1964 Marx 1972) and increase tolerance to environmentalstress such as drought (Tommerup and Bougher 2000)Several different types of mycorrhizas are known includingectomycorrhizas (ECM) arbuscular mycorrhizas (AMformerly known as vesicular arbuscular mycorrhizas VAM)ericoid mycorrhizas and orchid mycorrhizas each associationformed by particular groups of fungi and plants and eachwith a characteristic structure Apart from a few familiesnotably the Proteaceae most vascular plants and someliverworts form mycorrhizas of one sort or another Surveysof a variety of Australian ecosystems with Eucalyptus orAngophora overstoreys detected mycorrhizas in 66ndash96 ofthe plant species present with AM being the most commontype (Brundrett et al 1996a May and Simpson 1997)

Arbuscular mycorrhizal fungi are probably the mostwidespread and common mutualistic fungi Their responses tofire have been shown to be quite variable but fire usually impactsnegatively (Hart et al 2005 Cairney and Bastias 2007) In anopen sclerophyll shrubland in New South Wales spores of AMdeclined in abundance immediately after fire although no long-term effect on infectivity and spore abundance was recorded

(Bellgard et al 1994) In tropical savannah woodlands in theNorthern Territory inocula of fungi that formAMdid not seem tobe affected by fire intensity (Brundrett et al 1996a 1996b)Pattinson et al (1999) suggest that it is the loss of the mycelialnetwork after fire rather than modifications to inoculum potentialwhichdrives post-fire reductions inAMGiven that early seedlingdevelopment may well depend on arbuscular mycorrhizal fungi(Adjoud-Sadadou and Halli-Hargas 2000) such disruption to themycelia network may be important in the development of thepost-fire plant community (Bellgard et al 1994 Pattinson et al1999)

Ectomycorrhizas are common in eucalypt forests andwoodlands (Dell 2002) Ectomycorrhizal roots predominate inthe top 10ndash20 cm of soil and in leaf litter (Bastias et al 2006a)and can be impacted significantly at least in the short termby prescribed burning (Chen and Cairney 2002) Several studieshave noted a predominance of fruit-bodies formed byectomycorrhizal fungi in mature forest compared with recentlyburnt (2ndash3 years) sites (Glen 2002 Gates et al 2005) Onceforests have matured the species richness of macrofungi thatform ECM is similar in forests of different ages althoughindividual fungal species may favour forest stands of aparticular age (Packham et al 2002) Furthermore preferencesfor host plants by ectomycorrhizal fungi may affect plantcommunities (Allen et al 1995) For example host generalistectomycorrhizal fungi facilitate seedling establishment inlate succession forests but fire-dependent tree species such asPomaderris apetala and E regnans may competitively excludeeach other through the low compatibility of their respectiveectomycorrhizal fungi (Tedersoo et al 2008)

In Jarrah forest the number of mycorrhizal roots on trees wasdramatically reduced following removal of litter and soil organiclayers by fire (Reddell and Malajczuk 1984) and subsequentrecovery was related to time since fire and litter accumulation(Malajczuk and Hingston 1981) In E pilularis forest thecommunity of ECM in the upper soil from burnt plots differedfrom long unburnt plots (Bastias et al 2006a) Seedlings ofE maculata grown in heat-treated or untreated soils all hadabundant mycorrhizal associations but those grown in heat-treated soils had lower diversity and different types ofmycorrhizas (Warcup 1983) Similarly the frequency of ECMand the growth of E regnans seedlings were greater in burntblack soilwhen comparedwith unburnt soil whichwas attributedto changes in soil nutrition and the presence of differentectomycorrhizal fungi (Launonen et al 1999) In tropicalsavannah woodlands in the Northern Territory for sites thathad hot annual fires ectomycorrhizal fungi were restricted toinfrequent patches in the surface horizon and in unburnt sitesthe inocula of ectomycorrhizal fungi were more frequent(Brundrett et al 1996a 1996b)

Mycelia of most ectomycorrhizal fungi grow in the upperorganic layer of the soil However CWD may house a range ofectomycorrhizal fungal mycelia particularly when associatedwith seedlings of trees such as Nothofagus cunninghamii(Tedersoo et al 2009a) In addition some ectomycorrhizalfungi in the Thelephoraceae produce their fruit-bodies onCWD (Tedersoo et al 2003) This suite of ectomycorrhizalfungi will be particularly sensitive to changes andor loss ofCWD after fire


Orchids are particularly dependent on their fungal symbiontsfor survival (Rasmussen 1995 Smith and Read 2002 Dearnaleyand Le Brocque 2006) Fire stimulates the flowering of someterrestrial orchids such that declining populations may needregular disturbance by fire to maintain their on-going viabilityFor other terrestrial orchids fires that occur too frequently have anegative impact on populations presumably due to the effect offire on fungal symbionts as well as on the host plants (Brundrett2007) High fire frequency has been shown to reduce the numberof epiphytic orchids in tropical savannah both directly andindirectly due to decreased numbers of host trees on burntsites (Cook 1991) Many orchid species are listed as beinglsquothreatenedrsquo and altered fire regimes are considered to be oneof the causes for this status An understanding of the ecology ofthe fungal symbionts of orchids has an important role inpromoting orchid conservation (Brundrett 2006) Ericaceaewhich are partners in ericoid mycorrhizas (Cairney and Burke1998 Chambers et al 2008) are common plants in Australianwoodlands and heathlands but there is no information on theresponse of their mycorrhizal fungi to fire

Lichenised fungi

Lichenised fungi often have strong associations with particularsubstrates and habitats (Brodo 1973 Brodo et al 2001) Whenfires modify microclimatic and substrate conditions lichencommunities are often greatly altered particularly as fewlichens survive fire and are slow to recover (Stevens 1997)Lichens are so sensitive to changes in vegetation occurringover time since fire that they have been used as bio-indicatorsfor determining the age of cerrado vegetation in central Brazil(Mistry 1998) However where knowledge about distributionand ecology is limited care needs to be taken when assigningindicator species For example two lichenswhichwere thought tobe indicators of old growth wet sclerophyll forest in Tasmania(Kantvilas and Jarman 2004) were later found to be common onrecently burnt sites (Kantvilas and Jarman 2006) In the latterstudy which surveyedwood- and tree-dwelling lichens the post-fire lichen community was dominated by common cosmopolitanspecies and drier climate specialists and species changes wereattributed to the different microclimate and habitat characteristicsafter fire (Kantvilas and Jarman 2006) Several rare lichens arelimited to rainforests where fire has not been present for100ndash500 years (Kantvilas and Jarman 1993 Rogers 1995Kantvilas 2000 Morley and Gibson 2004)

Soil lichens are a component of biological soil crusts that areparticularly important in arid and semiarid ecosystems becauseof their role in preventing soil erosion (Eldridge 2003) Soil cruststend to be damaged by extreme disturbances such as highintensity fires (Eldridge and Bradstock 1994 Eldridge 1996Eldridge and Tozer 1997) but local species richness may bemaintained with lower impact disturbances (Eldridge et al2000 2006 OrsquoBryan et al 2009) In Tasmanian tussockgrassland cover and abundance of soil cryptogams in generalwere found to remainhighafter a fuel reductionfire (Fergusonet al2009) In contrast soil crusts in the mallee region were dominatedby algae on sites burnt less than 10 years previously while lichencover and compositionwas highest 13ndash35years afterfire (Eldridgeand Bradstock 1994) Lichen species and communities are rarely

included in studies onfire ecology inAustralia and further researchon their fire responses is required across a range of ecosystems

Fire and fungal-faunal interactions

Fungi-invertebrate interactions

Despite the prevalence of larvae and adults of many invertebratesutilising macrofungal fruit-bodies for habitat and food there isonly limited information about the interactions of invertebrateswith fungi and the effect that fire may have on these relationships(Wardle et al 2004) Frequent fire has recently been shown todisrupt the nature of fungal-invertebrate interactions in leaf litterleading to substantial changes in rates of decomposition (Brennanet al 2009) Belowground larger organisms such as earthwormsmites and collembola (springtails) are likely to be involved intransport of spores (Brown 1995 Dighton et al 1997 Dighton2003 Dromph 2003) Fire can reduce the abundance of theseand other soil- and litter-dwelling invertebrates (Neumann 1991Neumann and Tolhurst 1991 Collett et al 1993 York 1999)Grazing of soil-borne mycelia by invertebrates includingcollembola mites and nematodes can influence fungal biomassand community composition (Dighton 2003) This could haveflow-on effects on leaf litter decomposition and the efficiency ofmycorrhizas to facilitate nutrient uptake of host plants (Hanski1989 Shaw 1992 Brennan et al 2009) Preferential grazingby collembola can affect interactions between fungi thatform AM and saprotrophic fungi (Tiunov and Scheu 2005) Asingle low intensity fire was shown to alter the abundanceand composition of collembolan communities (Greenslade1997) and long-term frequent burning can reduce collembolannumbers bymore than half (York 1999) These changesmay alternutrient availability and in turn alter plant growth and vigour andcommunity structure

Fungindashvertebrate interactions

The interactions of fire fungi and mycophagous animals arecomplex Many native Australian animals consume fruit-bodiesof hypogeal truffle-like and some epigeal fungi Spores aresubsequently dispersed in scats sometimes a considerabledistance away where they germinate and form mycorrhizaswith trees or shrubs (Claridge and May 1994 Blaney 1996Johnson 1996 Vernes 2009 Vernes and Dunn 2009) As aresult of digging for hypogeal fruit-bodies soil aeration andwater incursion are enhanced (Garkaklis et al 2000 2003)and consequent changes to soil surface topography assist seedsettlement andestablishment It hasbeen suggested that the loss orreduction of mycophagous animal populations may have adeleterious impact upon the long-term health viability anddiversity of truffle-like fungi and consequently on soilstructure and nutrient cycling and eventually mycorrhizal plantcommunities (Claridge 2002) although exotic rodents such asRattus rattus may also disperse fungal spores (Vernes andMcGrath 2009)

The focus of considerable research has been on mitigatingthe effects of fire on mycophagous animals and their habitat(Catling 1991) However there are still large gaps in theunderstanding of interactions among mammals fungi fireevents and vegetation development In the only comprehensivelongitudinal study of the ecology of truffle-like fungi (Claridge


et al 2000a 2000b) fruiting of some individual species wasshown to be influenced by many factors including the time sincefire climatic variables such as temperature and moisture levelstopographic position geology soil fertility depth of litterdiversity of mycorrhizal hosts and abundance of mycophagousanimals Two taxa Cortinarius globuliformis and Mesophelliatrabalis (Fig 2f ) decreased in occurrence with increasing timesince fire (Claridge et al 2000a) However subsequent studiesshowed that C globuliformis was dominant on unburnt siteswhereasM trabalis appeared on both burnt and unburnt sites andfruit-body production of both species was influenced by manyother environmental factors (Claridge and Trappe 2004 Claridgeet al 2009b)

In regard to the community of truffle-like fungi studies in theAustralian Capital Territory and New SouthWales demonstratedthat prescribed burning decreased the overall diversity andabundance of fruit-bodies of truffle-like fungi (Claridge et al2000a Trappe et al 2006) Conversely several Tasmanianand Queensland studies suggest that prescribed burningmay stimulate the fruiting of some species of truffle-like fungi(Taylor 1992 Johnson 1994 1997 Vernes et al 2004)Comparison and interpretation of studies relating fire andtruffle-like fungi should be undertaken carefully as surveymethods can differ greatly and fungal communities should notbe assumed to be similar in different ecosystems (Claridge andTrappe 2004 Trappe et al 2005 2006)

Mycophagous macropods and rodents are likely to be criticalfor dispersal of fungal spores into disturbed habitats as wellas across the mosaics of vegetation types and ages that exist inmany Australian landscapes (Vernes and Trappe 2007 Vernesand Dunn 2009 Vernes andMcGrath 2009) Investigation of theforaging habits of mycophagous animals before and after fireshowed that they foraged preferentially on burnt ground andfrequently moved between burnt and adjacent unburnt habitat(Johnson 1994 1996 Vernes and Haydon 2001 Vernes andTrappe 2007) Exclusion of small animals from plots inQueensland rainforest resulted in lower seedling colonisationand an altered community composition of AM (Gehring et al2002) Comparable studies would be instructive to determine theresponse of truffle-like fungi in fire-prone forests whenmycophagous mammals are excluded

Future directions for research and managementin relation to fire and fungi

Fungi are clearly relevant to research programs on fire inecosystems because of their direct roles and interactions withother biota Providing clear management recommendations iscurrently hampered by the lack of comparability among existingstudies and the many gaps according to geography habitattype and ecosystem (especially for grasslands arid and alpineenvironments and northern Australian savannahs) There is alsoa lack of information for the full range of fungal taxonomicand trophic groups It is not practicable to sample all fungi inevery ecological study However it would assist to havecomprehensive data on the taxonomy biology and ecology ofselected groups of fungi representative of phylogeny and trophicmode as candidates for surveys As well as a sound taxonomicunderpinning autecological data for such selectedgroups suchas

in relation to life history characteristics (eg size of individualslongevity of spores and mycelia recolonisation strategies) andhost habitat and substrate specificity would assist greatly ininterpreting observations of fungi and fire Further practicalchallenges for studies of fire and fungi are the inclusion offungi in monitoring programs and the integration of molecularand morphological data

