CHAPTER3 MushroomsinTheir Natural Habitats

ushroomscan be placedin 4 basiccategories plantsoverhead onto the forestfloor, the decomposers saprophytic,parasitic, myconhizal, and endo residingin the soil processthis newly availablefood. phytic, dependingupon how they nourish themseives. (Competitionis intense:on the forestfloor, a single However, exceptions abound, since some species "habitat"can actuallybe matricesof fungalnetworks employ more than one strategy,rraking them difficrrlt sharingone space.) These fungi secrete enzymes and to categorize.Approximately 8,000 macrofungi (visibie acidsthat degradelarge molecules of deadplants into b the naked eye) are saprophytic, around 2,000 to sirnplermolecules, which the fungi can reassemble 3,()00 are rnycorrhizal, ancl the remaining are either inio buildingblocks, such as polysaccharides, for cell cnclophl4icor parasitic,although more speciesare con- walls.From deadplants, fungi recyclecarbon, hydro- stantl1,beingdiscovered and categorized.The balance gen,nitrogen, phosporus, and mineralsinto nutrients of pr.rpulationscan vary drasticallywith environmental for livingplants, insects, and other organisms shanng cliange, however: deforestationcauses a rise in sapro- ihat 1'rabitat. phrtcs and a clecline in mycorrhizal mushrooms, for cxarnplc.Now let'stake a shorttour through the 4 major c;rtcgoriesof mushrooms.

SaprophyticMushrooms: TheDecomposers

5;r1:xrplrr.ticmushrooms, the decomposers,steer the t:ourscfirr proliFcratingbiologicai communities,shap- irrg ancl torrtring tlre first menus in tl-refood web from clcnr.lplirrrti. insects, ancl other anirnals.Most gourmet lunrlrrri:tlicinal nlrshrooms :rre wood decornposers,the prctrticrrccvclers orr Lheplanet; building soils is the prttn;rn.outconrc of thc activitiesof thesesaprophytic ;1,FIGURE 28 filrtqi, lllur.sc filanrentous mycelial nehnrorksweave Turkeytail (Trametes versiclllr)fruiring ona coniferlog deep tltrrritghlncl bctn'ecnthe cell walls of plants.When inold-growth forest in 0lympic National Park. orqiltric lrrattcr fulls fronr the canopy of trees and

19 MYCELIUMRUNNING

FIGURE29

Thesetowertng old'growthtrees nearMount Fainier,grow out ofthin soil but gathernutrlents froma{ar from therrmycellum- supportedroots lnfact, most plantsare suP- portedbY vast andcomPlex coloniesoffungt workingincon- cert.Here I Point Io BndgeaPorus nobilissimus (fora closerview & lru,t seefigure 50), a mushroom exclusiveto old- growthhabitat andthe first fun- gusto be listed asan endan- geredsPecies MushroomsinTheir Natural Habitats 2l