Fungi in monitoring programs

Greater understanding of the effect of fire on fungi relies ongeneration of substantial datasets Ideally fungi should beintegrated into established monitoring and survey projects toachieve this At present the Department of Environment andConservation (DEC) in Western Australia is the only landmanagement agency in Australia that includes fungi inpermanent monitoring programs DEC has a permanent fungalecologist on staff with support staff to undertake monitoringand research in programs and projects including FORESTCHECK(Abbott andBurrows 2004) and theWalpole FireMosaic (WFM)project (Burrows 2006) The focus is on macrofungi and morethan 750 species are currently recognised many of which arenewly recorded or yet to be formally described

FORESTCHECK (see httpwwwdecwagovau accessed 10December 2010) was initiated in 2001 as an integrated long-term landscape-scale program devised to record and monitorthe status and response of key forest organisms and communitiesto both forestmanagement activities andnatural variation (Abbottand Burrows 2004) The WFM project was initiated in 2005 totest the notion that fine-grained mosaics representing variouspatches of vegetation at different post-fire seral stages burnt atvarying intensities and different seasons across a landscape canreduce the severity of wildfires as well as be beneficial to themaintenance of biodiversity (Burrows 2006) Macrofungi havebeen included in both these projects through fruit-bodysurveys leading to a significant increase in knowledge of howmacrofungal communities and key species respond to fire insouthern eucalypt forests (Robinson 2001 2006 Robinson andBougher 2003 Robinson and Tunsell 2007 Robinson et al2008) Ongoing review of survey methods has also providedtechniques that allow monitoring of this traditionally difficultgroup of organisms to be undertaken in a consistent and cost-effective manner

There is clearly an urgent need to include fungi in current andplanned long-termmonitoring programs in relation to the effect offire across Australia The taxonomic scope of surveys needs to bewidened to cover not onlymacrofungi but also the highly diverseand ecologically important leaf-inhabiting parasitic microfungiendophytes and saprotrophic soil microfungi Where it is notpractical to comprehensively survey for all fungi considerationshould be given to surveying for subsets of readily identifiabletaxa representative of trophic phylogenetic and morphologicalgroups particularly in adaptive management systems

Integration of information relatingto the fungal community

Collection of information about fungi is particularly difficult asfungi usually reside within their host or substrate and surveyand identification methods vary depending upon whether


reproductive structures or symptoms are present or not Recentstudies adopt two quite different approaches On the one handfruit-body surveys allow compilation of inventories across arange of sites but do not fully recover the species presentbecause of species not fruiting at the time of survey or everOn the other hand molecular methods allow a snapshot of allspecies present but usually do not identify the particular speciesinvolved

Currently there is poor integration between the molecularcharacterisation of known taxa and sequences that are beingisolated from environmental samples such as ectomycorrhizalroot tips or bulk soil samples The main problem is a lack ofsequence data from authoritatively named material fromAustralia which in the first instance usually requires samplingof fruit-bodies or cultures In the long term it is essential to havelocal accurate and comprehensive barcode databases backed upby voucher material In the meantime at the least it would beuseful to have target groups across phylogenetic and functionalgroups for which the taxonomy (species limits) is reasonablyworked out A barcode library for such targets can then be thebasis formolecular identification of at least a substantial subset ofenvironmental samples

For fruit-body surveys different studies are poorly integratedwith respect to a standard taxonomic framework While somecommon and readily recognisable species appear in species listsfrom across Australia many collections included in inventoriesare assigned tag or field names or are not identified to species atall While tag names can be used consistently within surveys it isnot possible to match them up across different studies withouttime-consuming examination of voucher material (where this isavailable) Improved documentation of the distinctive charactersof taxa to which tag names have been assigned would assist butin the end comprehensive taxonomic revisions are the best wayto provide reliable names to species encountered in ecologicalsurveys Molecular identification of fruit-bodies will also bepossible once comprehensive barcode libraries are available

At present no one method precisely characterises the fungalcommunity in soil or other substrates Even when differentmethods are used in combination there are many fungi thatremain unculturable unrecognisable unidentifiable or difficultto quantify The choice of sampling and identification methodsoften comes down to the resources and funding available withinan organisation Additional information on the cost-effectivenessand accuracy of different methods of isolation and identificationwould assist in the choice of appropriate survey techniques andidentification protocols

Significant data on fungi and their responses to fire currentlyreside in unlinked datasets such as databases and other materialheld by state management agencies and research institutionsand in unpublished reports and studies by fungal interest groupsA meta-analysis of such data would significantly increaseunderstanding of the distribution host and habitat associationsand responses to fire of individual species of fungi

Management of fire for fungi

Mosaic burning is becoming an established means of managingfire and biodiversity at the landscape scale (Grove et al 2002Bradstock 2008 Burrows 2008) Several studies are now under

way in the south-west of Western Australia to investigate theeffect of fire regimes on the diversity of Jarrah forest biotaincluding fungi (Burrows 2006 Wittkuhn et al in press)Fire is thought to increase small-scale heterogeneity of fungalcommunities (Friese et al 1997) and results to date in WesternAustralia indicate that fire mosaics contribute to maintainingdiversity of macrofungi (Robinson 2006 Robinson et al 2008)

In management strategies for the use of prescribed fire inrelation to conservation of biodiversity that are currently beingdeveloped in Victoria and Western Australia fire intervals arebased on lsquovital attributesrsquo (ie life history characteristics) ofplant species and habitat preferences for endangered animals(Fire Ecology Working Group 1999 Burrows 2008) Someknowledge exists of the seral stages favoured by differentfungal species (McMullan-Fisher et al 2002 Ratkowsky2007 Robinson et al 2008) but there is limited understandingof the biology governing these preferences and distributions Inaddition fire may affect fungi in different ways at differentstages in their life cycles For example in sclerotia-producingmacrofungi such as Laccocephalum reproduction through fruit-body production is stimulated by fire but frequent burning mayhave a negative impact because of the requirement during thevegetativephase for larger logs characteristic of longunburnt sites(Grove and Meggs 2003) Fungi should be incorporated intomanagement schemes that use vital attributes and collection ofthe necessary data made a priority

As an interim measure the close correlation of substratecondition (including quality and quantity) with time since firesuggests that management of substrate diversity in differentvegetation types may be an appropriate approach whilespecific requirements of fungi are being investigated InAustralian forests differences in species richness and fungalassemblages at different times since fire have been linked tothe availability of suitable substrates (Tommerup et al 2000McMullan-Fisher et al 2002 Packham et al 2002 Gates et al2005 Robinson et al 2008 Gates et al 2010a 2010b) In thenorthern hemisphere a link has been demonstrated betweenspecies rarity and loss of suitable substrates (Berg et al 1994Jonsson et al 2005Raphael andMolina 2007) The retention andmaintenance of a diverse range of substrates within the landscapehas been highlighted as important for the conservation of fungiand other organisms in Australian forests (Grove and Meggs2003)

Conclusions

Fire impacts directly on all elements of the biota including fungias well as indirectly by inducing changes in soil structure andwater and nutrient availability and cycling Fire ecology isacknowledged as complex and highly variable with responsesof particular species usually dependent on site and speciescharacteristics Not surprisingly the effects of fire on fungalspecies and communities are also complex but are less wellunderstood than for vascular plants This review highlights thatthe effects of fire are often multifaceted with abiotic and bioticinteractions often mediated by particular trophic groups of fungiThese interactions are epitomised by the complex relationshipsamong fire vascular plants mycorrhizal fungi andmycophagousmammals


Fungal habitats and substrates are lost modified or created byfires and the degree of change depends on fire intensity fireregime and the age and type of vegetation on site Short-termeffects include sterilisation of upper soil layers increased pH andreduction or loss of host plants litter and small woody debris Inthe longer term other elements such as standing dead wood andCWD may be consumed or initiated by fire

Based on their importance in ecosystems as symbioticpartners decomposers nutrient cyclers and as a food resourcefor vertebrates and invertebrates fungi should be included in landmanagement decisions However an improved understanding ofthe functional roles of fungi the effects of fire on fungi and thepost-fire interactions between fungi and biotic and abioticcomponents of ecosystems is needed to help managers makeinformed decisions on best management practices

Integration of taxonomic and ecological research is needed tofacilitate better management of fungi Closer coordination ofresearch priorities between management agencies and researchorganisations would assist in this integration Ideally futurefungal research would also integrate both traditional andmolecular techniques to develop a clearer understanding of thecomplex nature of communities and ecosystems particularly insoil and other important fungal substrates such as litter andCWD

Acknowledgements

The reviewwould not have been possible without the generous support of theVictorian National Parks Association to SM-F We thank Frank Udovicic(Royal Botanic GardensMelbourne) for helpful comments on themanuscriptand twoanonymous referees for the considerable amountofvaluable feedbackthey provided

References

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Abbott I Burrows N (2004)Monitoring biodiversity in jarrah forest in south-westWesternAustralia theForestcheck initiative In lsquoTheconservationofAustraliarsquos faunarsquo (Ed D Lunney) pp 947ndash958 (Royal Society of NewSouth Wales Mosman)

Abbott I Loneragan O (1983) Influence of fire on growth rate mortality andbutt damage inMediterranean forest ofWesternAustraliaForest Ecologyand Management 6 139ndash153 doi1010160378-1127(83)90018-X

Adjoud-Sadadou D Halli-Hargas R (2000) Occurrence of arbuscularmycorrhiza on aged Eucalyptus Mycorrhiza 9 287ndash290doi101007PL00009993

Allen EB AllenMF HeelmDJ Trappe JM (1995) Patterns and regulation ofmycorrhizal plant and fungal diversity Plant and Soil 170 47ndash62doi101007BF02183054

Anderson IC Cairney JWG (2004) Diversity and ecology of soil fungalcommunities increased understanding through the application ofmolecular techniques Environmental Microbiology 6 769ndash779doi101111j1462-2920200400675x

Anderson IC Bastias BA Genney DR Parkin PI Cairney JWG (2007)Basidiomycete fungal communities in Australian sclerophyll forest soilare altered by repeated prescribed burning Mycological Research 111482ndash486 doi101016jmycres200702006

Artz RRE Reid E Anderson IC Campbell CD Cairney JWG (2009) Long-term repeated prescribed burning increases evenness in the basidiomycetelaccase gene pool in forest soils FEMS Microbiology Ecology 67397ndash410 doi101111j1574-6941200900650x

Ashton DH (1975) Studies of litter in Eucalyptus regnans forests AustralianJournal of Botany 23 413ndash433 doi101071BT9750413

Bastias BA Huang ZQ Blumfield T Xu Z Cairney JWG (2006a) Influenceof repeated prescribed burning on the soil fungal community in aneastern Australian wet sclerophyll forest Soil Biology amp Biochemistry38 3492ndash3501 doi101016jsoilbio200606007

Bastias BA Xu ZH Cairney JWG (2006b) Influence of long-term repeatedprescribed burning on mycelial communities of ectomycorrhizal fungiNew Phytologist 172 149ndash158 doi101111j1469-8137200601793x

Bastias BA Anderson IC Rangel-Castro JI Parkin PI Prosser JI CairneyJWG (2009) Influence of repeated prescribed burning on incorporation of13C from cellulose by forest soil fungi as determined by RNA stableisotope probing Soil Biology amp Biochemistry 41 467ndash472doi101016jsoilbio200811018

Bellgard SE Whelan RJ Muston RM (1994) The impact of wildfire onvesicular-arbuscular mycorrhizal fungi and their potential to influence there-establishment of post-fire plant communitiesMycorrhiza 4 139ndash146doi101007BF00203532

Berg A Ehnstroumlm B Gustafsson L Hallingbaumlck T Jonsell M Weslien J(1994) Threatened plant animal and fungus species in Swedish forestsdistribution and habitat preferences Conservation Biology 8 718ndash731doi101046j1523-1739199408030718x

Berg B Laskowski R (Eds) (2006) lsquoLitter decomposition a guide to carbonand nutrient turn overrsquo Advances in Ecological Research 38 (AcademicPress London)

Blaney B (1996) Fungal toxins and animals In lsquoFungi of Australia 1B ndash

Introduction ndash Fungi in the environmentrsquo (Eds K Mallett CAGrgurinovic) pp 225ndash238 (ABRS Australia Canberra)

Boddy L (1984) The micro-environment of basidiomycete mycelia intemperate deciduous woodlands In lsquoThe ecology and physiology ofthe fungal myceliumrsquo (Eds D Jennings A Rayner) pp 261ndash289(Cambridge University Press Cambridge UK)

Boddy L (2001) Fungal community ecology and wood decompositionprocesses in angiosperms from standing tree to complete decay ofcoarse woody debris In lsquoEcology of woody debris in boreal forestsrsquo(Eds BG Jonsson N Kruys) pp 43ndash56 (Blackwell Science Oxford)

Bradstock RA (2008) Effects of large fires on biodiversity in south-easternAustralia disaster or template for diversity International Journal ofWildland Fire 17 809ndash822 doi101071WF07153

Bradstock R Auld R Ellis M Cohn J (1992) Soil temperatures duringbushfires in semi-arid mallee shrublands Australian Journal ofEcology 17 433ndash440 doi101111j1442-99931992tb00826x

Bradstock RAWilliams JE Gill MA (Eds) (2002) lsquoFlammableAustralia thefire regimesandbiodiversityof a continentrsquo (CambridgeUniversityPressCambridge UK)

Brennan KEC Christie FJ York A (2009) Global climate change and litterdecomposition more frequent fire slows decomposition and increasesthe functional importance of invertebrates Global Change Biology 152958ndash2971 doi101111j1365-2486200902011x

Bridge P Spooner B (2001) Soil fungi diversity and detectionPlant and Soil232 147ndash154 doi101023A1010346305799

Brodo IM (1973) Substrate ecology In lsquoThe lichensrsquo (Eds V AhmadjianME Hale) pp 401ndash441 (Academic Press London)

Brodo IM Duran Sharnoff S Sharnoff S (2001) lsquoLichens of North Americarsquo(Yale University Press New Haven)

Brown GG (1995) How do earthworms affect microfloral and faunalcommunity diversity In lsquoThe significance and regulation of soilbiodiversityrsquo (Eds HP Collins GP Robertson MJ Klug) pp 247ndash269(Kluwer Academic Publishers Dordrecht)

BrundrettMC (2004)Diversity and classification ofmycorrhizal associationsBiological Reviews of the Cambridge Philosophical Society 79 473ndash495doi101017S1464793103006316

Brundrett MC (2006) Role of symbiotic relationships in Australian terrestrialorchid conservation Australasian Plant Conservation 15 2ndash7

BrundrettMC (2007) Scientific approaches to Australian temperate terrestrialorchid conservation Australian Journal of Botany 55 293ndash307doi101071BT06131


Brundrett MC (2009) Mycorrhizal associations and other means of nutritionof vascular plants understanding the global diversity of host plants byresolving conflicting information and developing reliable means ofdiagnosis Plant and Soil 320 37ndash77 doi101007s11104-008-9877-9

Brundrett MC Ashwath N Jasper DA (1996a) Mycorrhizas in the Kakaduregion of tropical Australia I Propagules of mycorrhizal fungi and soilproperties in natural habitats Plant and Soil 184 159ndash171doi101007BF00029285

Brundrett MC Ashwath N Jasper DA (1996b) Mycorrhizas in the Kakaduregion of tropical Australia II Propagules of mycorrhizal fungi indisturbed habitats Plant and Soil 184 173ndash184

Bruns TD (1995) Thoughts on the processes that maintain local speciesdiversity of ectomycorrhizal fungi Plant and Soil 170 63ndash73doi101007BF02183055

Burrows ND (2006) Burning for biodiversity investigating fine grainmosaics in south-west Australia Australasian Plant Conservation 145ndash6

Burrows ND (2008) Linking fire ecology and fire management in south-westAustralian forest landscapes Forest Ecology and Management 2552394ndash2406 doi101016jforeco200801009

Cairney JWG (2005) Basidiomycete mycelia in forest soils dimensionsdynamics and roles in nutrient distribution Mycological Research 1097ndash20 doi101017S0953756204001753

Cairney JWG Bastias BA (2007) Influences of fire on forest soil fungalcommunities Canadian Journal of Forest Research 37 207ndash215doi101139X06-190

Cairney JWGBurkeRM(1998)Extracellular enzyme activities of the ericoidmycorrhizal endophyte Hymenoscyphus ericae (Read) Korf amp Kernantheir likely roles in decomposition of dead plant tissue in soil Plant andSoil 205 181ndash192 doi101023A1004376731209

Campbell CDCameronCMBastiasBAChenCCairney JWG(2008)Longterm repeated burning in a wet sclerophyll forest reduces fungal andbacterial biomass and responses to carbon substrates Soil Biology ampBiochemistry 40 2246ndash2252 doi101016jsoilbio200804020

Catcheside PS (2009) The phoenicoid discomycetes on Kangaroo IslandFungimap Newsletter 38 5ndash7

Catcheside PS CatchesideDE (2008)A fungal hotspot StringybarkWalkingTrail Deep Creek Conservation Park South Australia and theconservation status of its macrofungi Journal of the Adelaide BotanicGardens 22 9ndash30

Catcheside PS May TW Catcheside DE (2009) The larger fungi in FlindersChase National Park Kangaroo Island Report for the WildlifeConservation Fund and Native Vegetation Council Bellevue HeightsSA

Catling PC (1991) Ecological effects of prescribed burning practices on themammals of southeasternAustralia In lsquoConservation ofAustraliarsquos forestfaunarsquo (Ed D Lunney) pp 353ndash363 (Royal Zoological Society ofNew South Wales Mosman)

Certini G (2005) Effects of fire on properties of forest soil a reviewOecologica 143 1ndash10 doi101007s00442-004-1788-8

Chambers SM Curlevski NJA Cairney JWG (2008) Ericoid mycorrhizalfungi are common root inhabitants of non-Ericaceae plants in a south-eastern Australian sclerophyll forest FEMS Microbiology Ecology 65263ndash270 doi101111j1574-6941200800481x

Chapman AD (2009) lsquoNumbers of living species in Australia and the worldrsquo(Australian Biological Resources Study Canberra)

Cheewangkoon R Groenewald JZ Summerell BA Hyde KD To-anun CCrous PW (2009) Myrtaceae a cache of fungal biodiversity Persoonia23 55ndash85 doi103767003158509X474752

Chen DM Cairney JWG (2002) Investigation of the influence of prescribedburning on ITS profiles of ectomycorrhizal and other soil fungi at threeAustralian sclerophyll forest sitesMycological Research 106 532ndash540doi101017S0953756202005890

Claridge AW (2002) Ecological role of hypogeous ectomycorrhizal fungi inAustralian forests and woodlands Plant and Soil 244 291ndash305doi101023A1020262317539

Claridge AW Barry SC (2000) Factors influencing the distribution ofmedium-sized ground-dwelling mammals in southeastern mainlandAustralia Austral Ecology 25 676ndash688

Claridge AW May TW (1994) Mycophagy among Australian mammalsAustralian Journal of Ecology 19 251ndash275doi101111j1442-99931994tb00489x

Claridge AW Trappe JM (2004) Managing habitat for mycophagous(fungus-feeding) mammals a burning issue In lsquoConservation ofAustraliarsquos forest faunarsquo (Ed D Lunney) pp 936ndash946 (RoyalZoological Society of New South Wales Mosman)

Claridge AW Barry SC Cork SJ Trappe JM (2000a) Diversity and habitatrelationships of hypogeous fungi II Factors influencing the occurrenceand number of taxa Biodiversity and Conservation 9 175ndash199doi101023A1008962711138

ClaridgeAWCorkSJ Trappe JM(2000b)Diversity and habitat relationshipsof hypogeous fungi I Study design sampling techniques and generalsurvey results Biodiversity and Conservation 9 151ndash173doi101023A1008941906441

Claridge AW Trappe JM Hansen K (2009a) Do fungi have a role as soilstabilizers and remediators after forest fire Forest Ecology andManagement 257 1063ndash1069 doi101016jforeco200811011

Claridge AW Trappe JM Mills DJ Claridge DL (2009b) Diversity andhabitat relationships of hypogeous fungi III Factors influencing theoccurrence of fire-adapted speciesMycological Research 113 792ndash801doi101016jmycres200902014

ClarkeMF (2008)Catering for the needs of fauna infiremanagement scienceor just wishful thinking Wildlife Research 35 385ndash394doi101071WR07137

Collett NG Neumann FG Tolhurst KG (1993) Effects of two short rotationprescribed fires in spring on surface-active arthropods and earthwormsin dry sclerophyll eucalypt forest of west-central Victoria AustralianForestry 56 49ndash60

Cook GD (1991) Effects of fire regime on two species of epiphytic orchids intropical savannas of theNorthernTerritoryAustral Ecology 16 537ndash540doi101111j1442-99931991tb01083x

Dahlberg A (2001) Community ecology of ectomycorrhizal fungi anadvancing interdisciplinary field New Phytologist 150 555ndash562doi101046j1469-8137200100142x

Dahlberg A (2002) Effects of fire on ectomycorrhizal fungi in Fennoscandianboreal forests Silva Fennica 36 69ndash80

Dearnaley JDW Le Brocque AF (2006) Endophytic fungi associated withAustralian orchids Australasian Plant Conservation 15 7ndash9

Dell B (2002) Role of mycorrhizal fungi in ecosystems Chiang MaiUniversity Journal 1 47ndash60

Department of Sustainability and Environment (2003) lsquoEcological effects ofrepeated low-intensity fire in a mixed eucalypt foothill forest in south-eastern Australiarsquo Fire research report no 57 (Department ofSustainability and Environment Melbourne Vic)

Dighton J (2003) Fungi secondary productivity and other fungal-faunalinteractions In lsquoFungi in ecosystem processesrsquo (Ed J Dighton)pp 185ndash242 (Marcel Dekker New York)

Dighton J Jones HE Robinson CH Beckett J (1997) The role of abioticfactors cultivation practices and soil fauna in the dispersal of geneticallymodified microorganisms in soils Applied Soil Ecology 5 109ndash131doi101016S0929-1393(96)00137-0

Dix N Webster J (1995) Colonization and decay of wood In lsquoFungalecologyrsquo (Eds N Dix J Webster) pp 145ndash169 (Chapman and HallCambridge UK)

Dromph KM (2003) Collembolans as vectors of entomopathogenic fungiPedobiologia 47 245ndash256 doi1010780031-4056-00188


El-Abyad MSH Webster J (1968a) Studies on pyrophilous discomycetesI Comparative physiological studies Transactions of the BritishMycological Society 51 353ndash367doi101016S0007-1536(68)80002-6

El-Abyad MSH Webster J (1968b) Studies on pyrophilous discomycetesII Competition Transactions of the British Mycological Society 51369ndash375 doi101016S0007-1536(68)80003-8

Eldridge DJ (1996) Distribution and floristics of terricolous lichens in soilcrusts in arid and semi-arid New South Wales Australia AustralianJournal of Botany 44 581ndash599 doi101071BT9960581

Eldridge DJ (2003) Biological soil crusts ndash naturersquos natural soil healers InlsquoPlant conservation ndash approaches and techniques from an Australianperspectiversquo (Eds CL Brown F Hall J Mill) pp Module 9 1ndash12(Australian Network for Plant Conservation Canberra)

EldridgeDJ BradstockR (1994) The effect of time since fire on the cover andcomposition of cryptogamic soil crusts on a eucalypt shrubland soilCunninghamia 3 521ndash527

Eldridge DJ Tozer ME (1997) lsquoA practical guide to soil lichens andbryophytes of Australiarsquos dry countryrsquo (Department of Land andWater Conservation Sydney)

Eldridge DJ Semple WS Koen TB (2000) Dynamics of cryptogamic soilcrusts in a derived grassland in south-eastern Australia Austral Ecology25 232ndash240

Eldridge DJ Freudenberger D Koen TB (2006) Diversity and abundance ofbiological soil crust taxa in relation to fine and coarse-scale disturbancesin grassy eucalypt woodland in eastern Australia Plant and Soil 281255ndash268 doi101007s11104-005-4436-0

Ferguson AV Pharo EJ Kirkpatrick JB Marsden-Smedley JB (2009) Theearly effects of fire and grazing on bryophytes and lichens in tussockgrassland and hummock sedgeland in north-eastern TasmaniaAustralianJournal of Botany 57 556ndash561 doi101071BT09131

FireEcologyWorkingGroup (1999) lsquoManagementoffire for the conservationof biodiversityrsquo (Department of Natural Resources and EnvironmentMelbourne Vic)

FrieseCMorris SAllenM(1997)Disturbance in natural ecosystems scalingfrom fungal diversity to ecosystem functioning In lsquoThe mycotaa comprehensive treatise on fungi as experimental systems for basicand applied researchrsquo (Eds D Wicklow BE Soumlderstroumlm) pp 47ndash63(Springer-Verlag Berlin)

Fuhrer B Robinson R (1992) lsquoRainforest fungi of Tasmania and south-eastAustraliarsquo (CSIRO Publishing Melbourne)

GardesMBrunsTD (1996)Community structure of ectomycorrhizal fungi ina Pinus muricata forest above- and below-ground views CanadianJournal of Botany 74 1572ndash1583 doi101139b96-190

Garkaklis MJ Bradley JS Wooller RD (2000) Digging by vertebrates as anactivity promoting the development of water-repellent patches in sub-surface soil Journal of Arid Environments 45 35ndash42doi101006jare19990603

Garkaklis MJ Bradley JS Wooller RD (2003) The relationship betweenanimal foraging and nutrient patchiness in south-west Australianwoodland soils Australian Journal of Soil Research 41 665ndash673doi101071SR02109

Gates G Ratkowsky DA (2004) A preliminary census of the macrofungi ofMt Wellington Tasmania ndash the non-gilled Basidiomycota Papers andProceedings of the Royal Society of Tasmania 138 53ndash59

Gates G Ratkowsky DA (2005) A preliminary census of the macrofungi ofMt Wellington Tasmania ndash the Ascomycota Papers and Proceedings ofthe Royal Society of Tasmania 139 49ndash52

Gates GM Ratkowsky DA Grove SJ (2005) A comparison of macrofungi inyoung silvicultural regeneration andmature forest at theWarra LTER sitein the southern forests of Tasmania Tasforests 16 127ndash152

Gates GM Ratkowsky DA Grove SJ (2009) Aggregated retention andmacrofungi a case study from the Warra LTER site TasmaniaTasforests 18 33ndash54