As decomposers,saprophytic mushrooms can be SecondaryDecomposers separatedinto 3 key groups:primary, secondary and Secondarydecomposers rely on the activityof primary tertiary,although some mushroomspecies can cross fungi that initially, although partially,break down over fiom one categoryto another, dependingupon plant and animal tissues.Secondary decomposers all circumstances.Primary secondary and tertiary decom- work in concertwith actinomycetes,other bacteria, poserscan all coexistin onelocation. Primary and sec- andfungi, including yeasts, in soilin the forestfloor or ondary decomposerssuch as oyster and meadow in compostpiles. Heat, water, carbon dioxide, ammo- mushroomsare the easiestto cultivate. nia, and othergases are emittedas by-products of the compostingprocess. Once the microorganisms(espe- PrimaryDecomposers cially actinomycetes)in the compostpiles complete Thesesaprophytes are typicallythe first to grow on a their life cycles,the temperaturedrops, encouraging a twig, a bladeof grass,a chip of wood,a log, a sturrrp, new waveofsecondary decomposers. a dead insect or other animal. Primary or decom- Cultivatorsexploit this sequence to growthe white are typicallyfast growing, sending posers out rapidly button mushroom(Agaicus bisporus),the mostwidely strandsof mycelium that extending quickly attachto cultivatedmushroom in the world. Other secondarv anddecompose plant tissue. These woodland species irrcltrdeoyster mushro oms (Pleurotus species), shiitake (Lentinula edodes),and maitake(Grifola frondosa). Y FIGURE30 However,species employ different setsof enzymes DavidArora, author of MushroomsDemystified and All down plant matter into io break varying stagesof Thatthe Bain Promises and Mlre is positionedt0 take a decornposition. photographofa familyof ambiguousStropharias, Stropharia ambigua,near my home. MYCELIUMRUNNING mush- saprophytesthat competewith compost-grown family Copri- ,-r.ti ,.. inky caps(belonging to the edible shaggy naceae,which includes the choice' others including the mane fCoprinus comatusland and Panaeolus i"tttt"inog"ni c Panaeolussubbalteatus chip beds' the ,yonnrru*), and, in outdoor wood lndts- ,*big,to,-t, Stropharia (Stropharia ambigua)' theseundesired invaders by trial irowerstry to thwart inhos- heat steamingtheir composisto temperatures Ditabieto their sPores' seemmore Secondarydecomposers, as a group' for dealingwith versatilethan pnmary decomposers since they complex arrortmerttsof microorganisms' rich have evoivedin direct contactwith microbially trom com- soils.Secondary decomposers typically grow species' postedmaterial. The bestculinary Stropharia (Stropharia i. th" g"rd.n giant, or king Stiopharia' intermediary *soro ornulata) is an exampleof an slnce b"tt..., primary and secondarydecomposers i then conttn- this speciesfirst digestsfresh debris and ,i of microbesioin ,il uesto thriveas complex communities ; FIGURE31 : with it to createsoil. thegarden gtant +i Mvdauqhter, LaDena Stamets, sits beside isdeep burgundy rncolor tsi*iii,i, rugosoannulatal' which achievinga ;ily;;;s u"ndfud.t as it matures'sometimes a primaryand a rulri,, t,u,u".This mushroom can be both soilmicrobes ,.Jo.lututuottphyte but is dependent upon for fruiting

r FIGURE32 brsPorus/cu ltivatton ;\ FIGUBE33 Commercialbutton mushroom (Aga ri c u s Thismushroom isa classicsecondarY saProPhYte, froma stump' inHolland. Thehoney mushroom (Armillaria ostoyae/fruiting growingon compost Mushroomsin lheir NaturalHabitats 23

TertiaryDecomposers to be 1,500years old. In Oregon,a far largerhoney This difficult-to-categorizegroup includes fungi mushroom(Armillaria ostoyae) mycelial mat found on found towardthe end of the decompositionprocess. a mountaintopcovers more than 2,400 acresand is They thrive in habitatscreated by primary and sec- possiblymore than 2,200 years old (seefigure 60). Each ondarydecomposers over a periodofyears, often pop- time thisfungus blight sweepsthrough, nurse logs are ping up from soils holding little decomposable created,soil depth increases, and centimetersof soil material.Tertiary decomposers include speciesof accumulateto createever-richer habitats where orrce Conocybe,Agrocybe, Mycena, Pluteus, and Agaricus. onlyltt.l barren rock stood. (For fuither discussionof Tertiary decomposersrely upon highly con-rplex Armillaria blights,see page 47.) What makesmush- microbialenvironments. The divisionbetweerr sec- room myceliadifferent from the myceliafrom mold ondary and tertiary decomposersis often obscure; fungi is that somemushroom species can grow into mycologistssimply call tertiary decomposers"soil massivemembranes, thousands of acresin size,hun- dwellers,"for lackof a betterdescription. Some mush- dredsof tonsin mass,and thousandsof yearsold. rooms initially act as parasites,and once tl-reyhave Many saprophyticfungi can be weaklyparasitic, killedtheir hosts,they act like saprophytes,growing on especiallyifa hosttree is dying from othercauses, such their deadremains. Honey mushrooms belonging to as environmentalstress or parasiteinfestation. Sapro- tlreger-rus Amillaria aregood examples of speciesthat phytesthat can take advantageof a dying tree are growboih parasiticallyand saprophytically. termedfacultatite parasites. For example, oyster mush- rooms (Pleurotusostreatus) are classicsaprophytes, ParasiticMushrooms: Blights of the althoughthey are frequentlyfound on dying cotton- Forestor Agents for HabitatRestoration? wood,oak, poplar, birch, maple,and aldertrees. And althoughreishi (Ganoderma lucidum) is considereda Parasitesare predators that endar-rgerthe host'shealth. true saprophyteby most mycologists,the Australian In the past,foresters saw all parasiticfungi ashostile to QuarantineInspection Service has classifiedthis the iong-tern-rhealth of forests.Although they do para- medicinalspecies as a parasiteand has banned its sitize trees,they nourish other organisms.Parasitic importation.Authorities on otherislands ir-rcluding New fr-rngistrci.r as the honeynushroom, which can destroy Zealandand Hawaii also considerthis mushroorrra thotrsanclsof acresof forest,are stigmatized as blights. threatto theirnative trees. Some parasitic fungi behave , However,rnore foresters are realizing that a rottingtree like saprophytes,such as honey mushrooms (Armillaria in the nidst of zrcanopied forest is, in fact,more sup- mellea and Armillaria ostoyae),which may be found portiveof biocliversitythan a livingtree. Parasitic mush- thrivingon the corpseoftheir treehost. :' :r rOonlsmat' be uatttre'sway of selectingthe strongest Most parasiticfungi, however,are microfungi, ,,, plalti andrepairing damaged habitats. Uitimately, par- barelyvisible to the nakedeye, but en massethey inflict :..,9*itic mushroorlsset the stagefor the revivalof weal<- cankersand lesionson tl-reshoots and leavesof trees. ,'. -elrcdhabitats that:rre too stressedto thrive. Oftentheir prominence in a middle-agedforest is symp- Otall thc parasiticblight mushrooms that are edi- tomatico[ otherimba]ances in the ecosystem,such as ble br huurarrs,the assortedhoney mushroons such acid rain, groundwaterpollution, and insectdamage. ..:.,, Armil/cria nrclleaand Armillaria ostoyaeare the best After a tree dies,parasitic fungi may inhabit the tree, knonrr.()rc urvcclialmat from a honeymushroom competingwith saprophytesfor dominance.Since the {Armillariubttlbos,t) made national headlines when a hostsfor some parasitescan be shortlived, natural speciincrr u asfrrurrtl in a Nzlichiganforest that covered selectionsometimes favors fast growers. Foresters have . -.3T.3crcs,ucighccl at lcast50 tons,and was estimated obsewedthis with Phytophthoraramorum, the causeof .. MYCELIUMRUNNING

FIGURE34 to Matsutake,which are myconhizal mushrooms known deepin the mycologistsas Tricholona magnivelare, growing old-growthforest of Washington State

myconhizae termfor thesefungi is vesiculararbuscular this (VAM). Both plant and mycorrhizaebenefit from grows association.Because ectomycorrhizal mycelium and beyondthe plant'sroots, it bringsdistant nutrients absorption moistureto the host plant' extendingthe zone well beyondthe root structureThe mycelium of nutrr- dramaticaliylncreases the plant'singestion elements ents,nttrogenoLls comPounds' and essential sur- (phosphorus,coPper' and zinc) as it decomposes thatthe ,o.u-tiingdebris- David Perry (1994) postulates myc- surfacearea-hence itsabsorption capability-of than the orrhizalfungi maybe 10to 100times greater the growth surfacearea ofleaves in a forestAs a result' mycor- of plant partnersis accelerated'Plants with far better rhizal fungalpartners can alsoresist diseases relation- than thosewithout. Fungi benefit from the to piant-secretedsug- can shipbecause it givesthem access oak disease;this downy mildew pathogen a: sr-rdden thatthe fr-ingiconvert to mannitols' ancestralforest in ars,mostly h."or", kill an ancient oak tree in daysand an ',: dead carcassesof arabitols.and erythritols few weeks,and remair-rviable on the a most excitingdiscoveries in the field platform for infectron One of the its victims, allowing a new staging transport of mycologyis that the mycorrhizaecan ftrrther into the forest' nutrientsto treesof differentspecies One mushroom a contin- speclescan connectmany acres of a forestin and researchers MycorrhizalMushrooms: Fungus .,ousnehtork of cells'In one experiment' myceliulr.r PlantPartnershiPs comparedthe flow of nutrientsvia the menziesii)' behveen3 trees:a Dougiasfir (Pseudotsuga mushrooms(myco means "mushroom"; Mycorrhizai papyrifera),and a westernrecl as matsutake' a paper birch (Betula ,hizol ^"nn "relatedto roots")'such gt plicata).TheDouglas fir andpaper birch form mutually beneficial uio boletus,and chanterelles, ""au cedarl-r:rcl sharedthe sameectomycorririza, while the with pinesand other plants' ln fact' most relationships (VAM) The researchers-coveLtcl Dor-rglasfirs havemycorrhizal an endomyconhiza plantsfrom grrrr., to lowerttrg the Douglasfir to simulatedeep shade' thus myceliaof fungalspecies that form exte- prrtr,"rr.The sugarsln respotrsc' plants are the tree'sability to photosynthesize ,ior sireathsaround t1-reroots of partner channeledsugars, tracked by radioac- The mycorrhizal fur-rgithat the mycorrhizae termecl ectomycorrhizal the birch to the root labeled tive carbon,from the rootzone of the interiorroot cellsofhost plantsare invade 9 percentof the net caLbt'tr the preferred zoneof the fir. More than endomycorrhizal,although currently luusnro0msIn lhetrNatural Habitats 25