GatesGMMohammedCWardlawTDavidsonNJRatkowskyDA (2010a)Diversity and phenology of the macrofungal assemblages supportedby litter in a tall wet Eucalyptus obliqua forest in southern TasmaniaAustralia Fungal Ecology in press doi101016jfuneco201008001

GatesGMMohammedCWardlawTRatkowskyDADavidsonNJ (2010b)The ecology and diversity of wood-inhabiting macrofungi in a nativeEucalyptus obliqua forest of southern Tasmania Australia FungalEcology in press doi101016jfuneco201007005

Gehring CA Wolf JE Theimer T (2002) Terrestrial vertebrates promotearbuscular mycorrhizal fungal diversity and inoculum potential in arainforest soil Ecology Letters 5 540ndash548doi101046j1461-0248200200353x

George P (2008) Fungimap surveys on Kangaroo Island FungimapNewsletter 36 13ndash14

Gill AM Groves RH Noble IR (Eds) (1981) lsquoFire and the Australian biotarsquo(Australian Academy of Science Adelaide)

GlenM (2002)Genetic variation in ectomycorrhizal fungi and its exploitationin ecological investigations of eucalypt forests PhD Thesis MurdochUniversity Perth WA

Glen M Tommerup IC Bougher NL OrsquoBrien PA (1998) A survey of forestectomycorrhizal fungal communities by PCR-RFLP analysis of root tips(Abstract) In lsquoProgramme and Abstracts of the Second InternationalConference on Mycorrhizarsquo Uppsala Sweden p 70

GlenMTommerup ICBougherNLOrsquoBrienPA (1999)PCR-RFLPanalysisof root tip fungi in forests (Abstract) In lsquoProceedings of the AustralasianPlant Pathology Society 12th Biennial Conferencersquo Australasian PlantPathology Society Canberra Australia p 127

Glen M Tommerup IC Bougher NL OrsquoBrien PA (2001) Interspecific andintraspecific variation of ectomycorrhizal fungi associated withEucalyptus ecosystems as revealed by ribosomal DNA PCR-RFLPMycological Research105 843ndash858 doi101017S095375620100418X

Gochenaur S (1981) Response of soil fungal communities to disturbance InlsquoThe fungal community its organization and role in the ecosystemrsquo (EdsGC Wicklow DT Carroll) pp 459ndash479 (Marcel Dekker New York)

Gray D Dighton J (2006) Mineralization of forest litter nutrients by heat andcombustion Soil Biology amp Biochemistry 38 1469ndash1477doi101016jsoilbio200511003

GreensladeP (1997) Short term effects of a prescribedburn on invertebrates ingrassy woodland in south-east Australia Memoirs of the Museum ofVictoria 56 305ndash312

GroveSMeggs J (2003)Coarsewoodydebris biodiversity andmanagementa review with particular reference to Tasmanian wet eucalypt forestsAustralian Forestry 66 258ndash272

Grove SMeggs J GoodwinA (2002) lsquoA review of biodiversity conservationissues relating to coarse woody debris management in the wet eucalyptproduction forests of Tasmaniarsquo (Forestry Tasmania Hobart)

Hanski I (1989) Fungivory fungi insects and ecology In lsquoInsectndashfungusinteractionsrsquo (Eds N Wilding NM Collins PM Hammond JF Webber)pp 25ndash68 (Academic Press London)

Hart SC DeLuca TH Newman GS MacKenzie MD Boyle SI (2005) Post-fire vegetative dynamics as drivers of microbial community structure andfunction in forest soils In lsquoForest soils research theory reality and its rolein technology transferrsquo (Eds MR Gale RF Powers JR Boyle)pp 166ndash184 (Elsevier London)

Hilton RNMalajczuk N PearceMH (1989) Larger fungi of the jarrah forestan ecological and taxonomic survey In lsquoThe jarrah forest ndash a complexmediterranean ecosystemrsquo (Eds B Dell JJ Havel N Malajczuk)pp 89ndash109 (Kluwer Academic Publishers Dordrecht)

Hollis J Whitford K Robinson R Danti S (2008) Down but not outdiscovering the significance of dead wood Landscope 24 10ndash16

Hopkins AJM Harrison KS Grove SJ Wardlaw TJ Mohammed CL (2005)Wood-decay fungi and saproxylic beetles association with livingEucalyptus obliqua trees early results from studies at the Warra LTERsite Tasmania Tasforests 16 111ndash125


Howell J Humphreys G Mitchell P (2006) Changes in soil water repellenceand its distribution in relation to surface microtopographic units after alow severity fire in eucalypt woodland Sydney Australia AustralianJournal of Soil Research 44 205ndash217 doi101071SR04176

Huang PM Wang MC Wang MK Daniel H (2005) Mineral-organic-microbial interactions In lsquoEncyclopedia of soils in the environmentrsquo(Ed D Hillel) pp 486ndash499 (Elsevier Oxford)

Hughes KW Petersen RH Lickey EB (2009) Using heterozygosity toestimate a percentage DNA sequence similarity for environmentalspeciesrsquo delimitation across basidiomycete fungi New Phytologist 182795ndash798 doi101111j1469-8137200902802x

Humphreys FR LambertMJ (1965) Soil temperature profiles under slash andlog fires of various intensities Australian Forest Research 1 23ndash29

Huston MA (1996) Models and management implications of coarse woodydebris impacts on biodiversity In lsquoWorkshop on coarse woody debris insouthern forests effects onbiodiversityrsquo (Eds JWMcMinnDACrossley)pp 139ndash143 (USDA Forest Service Athens)

Ing B (1993) Towards a RED list of endangered European macrofungi InlsquoFungi of Europe investigation recording and conservationrsquo (Eds DNPegler L Boddy B Ing PM Kirk) pp 231ndash237 (The Royal BotanicGardens Kew UK)

Izzo A Nguyen D Bruns T (2006) Spatial structure and richness ofectomycorrhizal fungi colonizing bioassay seedlings from resistantpropagules in a Sierra Nevada forest comparisons using two hosts thatexhibit different seedling establishment patterns Mycologia 98374ndash383 doi103852mycologia983374

Jobbaacutegy EG Jackson RB (2001) The distribution of soil nutrients with depthglobal patterns and the imprint of plants Biogeochemistry 53 51ndash77doi101023A1010760720215

Johnson CN (1994) Fruiting of hypogeous fungi in dry sclerophyll forestin Tasmania Australia seasonal variation and annual productionMycological Research 98 1173ndash1182doi101016S0953-7562(09)80201-3

Johnson CN (1996) Interactions between mammals and ectomycorrhizalfungi Trends in Ecology amp Evolution 11 503ndash507doi101016S0169-5347(96)10053-7

Johnson CN (1997) Fire and habitat management for a mycophagousmarsupial the Tasmanian bettong (Bettongia gaimardia) AustralianJournal of Ecology 22 101ndash105doi101111j1442-99931997tb00645x

Johnston PR (2001) Rhytismatales of Australasia Australian SystematicBotany 14 377ndash384 doi101071SB99035

Johnston PRMay TW ParkDHorak E (2007)Hypocreopsis amplectens spnov a rare fungus fromNewZealandandAustraliaNewZealandJournalof Botany 45 715ndash719 doi10108000288250709509746

Jonsson BG Kruys N Ranius T (2005) Ecology of species living on deadwood ndash lessons for dead wood management Silva Fennica 39 289ndash309

Kantvilas G (1990) The genus Roccellinastrum in Tasmania Lichenologist(London England) 22 79ndash86 doi101017S0024282990000044

Kantvilas G (2000) Conservation of Tasmanian lichens Forest Snow andLandscape Research 75 357ndash367

Kantvilas G Jarman SJ (1993) The cryptogamic flora of an isolated rainforestfragment in Tasmania Botanical Journal of the Linnean Society 111211ndash228 doi101111j1095-83391993tb01899x

KantvilasG JarmanSJ (2004)Lichens andbryophytes onEucalyptus obliquain Tasmania management implications in production forests BiologicalConservation 117 359ndash373 doi101016jbiocon200308001

Kantvilas G Jarman SJ (2006) Recovery of lichens after logging preliminaryresults fromTasmaniarsquoswet forestsLichenologist (LondonEngland) 38383ndash394 doi101017S0024282906005949

Kile GA (1980) Behaviour of an Armillaria in some Eucalyptus obliqua-Eucalyptus regnans forests in Tasmania and its role in their declineEuropean Journal of Forest Pathology 10 278ndash296doi101111j1439-03291980tb00040x

Kile GA (1981) Armillaria luteobubalina a primary cause of the decline anddeath of trees in mixed species eucalypt forests in central VictoriaAustralian Forest Research 11 63ndash77

Kile GA JohnsonGC (2000) Stem and butt rot of eucalypts In lsquoDiseases andpathogens of eucalyptsrsquo (Eds PJKeaneGAKile FDPodger BNBrown)pp 307ndash338 (CSIRO Publishing Melbourne)

Kjoller R Bruns TD (2003) Rhizopogon spore bank communities within andamong California pine forests Mycologia 95 603ndash613doi1023073761936

Klopatek C DeBano L Klopatek J (1988) Effects of simulated fire onvesicular-arbuscular mycorrhizae in pinyon-juniper woodland soilPlant and Soil 109 245ndash249 doi101007BF02202090

Launonen TM Ashton DH Keane PJ (1999) The effect of regenerationburns on the growth nutrient acquisition and mycorrhizae of Eucalyptusregnans F Muell (Mountain Ash) seedlings Plant and Soil 210273ndash283 doi101023A1004609912315

Lee KE Foster RC (1991) Soil fauna and soil structure Australian Journalof Soil Research 29 745ndash775 doi101071SR9910745

Mackensen J Bauhus J Webber E (2003) Decomposition rates of coarsewoody debris ndash a review with particular emphasis on Australian treespecies Australian Journal of Botany 51 27ndash37 doi101071BT02014

Malajczuk N Hingston FJ (1981) Ectomycorrhizae associated with jarrahAustralian Journal of Botany 29 453ndash462 doi101071BT9810453

Marx DH (1972) Ectomycorrhizae as biological deterrents to pathogenicroot infections Annual Review of Phytopathology 10 429ndash454doi101146annurevpy10090172002241

MayTW(1997)Fungi In lsquoAconservationoverviewofAustraliannon-marinelichens bryophytes algae and fungirsquo (Eds GAM Scott TJ EntwisleTW May GN Stevens) pp 49ndash74 (Environment Australia Canberra)

May TW (2003) Conservation of Australian fungi ndash knowledge is the key InlsquoPlant conservation ndash approaches and techniques from an Australianperspectiversquo (Eds CL Brown F Hall J Mill) Module 9 pp 1ndash16(Australian Network for Plant Conservation Canberra)

May TW Simpson JA (1997) Fungal diversity and ecology in eucalyptecosystems In lsquoEucalypt ecology individuals to ecosystemsrsquo (Eds JEWilliams JCZ Woinarski) pp 246ndash277 (Cambridge University PressCambridge UK)

McCaw WL (1983) Wood defect associated with fire scares on jarrahAustralian Forest Research 13 261ndash266

McCaw WL Neal JE Smith RH (1996) Fuel accumulation followingprescribed burning in young even-aged stands of karri (Eucalyptusdiversicolor) Australian Forestry 59 171ndash177

McCaw WL Neal JE Smith RH (2002) Stand characteristics and fuelaccumulation in a sequence of even-aged karri (Eucalyptusdiversicolor) regrowth stands in south-west Western Australia ForestEcology and Management 158 263ndash271doi101016S0378-1127(00)00719-2

McGee PA Markovina A Jeong GCE Cooper ED (2006) Trichocomaceaein bark survive high temperatures and fire FEMSMicrobiology Ecology56 365ndash371 doi101111j1574-6941200600079x

McMullan-Fisher SJM May TW Keane PJ (2002) The macrofungalcommunity and fire in a Mountain Ash forest in southern AustraliaFungal Diversity 10 57ndash76

McMullan-Fisher SJM May TW Kirkpatrick JB (2003) Some macrofungifrom alpine Tasmania Australasian Mycologist 22 44ndash52

Mistry J (1998) A preliminary lichen fire history (LFH) key for the cerrado oftheDistrito Federal centralBrazil Journal of Biogeography25 443ndash452doi101046j1365-269919982530443x

Morley S Gibson M (2004) Cool temperate rainforest lichens of VictoriaAustralia floristics and distribution The Bryologist 107 62ndash74doi1016390007-2745(2004)107[62CTRLOV]20CO2

Neary DG Klopatek CC Debano LF Ffolliot PF (1999) Fire effects onbelowground sustainability a review and synthesis Forest Ecology andManagement 122 51ndash71 doi101016S0378-1127(99)00032-8


NearyDGRyanKCDeBanoLF (2005) lsquoWildlandfire in ecosystems effectsof fire on soils and waterrsquo p 250 (US Department of Agriculture ForestService Rocky Mountain Research Station Ogden Utah US)

Neumann FG (1991) Responses of litter arthropods to major natural orartificial ecological disturbances in Mountain Ash forest AustralianJournal of Ecology 16 19ndash32 doi101111j1442-99931991tb01478x

Neumann FG Tolhurst K (1991) Effects of fuel reduction burning on epigealarthropods and earthworms in dry sclerophyll eucalypt forest of west-central Victoria Australian Journal of Ecology 16 315ndash330doi101111j1442-99931991tb01060x

New TR Yen AL Sands DPA Greenslade P Neville PJ York A Collett NJ(2010) Plannedfires and invertebrate conservation in south east AustraliaJournal of Insect Conservation 14 567ndash574doi101007s10841-010-9284-4

OrsquoBryan K Prober S Lunt I Eldridge D (2009) Frequent fire promotesdiversity and cover of biological soil crusts in a derived temperategrassland Oecologia 159 827ndash838 doi101007s00442-008-1260-2

Odor P Heilmann-Clausen J ChristensenM Aude E van Dort KW PiltaverA Siller I Veerkamp MT Walleyn R Standovar T van Hees AFMKosec J Matocec N Kraigher H Grebenc T (2006) Diversity of deadwood inhabiting fungi and bryophytes in semi-natural beech forests inEurope Biological Conservation 131 58ndash71doi101016jbiocon200602004

OoiMKJWhelanRJAuldTD(2006)Persistenceofobligate-seeding speciesat the population scale effects of fire intensity fire patchiness and longfire-free intervals International Journal of Wildland Fire 15 261ndash269doi101071WF05024

Osborn M (2007) Long-term effects of frequent burning on fungalcommunities and the role of fungi in fire-prone forests PhD ThesisUniversity of Melbourne

Packham JM May TW Brown MJ Wardlaw TJ Mills AK (2002)Macrofungal diversity and community ecology in mature and regrowthwet eucalypt forest in Tasmania amultivariate studyAustral Ecology 27149ndash161 doi101046j1442-9993200201167x

Parmenter JR Jr (1977) Effects of fire on pathogens In lsquoProceedings ofthe symposium on the environmental consequences of fire and fuelmanagement in Mediterranean ecosystemsrsquo (Eds HA Mooney CEConrad) pp 58ndash64 (USDA Forest Service Palo Alto CA)

Pattinson GS Hammill KA Sutton BG McGee PA (1999) Simulatedfire reduces the density of arbuscular mycorrhizal fungi at the soilsurface Mycological Research 103 491ndash496doi101017S0953756298007412

Penttilauml R Kotiranta H (1996) Short-term effects of prescribed burning onwood-rotting fungi Silva Fennica 30 399ndash419

Petersen PM (1970) Danish fireplace fungi an ecological investigationDansk Botanisk Arkiv 27 1ndash97

Petersen PM (1971) Changes of the fungus flora after treatment with variouschemicals Botanisk Tidsskrift 65 264ndash280

Raison RJ (1979) Modification of the soil environment by vegetation fireswith particular reference to nitrogen transformations a review Plant andSoil 51 73ndash108 doi101007BF02205929

Raison RJ Khanna PKWoods PV (1985)Mechanisms of element transfer tothe atmosphere during vegetation fires Canadian Journal of ForestResearch 15 132ndash140 doi101139x85-022

Raphael MG Molina R (Eds) (2007) lsquoConservation of rare or little-knownspeciesrsquo (Island Press Washington)

Rasmussen HN (1995) lsquoTerrestrial orchids from seed to mycotrophic plantrsquo(Cambridge University Press Cambridge UK)

Ratkowsky DA (2007) Visualising macrofungal species assemblagecompositions using canonical discriminant analysis AustralasianMycologist 26 75ndash85

RatkowskyDA Gates GM (2002) A preliminary census of the macrofungi ofMount Wellington Tasmania ndash the Agaricales Papers and Proceedingsof the Royal Society of Tasmania 136 89ndash100

Ratkowsky DA Gates GM (2008) Generalised canonical correlationsanalysis for explaining macrofungal species assemblages AustralasianMycologist 27 33ndash40

Ratkowsky DA Gates GM (2009) Macrofungi in early stages of forestregeneration in Tasmaniarsquos southern forests Tasforests 18 55ndash66

Rayner ADM Boddy L (1988) lsquoFungal decomposition of wood its biologyand ecologyrsquo (John Wiley and Sons Brisbane)

Reddell P Malajczuk N (1984) Formation of Mycorrhizae by Jarrah(Eucalyptus marginata Donn ex Smith) in litter and soil AustralianJournal of Botany 32 511ndash520 doi101071BT9840511

Robinson RM (2001) Fruits of fire Landscope 16 48ndash53Robinson RM (2006) Varying fire regimes to promote fungal diversity across

the landscape Australasian Plant Conservation 14 16ndash17Robinson RM (2009) Laccocephalums on Kangaroo Island Fungimap

Newsletter 37 6ndash7Robinson RM Bougher NL (2003) The response of fungi to fire in jarrah

(Eucalyptus marginata) and karri (Eucalyptus diversicolor) forests ofsouth-west western Australia In lsquoFire in ecosystems of south-westwestern Australia impacts and managementrsquo (Eds I Abbott NBurrows) pp 269ndash289 (Backhuys Publishers Leiden)

Robinson RM Tunsell VL (2007) A list of macrofungi recorded in burntand unburnt Eucalyptus diversicolor regrowth forest in the southwest ofWestern Australia 1998ndash2002 Conservation amp Society 6 75ndash96

Robinson RM Mellican AE Smith RH (2008) Epigeous macrofungalsuccession in the first five years following a wildfire in karri(Eucalyptus diversicolor) regrowth forest in Western Australia AustralEcology 33 807ndash820 doi101111j1442-9993200801853x

Rogers RW (1995) Lichen succession on leaves of the rainforest shrubCapparis arborea (Capparaceae) Australian Journal of Botany 43387ndash396 doi101071BT9950387

Schenck NC Graham SO Green NE (1975) Temperature and light effecton contamination and spore germination of VA mycorrhizal fungiMycologia 67 1189ndash1192 doi1023073758840

Shaw PJA (1992) Fungi fungivores and fungal food webs In lsquoThe fungalcommunity its organisation and role in the ecosystemrsquo (Eds GC CarrolDT Wicklow) pp 295ndash310 (Marcel Dekker New York)

SheaSRMcCormick J PortlockCC(1979)The effect offires on regenerationof leguminous species in the northern jarrah (Eucalyptus marginata Sm)forest of Western Australia Australian Journal of Ecology 4 195ndash205doi101111j1442-99931979tb01210x

Smith SE Read DJ (2002) lsquoMycorrhizal symbiosisrsquo (Academic PressLondon)

Spooner B (1987) lsquoHelotiales of Australasia Geoglossaceae OrbiliaceaeSclerotiniaceae Hyaloscyphaceaersquo (J Cramer Berlin)

Stevens GN (1997) Lichens In lsquoA conservation overview of Australian non-marine lichens bryophytes algae and fungirsquo (Eds GAM Scott TJEntwistle TW May GN Stevens) (Environment Australia Canberra)

Taylor RJ (1992) Reply fire mycorrhizal fungi and management ofmycophagous marsupials Australian Journal of Ecology 17 227ndash228doi101111j1442-99931992tb00802x

TedersooLKoljalgUHallenbergNLarssonK(2003)Fine scaledistributionof ectomycorrhizal fungi and roots across substrate layers includingcoarse woody debris in a mixed forest New Phytologist 159 153ndash165doi101046j1469-8137200300792x

Tedersoo L Jairus T Horton BM Abarenkov K Suvi T Saar I Kotildeljalg U(2008) Strong host preference of ectomycorrhizal fungi in a Tasmanianwet schlerophyll forest as revealed by DNA barcoding and taxon-specificprimers New Phytologist 180 479ndash490doi101111j1469-8137200802561x

Tedersoo L Gates G Dunk CW Lebel T May TW Kotildeljalg U Jairus T(2009a) Establishment of ectomycorrhizal fungal community onNothofagus cunninghamii seedlings regenerating on dead wood inAustralian wet temperate forests does fruit-body type matterMycorrhiza 19 403ndash416 doi101007s00572-009-0244-3


Tedersoo L May TW Smith ME (2009b) Ectomycorrhizal lifestyle in fungipatterns of evolution and distribution Mycorrhiza 20 217ndash263doi101007s00572-009-0274-x

Theodorou C Bowen GD (1982) Effects of a bushfire on the microbiology ofa South Australian low open (dry sclerophyll) forest soil AustralianForest Research 12 317ndash327

TiunovAVScheuS (2005)Arbuscularmycorrhizae andcollembolan interactin affecting community composition of saprotrophic microfungiOecologia 142 636ndash642 doi101007s00442-004-1758-1

Tommerup IC Bougher NL (2000) The role of ectomycorrhizal fungi innutrient cycling in temperate Australian woodlands In lsquoTemperateeucalypt woodlands in Australiarsquo (Eds RJ Hobbs CJ Yates)pp 190ndash224 (Surrey Beatty and Sons Pty Ltd Chipping Norton NSW)

Tommerup IC Bougher NL Syme K Syme A Fernie G (2000) Preliminaryguidelines for managing fungal biodiversity in remnant Eucalyptusmarginata or other Eucalyptus forest types using fire as a toolEcological Management amp Restoration 1 146ndash147

Trappe JM Claridge AW Jumpponen A (2005) Fire hypogeous fungi andmycophagous marsupials Mycological Research 109 516ndash518doi101017S0953756205233014

Trappe JM Nicholls AO Claridge AW Cork SJ (2006) Prescribed burningin a Eucalyptus woodland suppresses fruiting of hypogeous fungi animportant food source for mammals Mycological Research 1101333ndash1339 doi101016jmycres200607018

Trappe JM Bougher NL Castellano M Claridge AW Gates GM Lebel TRatkowsky DA (2008) A preliminary census of the macrofungi of MtWellington Tasmania ndash the sequestrate species Papers and Proceedingsof the Royal Society of Tasmania 142 85ndash95

van der Heijden MGA Sanders IR (Eds) (2002) lsquoMycorrhizal ecologyrsquoEcological studies (Springer London)

Vernes K (2009) Mycophagy in a community of macropod species InlsquoMacropods the biology of kangaroos wallabies and rat-kangaroosrsquo(Eds GM Coulson MDB Eldridge) (Surrey Beattie and Sons Sydney)

Vernes K Dunn L (2009) Mammal mycophagy and fungal spore dispersalacross a steep environmental gradient in eastern Australia AustralEcology 34 69ndash76 doi101111j1442-9993200801883x

Vernes K Haydon DT (2001) Effect of fire on northern bettong (Bettongiatropica) foraging behaviour Austral Ecology 26 649ndash659doi101046j1442-9993200101141x

Vernes K McGrath K (2009) Are introduced black rats (Rattus rattus) afunctional replacement for mycophagous native rodents in fragmentedforests Fungal Ecology 2 145ndash148 doi101016jfuneco200903001

Vernes K Trappe JM (2007) Hypogeous fungi in the diet of the red-leggedpademelon (Thylogale stigmatica) from a rainforest-open forest interfacein northeastern Australia Australian Zoologist 34 203ndash208

Vernes K CastellanoM Johnson CN (2001) Effects of season and fire on thediversity of hypogeous fungi consumed by a tropical mycophagousmarsupial Journal of Animal Ecology 70 945ndash954doi101046j0021-8790200100564x

Vernes K Johnson CN Castellano MA (2004) Fire-related changes inbiomass of hypogeous sporocarps at foraging points used by a tropicalmycophagous marsupial Mycological Research 108 1438ndash1446doi101017S0953756204000048

Warcup JH (1981) Effect of fire on the soil microflora and other non-vascularplants In lsquoFire and the Australian biotarsquo (Eds AM Gill RH GrovesIR Noble) pp 203ndash214 (Academy of Science Canberra)

Warcup JH (1983) Effect of fire and sun-baking on the soil microfloraand seedling growth in forest soils In lsquoSoils an Australian viewpointrsquo(Ed Division of Soils) pp 735ndash740 (CSIRO and Academic PressMelbourne)

Warcup JH (1990) Occurrence of ectomycorrhizal and saprophyticdiscomycetes after a wildfire in a eucalypt forest MycologicalResearch 94 1065ndash1069 doi101016S0953-7562(09)81334-8

Warcup JH Baker KF (1963) Occurrence of dormant ascospores in soilNature 197 1317ndash1318 doi1010381971317a0

Wardlaw TJ Grove SJ Hopkins A Yee M Harrison K Mohammed CL(2009) The uniqueness of habitats in old eucalypts contrasting wood-decay fungi and saproxylic beetles of young and old eucalypts Tasforests18 17ndash32

Wardle DA Bardgett RD Klironomos JN Setaumllauml H van der PuttenWHHallDH (2004) Ecological linkages between aboveground and belowgroundbiota Science 304 1629ndash1633 doi101126science1094875

Watson PJ Bradstock RA Morris EC (2009) Fire frequency influencescomposition and structure of the shrub layer in an Australiansubcoastal temperate grassy woodland Austral Ecology 34 218ndash232doi101111j1442-9993200801924x

Wicklow DT (1975) Fire as an environmental cue initiating ascomycetedevelopment in a tallgrass prairie Mycologia 67 852ndash862doi1023073758344

Williams RJ Bradstock RA (2008) Large fires and their ecologicalconsequences introduction to the special issue International Journalof Wildland Fire 17 685ndash687 doi101071WF07155

Wills RT (1983) The ecology of the wood-rotting Basidiomycete Polyporustumulosus with special reference to the significance of fire HonoursThesis University of Western Australia Perth WA