c" FIGURE35 ,:ri FIGURE36

DustyYao happily holds her harvest ofwild porcinis, the JimGouin is pleased tofind these delicious matsutakes /Zr- mycorrhizalBoletus edulis, collected inthe mountains above cholomanagnivelare), a myconhizal mushroom, inthe moun- Telluride.Colorado. tainssomewhere within 200 miles of Seattle, Washinqton.

A FICURE37

basketawaits a bountiful *::lli Yt collectionofthese apricot-smelling chanrerelles, probably Cantharellus fornosus,a mycor- speciesgrowing Tt:lyt,ht99l ina 40-year-oldDouglas firforest near 0lympia, Washington. Mypractice isto pick no more than 01the mushroomsofa wildpatch, leaving young ones, and when encountering pairs of mushrooms, onlypick one of $em.Lnanterelles111tt::1t tendt0 form as twins, so cutting one mushroom near to the ground saves the other twin, allowing it to mature, spo{$late,and spread. MYCELIUMRUNNING

Simardexperin'rent showed that a common fir origir-ratedfrom the The courPolrndstransferred to tl-re under- n-ryce1ialnet couid unite J speciesof treesand cedar receivedonly a small frac- birch's roots,Ii4.rile the of mycorrhizal fungi: pro- scored a remarkabieabiligv of sugartransferred was directly tion. Tl're an'iount keepdiverse species oftrees in forests (Simard et af i997)' mycorrhizaecan to the amount of shading portional particularlyyounger trees strugglir-rg for sunlight' Arnebrant and others(1993) fed, L .nrli., stuclyby Kristir-ra saplings Nowwe havea betterunderstanding of how bidirectional transfer of nitrogen- shon,ed a similar of eldertrees that towe r overhead glutinosa) to survivein theshaclows nutrients fron-r aldet (Alnus based .pine out essentiallight The factthat a singlemyc- a shared ectomycorrhizal anclblock (Pit-tuscontorta) through 2 different orrhizalmushroom nutritionally supported myceliun-r. trees-onea coniferand the otherdeciduous-shows health' that the mycelittmguards the forest'soverall budgetingand multidirectionally allocating nutrients' Another exampleof a fungusand plant partner- shipis the matsutake, which hasa uniquerelationship plant *ith the non-chlorophyll-producingcandystick its (Allotropavirgata).The candystick gains virtually all western $.t.$milfimilD.J. Roail sugarsfrom the matsutakenycelium and the hemlockand/or Sitka spruce with r'vhichit assoctates (Hosfordet al. 1997;Trudeli et al' 2003) One rnycol- of matsu- ogistI knowspecuiates that the spotfiuitings on a siopeof Oregon's ttrk lTrirholomamagnivelare) intercor- Mount Hood may, in fact, be fiom a vast of nectedmyceiial colony extendingover thousands (Acer acres.A further example is tl-rebigleaf maple roots macrophyllum),which proiectsvinelike aerial rain that ascer-rdto the canopyof PacificNorthwest forestsand areteamed with mycorrhizae' for- Our understandingof the role of fungi in the held estis far moreadvanced than the simplisticviews justa few clecadesago Most ecologistsnow recognize that a forest'svitality is directlyrelated to the presence' A large abundance,and variety of myceiialassociates ttt portion,one-terrtl'r to one-fifth,of the total biomass Pacific th" toproilof a healthyDouglas fir forestin the evert Northlvestmay be rnadeup of n-rycelium'and and ector.ur- FIGURE38 moreif we includethe endomycorrhizae forest corrhizaethat thrive in the canopy l doubt a excellentbook, the roots of a youngplne 0nthe cover of this r'vithoutits fungi' withthe mycelium 0{the can be defined treelpinus sylvestris)are enveloped provedto thetree's Growing mycorrhizalmushrooms has rutfr.ot Suillusbovinus'The mycelium extends a than firstanticipated due to lhc rangefor absorbing nutrients and water while conterrlng be a greaterlhallenge pair- fungiplav a cLit- tunqatdefense against invasive diseases This symbiotic .o,til." interdependenciesin which ingis the n0rm in nature, n0t tne excepll0n i.al role.Or-rce the hurdle of establishingmycorrlrizal rvtuDtil uuillJ lil r!otut dr I toutLdtJ e,

FIGURE39

Bigleafmaples (Acer macrophyllum) grow in the rain forest of the0lympic lvlountains, inWashington State. Research by Cobbet al. (2001 ) showed that this maple extends r00ts 0n itsouter trunk that climb into the canopy, essentially creaung a biospherehigh above the forest floor. The biomass ofthese aerialr00ts is similar to the biomass ofthe subterranean roots.Up0n these aerial roots, a complexhabitat has evolved, includingmosses (nonvascular epiphytes) and licorice ferns lPolypodiunglycyrrhiza, a vascular epiphyte), once thought to beparasitic t0the tree but now known to bepart of the trees healthvecosvstem.

m1'celir-rrnl-ras been overcome,decades may pass beforea singlemushroom forms. Nuances of climate, qoilchemistry, and predominantmicroflora limit our . successin cultivating mycorrhizal mushroomsin - 'naturalsettings. The challengewe face is to tilt the balanceso that a speciesofour choicecan take up res- ;;l .iden..ir sucha complexnatural setting-to design it| FIGURE40 , ,habitatsin which it can grow. Speciesnative to a : : regionare more , likely than importedspecies to adapt ThePerigord Iruffle (Tuber melanosporum), isone of themost . feadilyto thesedesigned habitats. sought-afterandhighly regarded gourmet mushrooms inthe .::,, ; Manv American growershope for huge profits world.This mushroom ismycorrhizal, growing inassociation :.'..ilvhenthev tn to growEuropean truffles, mycorrhizal withfilberts and oak trees. tnrrsiiroonrsthat sell at veryhigh prices.In an attempt to dr-rplicrtr. the well-establishedtruffle orchardsin eralcompanies now markettruffle-inoculated trees for Sprrin. perigord -F1ancc, andItaly, where the renowned commercialuse, and calcareous (high in calcium)soils blacktrLrffle (firber melanosporum)fetches up to $500 in Texas,Washington, and Oregon have been suggested . per pound, dozensof growershave tried to cultivate as ideal sitesfor these.One company(www.truffle- ftonuativeFiuropean trufflesaround the American oaks tree.com)that seernson top of its gameconfirms that on thcir ,_-,.,4,l}berts land.Capitalizing on thisdesire, sev- the tree,inoculated with truffiemycorrhizae. is absent ' 1,. MYCELIUMRUNNING