Wittkuhn RS McCaw WL Wills A Robinson RM Andersen AN VanHeurck P Farr JD Liddelow GL Cranfield R (2010) Fire intervalsequences has minimal effects on species richness or communitycomposition in fire-prone landscapes of southern Western AustraliaForest Ecology and Management in press

WittkuhnRSMcCawLWills A RobinsonR AndersenAN VanHeurck PFarr J Liddelow G Cranfield R (in press) Variation in fire intervalsequences has minimal effects on species richness and composition infire-prone landscapes of south-west Western Australia Forest Ecologyand Management in press

Yates CP Edwards AC Russell-Smith J (2008) Big fires and their ecologicalimpacts in Australian savannas size and frequencymatters InternationalJournal of Wildland Fire 17 768ndash781 doi101071WF07150

York A (1999) Long-term effects of frequent low-intensity burning on theabundance of litter-dwelling invertebrates in coastal blackbutt forests ofsoutheastern Australia Journal of Insect Conservation 3 191ndash199doi101023A1009643627781

Zak B (1964) Role of mycorrhizae in root disease Annual Reviewof Phytopathology 2 377ndash392doi101146annurevpy02090164002113

Zak JC (1992) Response of soil fungal communities to disturbanceIn lsquoThe fungal community its organization and role in the ecosystemrsquo(Eds DT Carroll GC Wicklow) pp 403ndash425 (Marcel DekkerNew York)

Manuscript received 3 March 2010 accepted 7 December 2010


httpwwwpublishcsiroaujournalsajb

important in old highly weathered and nutrient-poor soils(Brundrett 2009) such as those found across the majority ofthe Australian landscape Fungi are essential participants in thecycling of carbon nitrogen phosphorus and other nutrientsin ecosystems (Berg and Laskowski 2006) Consumption offruit-bodies and mycelium of fungi by mycophagous animalsparticularly invertebrates also contributes to nutrient cyclingAnother critical role of fungi in ecosystem function is theirinfluence on soil in relation to ionic exchange particleaggregation carbon content water holding capacity and waterinfiltration (Bastias et al 2006a Claridge et al 2009a)

Fungi are a significant but often overlooked component ofthe Australian biota Some 11 846 described species of fungiare known from Australia of which 3495 form lichens(Chapman 2009) Estimated diversity is at least 10 000 speciesfor macrofungi (about twice as many as are currently known)and as high as 250 000 species for microfungi (Chapman 2009)Not only are there many species awaiting formal descriptionbut the biology and ecology of named species is usually poorlyknown Very few fungi are formally listed on state or nationalconservation schedules (May 1997 2003) Management of fungiin native ecosystems is compromised by the almost total lack ofstaff with expertise in or responsibility for fungi in conservationand management agencies

Fire both planned and unplanned is an integral part ofAustralian ecosystems and impacts on fungi in various waysthrough physical and chemical changes to habitats andsubstrates Heating during fire causes sterilisation of uppersoil layers and loss of nutrients and carbon via volatilisationand combustion of litter and soil organic matter (Certini 2005)Most soil nutrients are concentrated in these upper layers(Jobbaacutegy and Jackson 2001) as are most organisms (Lee andFoster 1991 Dahlberg 2001 Huang et al 2005) For most firesradiation provides enough heat to kill soil organisms includingfungi in the upper soil layer although heat intensity declinesrapidlywith depth (Humphreys and Lambert 1965 Raison et al1985) Heating to 60C is sufficient to kill living tissueparticularly unprotected fungal mycelia (Schenck et al 1975Klopatek et al 1988) and upper layers of soil often reach thistemperature during fires (Humphreys and Lambert 1965Bradstock et al 1992 Pattinson et al 1999) Neverthelessfungi that have evolved in fire-prone Australian environmentshave adaptations to cope with fire such as heat-resistant sporesor other resting structures (Warcup 1981) Fire can also affectfungi by removing and creating substrates for saprotrophs suchas litter and woody debris or by detrimental effects onmutualistic partners such as spore dispersers or mycorrhizalhosts (Dahlberg 2002)

Soil nutrients are altered by fire which in turn impacts theutilisation and genesis of nutrients by fungi and consequentlyother biota Despite a direct loss of nitrogen from soil viavolatilisation nitrogen and phosphorus availability in soilfollowing fire can increase or decrease depending on fireintensity (Raison 1979 Raison et al 1985 Gray and Dighton2006) Changes in nutrient availability subsequently impact onrecovery of vegetation and nutrient cycling (Neary et al 19992005Howell et al 2006) Fire also alters soil fungal communitiesinvolved in decomposition (Neary et al 1999 Boddy 2001)therefore altering carbon nitrogen ratios and mineralisation

rates which ultimately affect plant growth and productivity(Neary et al 1999 2005 Boddy 2001)

The understanding of fire regimes and the effects of fire onAustralian vegetation has developed rapidly from the generallyscience-based endeavour of lsquoFire and the Australian Biotarsquo(Gill et al 1981) to a much more management- consequence-and biodiversity-orientated approach (see Bradstock et al2002 Abbott and Burrows 2003) Recent extensive fires thathave occurred on most continents have prompted recognitionthat the incidence and characteristics of large fires may alter inthe future as a consequence of global climate change (Williamsand Bradstock 2008) Fires whether planned or unplannedindependent of season are patchy and the effects on soil areequally irregular (Bradstock 2008) It is now widely recognisedthat no single optimum fire regime will meet all managementobjectives (Burrows 2008) and diversity of fire regimes amonglandscapes including long unburnt patches is now beingaccepted by land management agencies in adaptive firemanagement programs Fire mosaics resulting from cumulativeheterogeneity (patchiness) from successive fires may contributeto increased local biodiversity (Grove et al 2002 Bradstock2008 Burrows 2008) Although fire management policies aim tomaximise biodiversity the data upon which decisions are basedpredominantly concernvegetation fuel quantities andconditionsand the likely locations of recognised threatened species Thusmost biota especially fungi are not explicitly covered by currentfire management policies

Recently concern has been expressed that management offire for vegetation may not provide the best outcomes forother biota (Clarke 2008 New et al 2010) In turn due to theinterconnectedness of fungi with the biota in general otherorganisms that rely on fungi could also be compromisedby knowledge deficiencies about their fungal partners Thepurpose of this review is to summarise the currentunderstanding of how fire affects fungi in Australianecosystems to identify knowledge gaps and to suggest stepsby which fungi and fire may be better managed After anoverview of the current literature on fungi and fire inAustralia and discussion of fungi that do or do not require fire(and the optimum fire frequency for fungi) we deal with fireand fungi in the framework of fungal trophic groups andsubstrates in order to highlight the functional roles of fungiand to suggest a context that will aid integration of fungi intoecosystem management

Fire and the fungal community in Australian ecosystems

Studies of the effect of fire on fungal communities in Australianecosystems are summarised in Table 1 The 30 field studiesidentified utilised a range of techniques but all compare thefungi present on sites with different times since fire Most studiesincluded recently burnt sites but in some cases (eg Ratkowskyand Gates 2008) the comparison was among sites of known firehistory that had not been burnt for several years to many decadesIn general long unburnt (commonly referred to as control) siteswere incorporated but some studies focussed only on recentlyburnt sites where the fire was less than 10 years ago (Theodorouand Bowen 1982 Warcup 1990 Bellgard et al 1994 Launonenet al 1999 Vernes and Haydon 2001 Vernes et al 2001 2004




















Pyrophilous fungi




Tab

le1

Stud

iesof

fung

alcommun

itiesin


know

nfire

history

Multip

lestud

ieson


ireages

with

anasterisk


dergon

erepeated

burns

Vegetationtype

(dom

inantvegetation)

State

Sites(age

sincefireinyears)

Fun

gigrou

psstud

ied

Sam

plingandidentifi

catio

ntechniqu

esandor

estim

ationof

fungalactiv

ityReference

Fire0ndash12

mon

ths

Fire

gt1ndash10

years

Mature

Trophic

group

Morph

o-logical

group

Sam

ple

collection

techniqu

e

Identifi

catio

ntechnique

(metho

dor

scop

e)

Estim

ateof

fungalactiv

ity(m

etho

d)

Wetscleroph

yllforest(Eo

bliqua

andor

Ed

elegatensis)

Tas

ndashndash

+(25ndash30

gt1

00)

All

EM

FB

MOR(SP)

ndashPackh

ametal(20

02)

Wetscleroph

yllforest(Eo

bliqua)

Tas

++(3)

+(65ndash10

1)L

LFB

MOR(SP)

ndashKan

tvila

san

dJa

rman

(200

6)Wetscleroph

yllforest(Eo

bliqua)

Tas

++

+(73)

All

EM

FB

MOR(SP)

ndashGates

etal(20

09)

Wetscleroph

yllforest(Eo

bliqua)

Tas

++(2ndash3)

All

EM

FB

MOR(SP)

ndashRatkowskyan

dGates

(200

9)Wetscleroph

yllforest(Eo

bliqua)

Tas

ndash+(2ndash3)

+(70)

All

EM

FB

MOR(SP)

ndashGates

etal(20

05)

Wetscleroph

yllforest(Eo

bliqua)

Tas

ndashndash

+(73

10920

0ndash30

0)All

EM

FB

MOR(SP)

ndashRatko

wskyandGates

(200

8)G

ates

etal

(201

0a2

010b)

Wetscleroph

yllforest(Eregnans)

Tas

++(24

7)

+(57)

All

EM

FB

MOR(SP)

ndashMcM

ullan-Fisheretal

(200

2)Wetscleroph

yllforest(Eregnans)

Vic

+(2

mon

ths)

+(14

25mon

ths)

ndashM

(ECM)

SF

SO

MOR(M

G)

BAB

M(ERG)

Launo

nenetal(19

99)

Sclerop

hyllforest(Ed

iversicolor

regrow

thforest)

WA

++

+(17ndash25

)All

EM

FB

MOR(SP)

ndashRob

insonan

dBou

gher

(200

3)R

obinsonetal

(200

8)Sclerop

hyllforest(Em

arginata)

WA

++(6)

+(45)

MSF

SC

MOR(M

G)

RT

Malajczuk

andHingston

(198

1)Sclerop

hyllforest(Em

arginata)

WA

ndash+(2ndash4)

+(28ndash30

)All

EM

FB

MOR(SP)

BM

(WEI)

ndashTom

merup

etal(20

00)

Sclerop

hyllforest(Em

arginata)

WA

ndash+

+All

EM

FB

MOR(SP)

ndashHilton

etal(19

89)

Sclerop

hyllforest(Em

arginata

Corym

biacaloph

ylla)

WA

ndash+(6ndash7)

+(66)

M(ECM)

SQE

MFB

MOR(SP)

MOL(RFLP)

Glenetal(20

01)

Dry


aculata)

SA

++(23

)ndash

All

EM

(Disc)

FB

MOR(SP)

ndashWarcup(199

0)Sclerop

hyllwoo

dland(Ecladocalyx

Eb

axteri)

SA

+ndash

ndashAll

SQE

MFB

MOR(SP)

ndashGeorge(200

8)C

atcheside

(200

9)C

atchesideetal

(200

9)R

obinson

(200

9)Low

open

drysclerophyllwoo

dland

(Eb

axteriE

obliqua)

SA

+(12

7mon

ths)

+(12and

20mon

ths)

ndashS

SF

SOC

UMOR(M

G)

ndashTheod

orou

andBow

en(198

2)Dry

sclerophyllwoo

dland(Erossii

Em

acrorhyncha)

ACT

++(4)

+(21)

All

SQE

MFB

MOR(SP)

ndashTrapp

eetal(20

06)

Woodland(Ea

lbensCallitris

glau

cophylla)riparian

strips

(Eviminalis)grassy

orshrubb

ywoo

dland(Em

ellio

dora)

woo

dland(En

ortonii)gradingto

open

forest(Ed

alrympleana)

NSW

++

ndashAll

EM

FB

MOR(SP)

ndashClaridg

eetal(20

09a)


Sclerop

hyllshrubland(Ang

opho

rahispida)

NSW

++(8)

+M

(AM)

SF

SS

MOR(M

G)


94)

Dry

open

sclerophyllforests

(Eg

ummiferaS

yncarpia

glom

uliferaE

squ

amosa

EresiniferaE

haemastoma

Eb

auerana

Allo

casuarina

littoralisA

ngop

hora

hispida)

NSW

+(2

weeks)

+(81

0)+(25)

All

SF

SO

MOL(RFLP

DNA)

ndashChenandCairney

(200

2)

Sclerop

hyllforest(Ep

ilularis)and

mixed

eucalypt

forest(Eo

bliqua

Erub

ida

Erad

iata)

NSWV

ic

ndash+(31

0)

+(gt60

)All

SF

FBS

OMOR(M

G)

MOL(IS

RFLP

T-RFLP)

BM

(ERG)

Osborn(200

7)

Sclerop

hyllforest(Ep

ilularis)

Qld

ndash+(2

4)

+(31ndash34

)SM

(ECM)

SF

SOH

IMOL(D

GGE

RFLPT

-RFLP

DNAS

IR)

BM

(PLFA)

GLP

OIS

Bastia

setal(20

06a

2006b)A

ndersonetal

(200

7)C

ampb

elletal

(200

8)A

rtzetal(200

9)Sclerop

hyllforest(Ep

ilularis)