patientlywaiting for 20 yearsuntil the (althoughit makes gibbosum),after o[competitorfungi before shipment commer- irst trufflescould be harvestedNevertheless' However'I knowof onlya few fr"-tt"t abouiyield)' gourmet mu5h1s6rn5has "" and one from cializationof mycorrhizal from North Carolina tniffle ,n.""rr"r-one little successoutside of the European have producedEuropean seen Northern Caiifornia-that thosein Franceand ltaly' a decadeof effort' In orchards,particulariy t ,-tffi"r,and only aftermore than of The realityis, though,that our nativespecies culturetechniques have increas- the past30 years tissue the foreign F'uro- tnrffle- mycorrhizaequickly outcompete *Oft."a the tradition of transplanting t"gft SinceEuropean truffles like basic(high d"u"lopment'most plant- oeantruffles. *pp"ut"* trees.Despite ihi' competi- pH; ,oit, the additionof calciumdiminishes of Europeantrufifles beyond their il; in"oculations but this alonewill not ", Showing tion fro* nativemushrooms, habit"t still fail to producemushrooms' ,raliu" In New Zea\and,where the repertotre far more successfulthan assuresuccess' that giowingnative species is just a few of competingmycorrhizae is limited to ones' a trufflateur in Washington growing nonnative arelikely to do betterthan rn truffie (Tuber species,inoculated trees i...r,tt"u produced the Oregon white with ,"g1o.,,of North Americathat are resplendent hundredsof competingmycorrhizal varieties' for One methodof inoculatingmycorrhizae calls oi proven plantingyoung seedlings near the rootzones mycorrhizae ir,tffl. ti."r. The new seedlingsacquire of from a neighboringtree, and a secondgeneration is prodrrced' treescarrying the mycorrhizaifungus replanted Aftera fewyears, the new treesare dug up and longesthis- in newlocations. This methodhas had the soils'trees' toryof successin Europeansites where the and fr,rngiare comPatible' guar- Anotherapproach, simple and elegant but not into water anteed,is to dip the exposedroots ofseedlings candi- enricl-redwith the sPoremass of a mycorrhizai wild' and date.First, mushrooms are gathered from tl're the fruiting the spore-bearingsurfaces are removed from of bodies,crr-rshed, and immersedin water'Tirousands broih of sporesare wasl-red off, resultingin an enriched mushroour' inoculum.A spore-massslurry from a single inocrriaterr dilutedin a 5-galionbucket of water'can are hr-rndredor moreseedlings Mycorrhized seediings oncs healthierancl grow fasterthan nonmycorrhized gros' (seefigure 42). Eveni[you arenot successfulin this pzrir- ing tt.rffle mushrooms,the treesbenefit from ;i, FIGURE41 ingwiththe introduced mYcelium' water as a carrier-on the themycelium of Tossingspores using Truffle"brule" surrounds this filbert tree As is thesurroundlng abovethe rootzones of likelytree candidates Tubermetanosporum consolidates itsdomain' ground inthe calcareous takeslittle time and effort Habi- u.qttutiondres, creating a noticeable zone anotherrrethod that takenroot' ,oir, u t.tttuttsign that truffle mycelium has tatsshoulclbesejectedonthebasisoftheirparallelsitr the wild. For instance,chanterelles can be found in Chanterellesare one of the n'rostpopular col- oakforests in the Midwestar-rd in Douglasfir forestsin lectedmushrooms. In the PacificNorthwest, harvest- the Northwest.Casting a sporemass of chanterelies ing chanterellesis a controversial,multimiliion-dollar into a forestsimilar to oner'vhere chanterelles naturally business.Unfortunately, the gourmet mycorrhizal proliferateis obviouslythe besichoice. However, the mushroomspecies are not readilycultured. Chanterelles successrate is not higl-r:eve n tree rootsconfirmed to be demonstratean unusual interdependenceon soil n'rycorrhizedwith gourn'retmycelia will not necessarily yeasts,rnaking tissue culture difficuit. At least4 organ- yield harvestablemr-rshrooms. Fungi and their lrost isns mustbe culturedsimultaneously: the hosttree, treesmay havebeneficial associations for long periods the rnushroon,pseudomonas bacteria, and soil yeasts of time with no ediblefrriiting bodies appearir-rg. h-roc- (red soil yeast,Rhodolorula glutinis, is needed for rilationsof mycorrhizaeby one generationof rnycolo- stimulatingspore germination and healthymycelial gistsrnay t'tot see fruition until tl-renext generation. development).Not onlydo otherrnicroorganisms play essentialroles, but the tining of their introductionis also critical to successin the fungal theater.IVIany expertsbelieve thzrt decades will passbefore the plan- tationsgrowing mycorrhizal species like chanterelles matureto a productivestate. No one hasyet grownchanterelles to the fruiting bodystage under sterile laboratory conditions, although greenhouse-grownpines have produceclchanterelles afterinoculation. In 1997Eric Danell (accompaniecl by F'.Caniacho) was the first to successfullycultivate a chanterelle,fruiting rnushroomswith a potted l6-nonth-old pine seecllingin a greenhouse.Soon thereafter,Danell patenteclthis particularlyvigorous strain,which showedcommercial potential. Field tests in 2'l locationsrevealed chanterelle mycelium in the seedlings'root zones 2 yearsafter inoculatior-r. Unfortu- natel1',he coulclnot stopgrazing anirnals, such as deer, squirrels,and beetles,from foragingand disturbinghis crops.More recently,Danell started a Swedishcompany called CantharellusAB to comr.nercializethis break- througl-rrnycotechnology in the creationof char-rterelle orchards.His grouphas planted thousands of treeswith the chanterellemycelir-rn in an attemptto createmush- roon.rplantations that producenushroorrs within a decadeof planting.For the time being,only the patient mightr,vant to investin mycorrhizalplantations. Giventhe long time involveclin honir-rglaboratory techniques,I lavor the lou.tech approachand tradi- Comparrsorrot big leal maples(Acer macrophvltum)wilhout tionalmethod of plantingseedlings adjacent to known tsmancriarrr j with ilarger) mycorrhizae. producersof chanterelles,matsutake, trr-rffles, and MYCELIUMRUNNING

Sincecertain fungi function as natural bactericides replantingthe seedlingsseveral years boletusand then themas allies in an not for its andfungicides, some insects engage later.In this way,we can value the forest coLrnterinfections from hostile bacteriaand lumber but for its potentialto effort to quantityof harvestabie (Cunie otherfungi. Studies at OregonState University harbormushroom colontes' leaf- et al. 2001)show that attineants, which include cutters,grow kpiota myceliumas a hostfor a benign MutualisticSpecies: Fungal Partnerships bacteriumthat producesan antibioticagainst destruc- work Mutualism occurswhen 2 or more organisms usuallyto directiytogether for their mutual benefit' nutrients' preventinfestatior-r by parasitesand gather fungi aspro- Many organisms,from plantsto ants'seek field of tectors,,"nd ui.. versaThis rapidlyexpanding studyis one of the mostexciting in mycology'

lnsectsand Mushrooms for incubat- Many insectsuse mushrooms as platforms casesof irrg and feedingtheir larvae One of the first mitl,alism to be noticed was the interrelationship mushroom behveenmushrooms and termites The of mush- gentsTermitomyces includes several spe cies espe- ioor,t,,rro.irteci with terrestrialtermite coionies' with ciallyin the tropicsTermites construct their nests aban- organicmatter and cultivatemycelium' When do"n.d, these n'rycelium-richnests sprout delicious mushrooms.

i:. FIGURE44 the This glantmushroom, Tern itonyces sp, is hi ghly Ia-vored by peopieof centralAfrica as a deliciousedible The primordtum [.gin ,.urrulfeet underground, deep in a termitenest' extend- ingupward as a long"taproot," and then forming a mushroont How FIGURE43 on"thesurface, especially when the nests are abandoned myco- termitescultivate this mushroom has befuddled the best Ternitomycesrobustus, a delicious choice edible mushroom' logicalmlnds in the world sproutsfrom an aged. abandoned termlte c0lony tivemicrofungal parasites (Escovopsis sp.), and thel, also live in closerharnony with or-rrenvironment. Both feedLepiota mycelium to their larvae.This complex ants and people ber-refitfrom the guardianshipof partneringhas allowed lineages of attineants to survive n-rycelium-by partnerir-rgwith fungi, many organ- for morethan 50 million years,and to establishmassive isms,inclr-rding humans, can resist disease. coloniesnumbering in themany millions of inhabitants. (Seechapter 14 for a discussionol Lepiotaprocera and Snailsas Fungus Farmers Lepiotarachodes growing from anthills inoculatedwith Snailsand slugslove mushrooms-anunfortunate srt- spawn;also see chapter 8 regardingmycopesticides.) uationfor manyof us mushroomlovers. Some snails The mutualism betweenants, n-iushrooms,and enlistfungi to help them digestplants. Silliman and bacteriais a useful mode'lfor how we humans can Newell (2003)found that a seasidesnail, the marsh periwinkle (Littoraria irrorata), damagesand then defecateson certain grass(Spartina alternifolia), wherea particularfungus soon grows. Days later, the snailsreturn to the grass,now overgrowingwith fun- gus, and consumeboth fungus and plant. Grasses without the snail-enabledfungus grew 50 percent fasterbut were less appealingto feeding snails, whereasthe plantscovered with fungus were more palatableand nutritiousfor the snails.As you can see, the snailand fungusrelationship affects other species in the marshenvironment, such as grasses.