NSW

+(48h)

ndash+

SMI(Tric)

CU

MOR(SP)

ndashMcG

eeetal(20

06)

TropicalAllo

casuarinaforest

Eucalyptuswoo

dlandandtropical

rainforest

Qld

++(4ndash5)

ndashM

(ECM)

SQ

FBS

PMOR(SP)

BM

(WEI)

Vernesan

dHaydon

(200

1)V

ernesetal

(200

120

04)

Tropicalsavann

asNT

++

+M

(ECM

andAM)

SF

SOS

SMOR(M

G)

BA

Brund

rettetal(19

96a

1996b)

Tem


australisP

oasieberiana)

NSW

ndash+(24

8)

+L

CS

FB

MOR(SP)

ndashOrsquoBryan

etal(20

09)

Tussock

grasslandandhummock

sedg

eland

Tas

+(3

mon

ths)

+L

LFB

MOR(SP)

ndashFergu

sonetal(20

09)

Alpinewoo

dland(Eucalyptus

pauciflora

Ed

alrympleana)tall

wetscleroph

yllforest(Eregnans

Efastig

ata)d

rysclerophyllforest

(Em

anniferaE

macrorhyncha)

Vicand

NSW

++(gt1ndash10

)+(11ndash20

21ndash30

gt30

)M

(ECM)

SQ

FB

MOR(SP)

ndashClaridg

ean

dBarry

(200

0)C

laridg

eetal

(200

0a2

000b)

Claridg

ean

dTrapp

e(200

4)Alpineheath

Tas

ndashndash

+(39

56)

All

EM

FB

MOR(SP)

ndashMcM

ullan-Fisheretal

(200


verrucosa)

NSW

++(34

7)

+(13

161

835

100

)L

CS

FB

MOR(SP)

ndashEldridg

eandBradstock

(199

4)

Referencesinbo

ldareforstud

iesthathave

replicated


licsareforstud

ieswith

replicated

samples

with

inun

replicated

fireeventsA

bbreviationsT

rophicgroupAll=all

trophicgroups

except

lichensL

=lichensM


=ectomycorrhizasA

M=arbu

scular


sMorph

olog

icalgrou

pCS=cryp

togamicsoilcrusts

EM

=epigeous






redFB=fruit-body

surveyH

I=myceliumfrom

hyph

alingrow


sfrom

soilsampleSO=bu


from


soilFun

galidentificatio

ntechniqu

eMOL=identifi

catio

noftaxa

bymolecularmethods



NA=DNAsequencing





MOR=identifi

catio

noftaxa

from

morph


identifi

edto


edtobroadmorph

ogroups)E

stim

ateof

fung

alactiv

ityB

A=Bioassayof

extentof


nof

seedlin

gsB

M=fungalbiom

ass(m

etho

dERG=ergo

sterol

analysisPLFA


fatty

acidsWEI=


diversity

oflaccasegenesIS


ityR


root

tips




(a) (b)

(c) (d )

(e ) (f )

















(a) (b)

(d )(c )

(e ) (f)












Saprotrophic fungi

















Mycorrhizal fungi









Lichenised fungi


































Conclusions






Acknowledgements


References
























































































































































































































































Pyrophilous fungi




Tab

le1

Stud

iesof

fung

alcommun

itiesin


know

nfire

history

Multip

lestud

ieson


ireages

with

anasterisk


dergon

erepeated

burns

Vegetationtype

(dom

inantvegetation)

State

Sites(age

sincefireinyears)

Fun

gigrou

psstud

ied

Sam

plingandidentifi

catio

ntechniqu

esandor

estim

ationof

fungalactiv

ityReference

Fire0ndash12

mon

ths

Fire

gt1ndash10

years

Mature

Trophic

group

Morph

o-logical

group

Sam

ple

collection

techniqu

e

Identifi

catio

ntechnique

(metho

dor

scop

e)

Estim

ateof

fungalactiv

ity(m

etho

d)

Wetscleroph

yllforest(Eo

bliqua

andor

Ed

elegatensis)

Tas

ndashndash

+(25ndash30

gt1

00)

All

EM

FB

MOR(SP)

ndashPackh

ametal(20

02)

Wetscleroph

yllforest(Eo

bliqua)

Tas

++(3)

+(65ndash10

1)L

LFB

MOR(SP)

ndashKan

tvila

san

dJa

rman

(200

6)Wetscleroph

yllforest(Eo

bliqua)

Tas

++

+(73)

All

EM

FB

MOR(SP)

ndashGates

etal(20

09)

Wetscleroph

yllforest(Eo

bliqua)

Tas

++(2ndash3)

All

EM

FB

MOR(SP)

ndashRatkowskyan

dGates

(200

9)Wetscleroph

yllforest(Eo

bliqua)

Tas

ndash+(2ndash3)

+(70)

All

EM

FB

MOR(SP)

ndashGates

etal(20

05)

Wetscleroph

yllforest(Eo

bliqua)

Tas

ndashndash

+(73

10920

0ndash30

0)All

EM

FB

MOR(SP)

ndashRatko

wskyandGates

(200

8)G

ates

etal

(201

0a2

010b)

Wetscleroph

yllforest(Eregnans)

Tas

++(24

7)

+(57)

All

EM

FB

MOR(SP)

ndashMcM

ullan-Fisheretal

(200

2)Wetscleroph

yllforest(Eregnans)

Vic

+(2

mon

ths)

+(14

25mon

ths)

ndashM

(ECM)

SF

SO

MOR(M

G)

BAB

M(ERG)

Launo

nenetal(19

99)

Sclerop

hyllforest(Ed

iversicolor

regrow

thforest)

WA

++

+(17ndash25

)All

EM

FB

MOR(SP)

ndashRob

insonan

dBou

gher

(200

3)R

obinsonetal

(200

8)Sclerop

hyllforest(Em

arginata)

WA

++(6)

+(45)

MSF

SC

MOR(M

G)

RT

Malajczuk

andHingston

(198

1)Sclerop

hyllforest(Em

arginata)

WA

ndash+(2ndash4)

+(28ndash30

)All

EM

FB

MOR(SP)

BM

(WEI)

ndashTom

merup

etal(20

00)

Sclerop

hyllforest(Em

arginata)

WA

ndash+

+All

EM

FB

MOR(SP)

ndashHilton

etal(19

89)

Sclerop

hyllforest(Em

arginata

Corym

biacaloph

ylla)

WA

ndash+(6ndash7)

+(66)

M(ECM)

SQE

MFB

MOR(SP)

MOL(RFLP)

Glenetal(20

01)

Dry


aculata)

SA

++(23

)ndash

All

EM

(Disc)

FB

MOR(SP)

ndashWarcup(199

0)Sclerop

hyllwoo

dland(Ecladocalyx

Eb

axteri)

SA

+ndash

ndashAll

SQE

MFB

MOR(SP)

ndashGeorge(200

8)C

atcheside

(200

9)C

atchesideetal

(200

9)R

obinson

(200

9)Low

open

drysclerophyllwoo

dland

(Eb

axteriE

obliqua)

SA

+(12

7mon

ths)

+(12and

20mon

ths)

ndashS

SF

SOC

UMOR(M

G)

ndashTheod

orou

andBow

en(198

2)Dry

sclerophyllwoo

dland(Erossii

Em

acrorhyncha)

ACT

++(4)

+(21)

All

SQE

MFB

MOR(SP)

ndashTrapp

eetal(20

06)

Woodland(Ea

lbensCallitris

glau

cophylla)riparian

strips

(Eviminalis)grassy

orshrubb

ywoo

dland(Em

ellio

dora)

woo

dland(En

ortonii)gradingto

open

forest(Ed

alrympleana)

NSW

++

ndashAll

EM

FB

MOR(SP)

ndashClaridg

eetal(20

09a)


Sclerop

hyllshrubland(Ang

opho

rahispida)

NSW

++(8)

+M

(AM)

SF

SS

MOR(M

G)


94)

Dry

open

sclerophyllforests

(Eg

ummiferaS

yncarpia

glom

uliferaE

squ

amosa

EresiniferaE

haemastoma

Eb

auerana

Allo

casuarina

littoralisA

ngop

hora

hispida)

NSW

+(2

weeks)

+(81

0)+(25)

All

SF

SO

MOL(RFLP

DNA)

ndashChenandCairney

(200

2)

Sclerop

hyllforest(Ep

ilularis)and

mixed

eucalypt

forest(Eo

bliqua

Erub

ida

Erad

iata)

NSWV

ic

ndash+(31

0)

+(gt60

)All

SF

FBS

OMOR(M

G)

MOL(IS

RFLP

T-RFLP)

BM

(ERG)

Osborn(200

7)

Sclerop

hyllforest(Ep

ilularis)

Qld

ndash+(2

4)

+(31ndash34

)SM

(ECM)

SF

SOH

IMOL(D

GGE

RFLPT

-RFLP

DNAS

IR)

BM

(PLFA)

GLP

OIS

Bastia

setal(20

06a

2006b)A

ndersonetal

(200

7)C

ampb

elletal

(200

8)A

rtzetal(200

9)Sclerop

hyllforest(Ep

ilularis)

NSW

+(48h)

ndash+

SMI(Tric)

CU

MOR(SP)

ndashMcG

eeetal(20

06)

TropicalAllo

casuarinaforest

Eucalyptuswoo

dlandandtropical

rainforest

Qld

++(4ndash5)

ndashM

(ECM)

SQ

FBS

PMOR(SP)

BM

(WEI)

Vernesan

dHaydon

(200

1)V

ernesetal

(200

120

04)

Tropicalsavann

asNT

++

+M

(ECM

andAM)

SF

SOS

SMOR(M

G)

BA

Brund

rettetal(19

96a

1996b)

Tem


australisP

oasieberiana)

NSW

ndash+(24

8)

+L

CS

FB

MOR(SP)

ndashOrsquoBryan

etal(20

09)

Tussock

grasslandandhummock

sedg

eland

Tas

+(3

mon

ths)

+L

LFB

MOR(SP)

ndashFergu

sonetal(20

09)

Alpinewoo

dland(Eucalyptus

pauciflora

Ed

alrympleana)tall

wetscleroph

yllforest(Eregnans

Efastig

ata)d

rysclerophyllforest

(Em

anniferaE

macrorhyncha)

Vicand

NSW

++(gt1ndash10

)+(11ndash20

21ndash30

gt30

)M

(ECM)

SQ

FB

MOR(SP)

ndashClaridg

ean

dBarry

(200

0)C

laridg

eetal

(200

0a2

000b)

Claridg

ean

dTrapp

e(200

4)Alpineheath

Tas

ndashndash

+(39

56)

All

EM

FB

MOR(SP)

ndashMcM

ullan-Fisheretal

(200


verrucosa)

NSW

++(34

7)

+(13

161

835

100

)L

CS

FB

MOR(SP)

ndashEldridg

eandBradstock

(199

4)

Referencesinbo

ldareforstud

iesthathave

replicated


licsareforstud

ieswith

replicated

samples

with

inun

replicated

fireeventsA

bbreviationsT

rophicgroupAll=all

trophicgroups

except

lichensL

=lichensM


=ectomycorrhizasA

M=arbu

scular


sMorph

olog

icalgrou

pCS=cryp

togamicsoilcrusts

EM

=epigeous






redFB=fruit-body

surveyH

I=myceliumfrom

hyph

alingrow


sfrom

soilsampleSO=bu


from


soilFun

galidentificatio

ntechniqu

eMOL=identifi

catio

noftaxa

bymolecularmethods



NA=DNAsequencing





MOR=identifi

catio

noftaxa

from

morph


identifi

edto


edtobroadmorph

ogroups)E

stim

ateof

fung

alactiv

ityB

A=Bioassayof

extentof


nof

seedlin

gsB

M=fungalbiom

ass(m

etho

dERG=ergo

sterol

analysisPLFA


fatty

acidsWEI=


diversity

oflaccasegenesIS


ityR


root

tips




(a) (b)

(c) (d )

(e ) (f )

















(a) (b)

(d )(c )

(e ) (f)












Saprotrophic fungi

















Mycorrhizal fungi









Lichenised fungi


































Conclusions






Acknowledgements


References






































































































































































































































Tab

le1

Stud

iesof

fung

alcommun

itiesin


know

nfire

history

Multip

lestud

ieson


ireages

with

anasterisk


dergon

erepeated

burns

Vegetationtype

(dom

inantvegetation)

State

Sites(age

sincefireinyears)

Fun

gigrou

psstud

ied

Sam

plingandidentifi

catio

ntechniqu

esandor

estim

ationof

fungalactiv

ityReference

Fire0ndash12

mon

ths

Fire

gt1ndash10

years

Mature

Trophic

group

Morph

o-logical

group

Sam

ple

collection

techniqu

e

Identifi

catio

ntechnique

(metho

dor

scop

e)

Estim

ateof

fungalactiv

ity(m

etho

d)

Wetscleroph

yllforest(Eo

bliqua

andor

Ed

elegatensis)

Tas

ndashndash

+(25ndash30

gt1

00)

All

EM

FB

MOR(SP)

ndashPackh

ametal(20

02)

Wetscleroph

yllforest(Eo

bliqua)

Tas

++(3)

+(65ndash10

1)L

LFB

MOR(SP)

ndashKan

tvila

san

dJa

rman

(200

6)Wetscleroph

yllforest(Eo

bliqua)