Endophytes:Mutualistic Symbionts Endophytesare primarily benevolent, nonmycorrhizal fungi that partnerwith many plants,lrom grassesto trees.Their mycelia thread betweencell walls but don't enterthem, enhancing a plant'sgrowth and abil- ity to absorbnutrients, while stavingoff parasites, infections,and predationfrom insects,other fungi, and herbivores.Cenerally, endophytes are not true saprophytesorparasites but arein a classoftheir own. In contrastto rnycorrhizalfungi, many endophytes grow well under laboratoryconditions, so we can make spawnby using methodslike those used for saprophyticmushrooms (Starnets 2000a). The vastmajority of endophytesare undescribed, and someappear to havelost the abilityto produce A FIGURE45 spores,living vegetativelyin a continuousmycelial TrevonStamets isexcited to havehis picture taken with the state. Most endophytesdescribed thus far are parasolmushroon (Lepiota procera)which iscultivated by ascomycetes.One exampleis Peziculaaurantiaca, a qfltst0 help thsmstave off infections. Many ants and termites smallcuplike mushroom that liveson healthyalder larn fungi trees.Like manyendophytes, ihis fungusis dimorphic, 4 MYCELIUMRUNNING

scientiststoday' are still far beyond the most sagacious can suddenly appear when trees mtishroom, "seedbails,"clay-soil peliets rich in A mycologistfriend of He invented chippedinto landscapemulch jump-startweak- cl-rips microbesand seeds,designed to a truckload of fresh' mostly alder *i,-t" nra Fukuokadid not selectthe Soontl-rereafter' ened habitats.Although deliveredto his housein the spring' of fr-rngiin his seedballs,he appreciated friendstook some home for populations his mostlyconservattve is to better were ih"i, co.,tribution. One of our tasks That fall mostof their mulchedbeds mulching. method and customizeit for potent Psilocy,bes'Where Fukuoka'spioneering fruiting i"ith h.t"d'"ds of straw' mycorestorationproiects Beginning with one have come from' n-rymycologist friend could they followedby mycor- is that the the saprophyteslead the charge, *ond.r"d? The only plausibieexplanation and endoPhYtes' in thewood' aboveground' while rhizae myceliawere already mushroom can As caretakersfor future generations' werealive' Perhaps Psilocybe rnycelium the trees treesgovern habitat progres- with mushrooms communitiessurrounding be endophyticbut delaysfruiting believefungi haveevolved to supporthabitats groundcontact' sion.I untii the treesmake hundredsof mycorrhizal' overthe long term,protecting generations When engagingfungai allies-the yearsinto tt," frrt.tr.. Saprophyticmushrooms gobble anJendophyticmushrooms-plants ben- saprophytic, andprevent invasion by rnycologicalsys- up debrisfallen from the trees ln l'*ryr. Thesecomplementary expand "fi, and parasites.The mycorrhizaechannei nutrients' temshelp plantssurvive stawation' dehydration' ,oot ,ot-t.r, and guard againstparasites' Similarly' The richer the fungus-plantpartner- parasitizaiion. understood'chemically can suPPort' endophyticfungi, less weil ships,the more organismsthe habitat reoelbacteria, tnsects' and otherfungi' After hundreds Fukuoka,the farmer-scientist'ecologicai Mrru.obr. fungalalliances have Revolution of millionsof yearsof evolution, visionary,and author of The One-Straw of naturekbody politic' It is time for our scientificreductionism failed becon-repart (1978),understood that wisdom' that are soeciesto partakein this ancientmycological to reflectbioiogical synergisms-processes

..1"FIGURE F

Psathyrellaaquailca nom. Prov' UntilrecentlY, conventional wis- domheld that gilled mushrooms didnot exist underwater. In2005, thisPsathYrella, a new species, wasdiscovered inthe clear, flow- ing,pristine waters of the Rogue Rivernear Crater Lake, 0regon' (SeeCoffan, Southworth, anc' Frank,2008, in Press ) This dis- coveryopens up a newbranch ofaquatic mYcologY, andraises manyquestions. How many othermushroom specles grov/ underwater?