Tas

++

+(73)

All

EM

FB

MOR(SP)

ndashGates

etal(20

09)

Wetscleroph

yllforest(Eo

bliqua)

Tas

++(2ndash3)

All

EM

FB

MOR(SP)

ndashRatkowskyan

dGates

(200

9)Wetscleroph

yllforest(Eo

bliqua)

Tas

ndash+(2ndash3)

+(70)

All

EM

FB

MOR(SP)

ndashGates

etal(20

05)

Wetscleroph

yllforest(Eo

bliqua)

Tas

ndashndash

+(73

10920

0ndash30

0)All

EM

FB

MOR(SP)

ndashRatko

wskyandGates

(200

8)G

ates

etal

(201

0a2

010b)

Wetscleroph

yllforest(Eregnans)

Tas

++(24

7)

+(57)

All

EM

FB

MOR(SP)

ndashMcM

ullan-Fisheretal

(200

2)Wetscleroph

yllforest(Eregnans)

Vic

+(2

mon

ths)

+(14

25mon

ths)

ndashM

(ECM)

SF

SO

MOR(M

G)

BAB

M(ERG)

Launo

nenetal(19

99)

Sclerop

hyllforest(Ed

iversicolor

regrow

thforest)

WA

++

+(17ndash25

)All

EM

FB

MOR(SP)

ndashRob

insonan

dBou

gher

(200

3)R

obinsonetal

(200

8)Sclerop

hyllforest(Em

arginata)

WA

++(6)

+(45)

MSF

SC

MOR(M

G)

RT

Malajczuk

andHingston

(198

1)Sclerop

hyllforest(Em

arginata)

WA

ndash+(2ndash4)

+(28ndash30

)All

EM

FB

MOR(SP)

BM

(WEI)

ndashTom

merup

etal(20

00)

Sclerop

hyllforest(Em

arginata)

WA

ndash+

+All

EM

FB

MOR(SP)

ndashHilton

etal(19

89)

Sclerop

hyllforest(Em

arginata

Corym

biacaloph

ylla)

WA

ndash+(6ndash7)

+(66)

M(ECM)

SQE

MFB

MOR(SP)

MOL(RFLP)

Glenetal(20

01)

Dry


aculata)

SA

++(23

)ndash

All

EM

(Disc)

FB

MOR(SP)

ndashWarcup(199

0)Sclerop

hyllwoo

dland(Ecladocalyx

Eb

axteri)

SA

+ndash

ndashAll

SQE

MFB

MOR(SP)

ndashGeorge(200

8)C

atcheside

(200

9)C

atchesideetal

(200

9)R

obinson

(200

9)Low

open

drysclerophyllwoo

dland

(Eb

axteriE

obliqua)

SA

+(12

7mon

ths)

+(12and

20mon

ths)

ndashS

SF

SOC

UMOR(M

G)

ndashTheod

orou

andBow

en(198

2)Dry

sclerophyllwoo

dland(Erossii

Em

acrorhyncha)

ACT

++(4)

+(21)

All

SQE

MFB

MOR(SP)

ndashTrapp

eetal(20

06)

Woodland(Ea

lbensCallitris

glau

cophylla)riparian

strips

(Eviminalis)grassy

orshrubb

ywoo

dland(Em

ellio

dora)

woo

dland(En

ortonii)gradingto

open

forest(Ed

alrympleana)

NSW

++

ndashAll

EM

FB

MOR(SP)

ndashClaridg

eetal(20

09a)


Sclerop

hyllshrubland(Ang

opho

rahispida)

NSW

++(8)

+M

(AM)

SF

SS

MOR(M

G)


94)

Dry

open

sclerophyllforests

(Eg

ummiferaS

yncarpia

glom

uliferaE

squ

amosa

EresiniferaE

haemastoma

Eb

auerana

Allo

casuarina

littoralisA

ngop

hora

hispida)

NSW

+(2

weeks)

+(81

0)+(25)

All

SF

SO

MOL(RFLP

DNA)

ndashChenandCairney

(200

2)

Sclerop

hyllforest(Ep

ilularis)and

mixed

eucalypt

forest(Eo

bliqua

Erub

ida

Erad

iata)

NSWV

ic

ndash+(31

0)

+(gt60

)All

SF

FBS

OMOR(M

G)

MOL(IS

RFLP

T-RFLP)

BM

(ERG)

Osborn(200

7)

Sclerop

hyllforest(Ep

ilularis)

Qld

ndash+(2

4)

+(31ndash34

)SM

(ECM)

SF

SOH

IMOL(D

GGE

RFLPT

-RFLP

DNAS

IR)

BM

(PLFA)

GLP

OIS

Bastia

setal(20

06a

2006b)A

ndersonetal

(200

7)C

ampb

elletal

(200

8)A

rtzetal(200

9)Sclerop

hyllforest(Ep

ilularis)

NSW

+(48h)

ndash+

SMI(Tric)

CU

MOR(SP)

ndashMcG

eeetal(20

06)

TropicalAllo

casuarinaforest

Eucalyptuswoo

dlandandtropical

rainforest

Qld

++(4ndash5)

ndashM

(ECM)

SQ

FBS

PMOR(SP)

BM

(WEI)

Vernesan

dHaydon

(200

1)V

ernesetal

(200

120

04)

Tropicalsavann

asNT

++

+M

(ECM

andAM)

SF

SOS

SMOR(M

G)

BA

Brund

rettetal(19

96a

1996b)

Tem


australisP

oasieberiana)

NSW

ndash+(24

8)

+L

CS

FB

MOR(SP)

ndashOrsquoBryan

etal(20

09)

Tussock

grasslandandhummock

sedg

eland

Tas

+(3

mon

ths)

+L

LFB

MOR(SP)

ndashFergu

sonetal(20

09)

Alpinewoo

dland(Eucalyptus

pauciflora

Ed

alrympleana)tall

wetscleroph

yllforest(Eregnans

Efastig

ata)d

rysclerophyllforest

(Em

anniferaE

macrorhyncha)

Vicand

NSW

++(gt1ndash10

)+(11ndash20

21ndash30

gt30

)M

(ECM)

SQ

FB

MOR(SP)

ndashClaridg

ean

dBarry

(200

0)C

laridg

eetal

(200

0a2

000b)

Claridg

ean

dTrapp

e(200

4)Alpineheath

Tas

ndashndash

+(39

56)

All

EM

FB

MOR(SP)

ndashMcM

ullan-Fisheretal

(200


verrucosa)

NSW

++(34

7)

+(13

161

835

100

)L

CS

FB

MOR(SP)

ndashEldridg

eandBradstock

(199

4)

Referencesinbo

ldareforstud

iesthathave

replicated


licsareforstud

ieswith

replicated

samples

with

inun

replicated

fireeventsA

bbreviationsT

rophicgroupAll=all

trophicgroups

except

lichensL

=lichensM


=ectomycorrhizasA

M=arbu

scular


sMorph

olog

icalgrou

pCS=cryp

togamicsoilcrusts

EM

=epigeous






redFB=fruit-body

surveyH

I=myceliumfrom

hyph

alingrow


sfrom

soilsampleSO=bu


from


soilFun

galidentificatio

ntechniqu

eMOL=identifi

catio

noftaxa

bymolecularmethods



NA=DNAsequencing





MOR=identifi

catio

noftaxa

from

morph


identifi

edto


edtobroadmorph

ogroups)E

stim

ateof

fung

alactiv

ityB

A=Bioassayof

extentof


nof

seedlin

gsB

M=fungalbiom

ass(m

etho

dERG=ergo

sterol

analysisPLFA


fatty

acidsWEI=


diversity

oflaccasegenesIS


ityR


root

tips




(a) (b)

(c) (d )

(e ) (f )

















(a) (b)

(d )(c )

(e ) (f)












Saprotrophic fungi

















Mycorrhizal fungi









Lichenised fungi


































Conclusions






Acknowledgements


References






































































































































































































































Sclerop

hyllshrubland(Ang

opho

rahispida)

NSW

++(8)

+M

(AM)

SF

SS

MOR(M

G)


94)

Dry

open

sclerophyllforests

(Eg

ummiferaS

yncarpia

glom

uliferaE

squ

amosa

EresiniferaE

haemastoma

Eb

auerana

Allo

casuarina

littoralisA

ngop

hora

hispida)

NSW

+(2

weeks)

+(81

0)+(25)

All

SF

SO

MOL(RFLP

DNA)

ndashChenandCairney

(200

2)

Sclerop

hyllforest(Ep

ilularis)and

mixed

eucalypt

forest(Eo

bliqua

Erub

ida

Erad

iata)

NSWV

ic

ndash+(31

0)

+(gt60

)All

SF

FBS

OMOR(M

G)

MOL(IS

RFLP

T-RFLP)

BM

(ERG)

Osborn(200

7)

Sclerop

hyllforest(Ep

ilularis)

Qld

ndash+(2

4)

+(31ndash34

)SM

(ECM)

SF

SOH

IMOL(D

GGE

RFLPT

-RFLP

DNAS

IR)

BM

(PLFA)

GLP

OIS

Bastia

setal(20

06a

2006b)A

ndersonetal

(200

7)C

ampb

elletal

(200

8)A

rtzetal(200

9)Sclerop

hyllforest(Ep

ilularis)

NSW

+(48h)

ndash+

SMI(Tric)

CU

MOR(SP)

ndashMcG

eeetal(20

06)

TropicalAllo

casuarinaforest

Eucalyptuswoo

dlandandtropical

rainforest

Qld

++(4ndash5)

ndashM

(ECM)

SQ

FBS

PMOR(SP)

BM

(WEI)

Vernesan

dHaydon

(200

1)V

ernesetal

(200

120

04)

Tropicalsavann

asNT

++

+M

(ECM

andAM)

SF

SOS

SMOR(M

G)

BA

Brund

rettetal(19

96a

1996b)

Tem


australisP

oasieberiana)

NSW

ndash+(24

8)

+L

CS

FB

MOR(SP)

ndashOrsquoBryan

etal(20

09)

Tussock

grasslandandhummock

sedg

eland

Tas

+(3

mon

ths)

+L

LFB

MOR(SP)

ndashFergu

sonetal(20

09)

Alpinewoo

dland(Eucalyptus

pauciflora

Ed

alrympleana)tall

wetscleroph

yllforest(Eregnans

Efastig

ata)d

rysclerophyllforest

(Em

anniferaE

macrorhyncha)

Vicand

NSW

++(gt1ndash10

)+(11ndash20

21ndash30

gt30

)M

(ECM)

SQ

FB

MOR(SP)

ndashClaridg

ean

dBarry

(200

0)C

laridg

eetal

(200

0a2

000b)

Claridg

ean

dTrapp

e(200

4)Alpineheath

Tas

ndashndash

+(39

56)

All

EM

FB

MOR(SP)

ndashMcM

ullan-Fisheretal

(200


verrucosa)

NSW

++(34

7)

+(13

161

835

100

)L

CS

FB

MOR(SP)

ndashEldridg

eandBradstock

(199

4)

Referencesinbo

ldareforstud

iesthathave

replicated


licsareforstud

ieswith

replicated

samples

with

inun

replicated

fireeventsA

bbreviationsT

rophicgroupAll=all

trophicgroups

except

lichensL

=lichensM


=ectomycorrhizasA

M=arbu

scular


sMorph

olog

icalgrou

pCS=cryp

togamicsoilcrusts

EM

=epigeous






redFB=fruit-body

surveyH

I=myceliumfrom

hyph

alingrow


sfrom

soilsampleSO=bu


from


soilFun

galidentificatio

ntechniqu

eMOL=identifi

catio

noftaxa

bymolecularmethods



NA=DNAsequencing





MOR=identifi

catio

noftaxa

from

morph


identifi

edto


edtobroadmorph

ogroups)E

stim

ateof

fung

alactiv

ityB

A=Bioassayof

extentof


nof

seedlin

gsB

M=fungalbiom

ass(m

etho

dERG=ergo

sterol

analysisPLFA


fatty

acidsWEI=


diversity

oflaccasegenesIS


ityR


root

tips




(a) (b)

(c) (d )

(e ) (f )

















(a) (b)

(d )(c )

(e ) (f)












Saprotrophic fungi

















Mycorrhizal fungi









Lichenised fungi


































Conclusions






Acknowledgements


References








































































































































































































































(a) (b)

(c) (d )

(e ) (f )

















(a) (b)

(d )(c )

(e ) (f)












Saprotrophic fungi

















Mycorrhizal fungi









Lichenised fungi


































Conclusions






Acknowledgements


References




















































































































































































































































(a) (b)

(d )(c )

(e ) (f)












Saprotrophic fungi

















Mycorrhizal fungi









Lichenised fungi


































Conclusions






Acknowledgements


References















































































































































































































































Saprotrophic fungi

















Mycorrhizal fungi









Lichenised fungi


































Conclusions






Acknowledgements


References




















































































































































































































































Mycorrhizal fungi









Lichenised fungi


































Conclusions






Acknowledgements


References







































































































































































































































Lichenised fungi


































Conclusions






Acknowledgements


References





























































































































































































































































Conclusions






Acknowledgements


References









































































































































































































































Acknowledgements


References






































































































































































































































fungi and fire in australian ecosystems: a review of ... · fungi and ﬁre in australian...

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