n.: Rok of File In ~or1l."" C,r<UmpO!>, Eoo.'>,em,

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CHAPTER 10

The Ecological Role ofFire in Jack PineForests

1.H, CAYFORD AND D.l. McRAE

ABSTRACT

Jack pine (Pin'" ""nks;ono Lam~, 1i< I spea.' .... n adlpt.d to fir. in ,he I><".al r,,,..,em·imnment. Cone ..ro'iny IOJ In Iblllt\' to 'u","e haM 'n' lronmental conJi'io",permits jack pin. '0 "generale ,u<U"lully al,., fI,.. lnde.d. ""hou' ,h. periodiCO<'Curr'nce of fir< lhe~... "'ould di",f'P""' hom ,h. boreal for.". fire in rhe Jac.pine fo..... "a hlihl~ "a"ab~ pher>om<non: behuiou, e,,"'lKteristia ar. dependentlarge)~ on the fuel .ha".ete,,";'" 01 indi'iJual "ond, and 0" fi ... "-.>lhe,, Pre",,,l>edfire l> be,ng u..J <u«<>sfuJl)· in ,h. jad pine lore.. as a" <eo""""..al Slte-pre"".."o"tool lor regene,."ng Iog;ed-owr ;i,«, Ir <in ..J In slash removal. ..eJbedpre"",ati01l. haz.a,d T.Juction.•"" compelltlon ,.duet""'. Be<;a".. 01 ,he ,-a"romplexi')' of ,he ,-..riable< i","OI.,.J in the jack pone lorest fire <ws)'''em. continued,~arch in rhi, fielJ is ..·.".",ed.

10.1 I",TROI)Ucrl0~

Of lhe 105 pine speci.,. di",us;ed by !>l"o>' (1967). l<:\Cn mal' be consideredas nonh..n Clrcumpol.r .pecie>_ Four of lhel<: ""cur in I'o"h Am..iea: jackpine (Pinus ""nksi"n" Lamb.) is b01h an eastern and. "'eSlern NorthAmerie.n specIes ....hite pIne (Pi"' ... strohu, L.) and red pine (Pinus ""i"os"Ait,).re eastern ,pc"ic<. "h'le lodgepole pinc (Pim<s Co"">rIa Dougl.) is aWe'tern spc.ies, The olher three nonhcrn pines are found in Eurasi., ScotsplOe (Pi"us .y/t'tslT;S l.) is lhe most widely distributed of all pines. occurringfrom Scotl.nd in lhe WCSt '0 'he PaCific coaM of S'beria , Two other pines arcfound in Siberia. Pmus "blTlca Ma~r and Pinus p"mi/a Regel. the latter a'-cry small trcc and more often. cr... ping shrub

Forest fir. pl.~,s an imponant role in the ...olog) of pine forests, B~

exposing favourable seedlxds for germinal ion. it often enhances p.ne­regcner.tmn. Lodgepole and jack pine a,e pafticularl~' well adapted torc~cn<ra'ion after fire. The)' are char.clerized by l<:rotioous cones lhat donOl nonnall~ open until the~ are exposed 10 high temperatures_ Because of

""

Th~ RoI~ of Fi.. in NOr/hun Ci,mmpvla' Ecosystrms

lheir unique abilil~' 10 disperse seed lollo"-ing hre. lhe} are a widelydimibuled species. This chapter will deal only "ith Jack pine.

Jack pme is the mo>l WIdely dlmibuted Canadian pine. ransmg from No,aSeolia and Ne" Brun,wick "-csl,,ard, to norlhern Briti'h Columbia. lheYukon Terrilory. and lhe Nonhwcst Territorics. In the Uniled Slates il i'mosl abundant in Ihe ,lales of MichIgan. Minnesota. and Wisconsin. Thevalue of jack pine was nOI recoV"ud unlll after the exten,i'e Slands ofea,tern while pine and red pine were ha"CSled. Jndeed. R01h (1902) referredto it as 'that frugal scrub among the stalel~' race of nonhern e"ergre.n", Thetree i' no,,' an imponant commercial ,pecics used lor pulpwood. constructiontimber. h~'dro and lelephone pole,. fence posIS. mine timbers. railwa} lies.and ChriSlmas trccs.

The species i' capable of gro"-ing on lhe driest and poore't sites ,,-ilhin itsrange. and its abilit} to produce merchantable stands where other SpeCICS canscarcely sur..-i,'e is One of its oumandlng "alucs_ The charactcristlc Jack pme,iles are dry sand plain, d..-eloped on glacial out"'ash. morainic. aeolian. andbeach deposils_ Jack pine i' also found on fresh 10 moi'll sand,. on till,. and onrelali...ly thin soils o>erl}ing rock oUlCrops (Chrosciewicz. 1963: Ca}'ford.t9l1).

Exten,i,-e areas of jack pine are generally con'idered rather unallracli"e.but less "lensi,-. 'land' thaI lend divenil}' to Ihe landscape. or occur onrock-oulcrop ridges. h.,'e high "isual appeal (Ohmann rI 01.. 1978), Jack pineareas prO\-'id. OpportUnlllCS for hlu.berr~· pICk loS. and pm. is generall)conSIdered 10 be a deer food of medium preference. Groups 01 mung trecswilh branches reaching lhe ground pro,-ide nesting sil'-'" for the endangeredK"t1and\ warhler In M"higan's Lo"'.r Peninsula (Bcnzi., 1977),

To a considerable eXlent, jack pine <lands are consolidated into larg. areasof flat to rolling lopograph}' in situalion, "'here the}' can be utilized.fficiently. and operated and managed On a ,-.ar-rou~d basis. Commercialoperations are seldom 10 direct conflict with re,reatlonal osc of lhe foreslMultiple-u,e valoes are normally enhanced b} access pro"ided by roadcontructio~. and lhe har"-CSling and regeneralion of old-growth jack pineconmhutcs 10 a varied habitat for ,,·lldlrf. (Y.alman. 1979).

Jack pin. is senerall~ a short.li..d ,peci.s and is inloleranl of shad•. Ondry. sandy ,ileS jack pine tend' 10 be a climax type. "hOleas on Olher ,ite' it i,a temporary lype Ihat is replaced b~' more shade-tolerant species as long asnalural succession is nOI interrupled b~- fire (Fow.lI,. 1%5. Ca~'ford. 1911:Benzie, 1913).

10,2 FIRE AI\D JACK PI:\'E

It has long been recognized Ihal fire is a panicularly importanl ecolosicalfactor fCsponsibl. for re~e"ing jack pine foresls. Indeed. as ea,l}' as 1911.

Jod Pm~ Fo'~sl5 ,~

Ellis reported thaI jack pine in Omario sre" p,im.ril~' On bumed-o>'er areaS,He also noted that Ihe oxtens,,'e !Hands of jack p.oe "ere due to paSI fi'e" .0facr, Ihe periodic occurrenee of wildfire is conside'ed by all autho'ities to bean indi,pensable pha.. in the nalurallife c)'de of thc species (Hcin>elman.1973, Ahlgrcn. 19U: Da~' and Woods. 1977)_

Bo'eal jack pine forests are rspecially prone to fi,e, particularly on dry 'ilesand al high densilies ,,'here Ii>'ing unde,gro"'th is ,pam: (Rowe and Sconer.1973), The foliage of jack pine is hig./tl) combuslible and bums read.ly "henIhe tree crownS are suffiClenrly ck>sd) spaced 10 carr~ the fire (Van Wagner.1967). Jack pine nees of almost an~' age are usuall~' girdled and killed b)wildfi,e and relali"ely few SUf"i"e a crown fire (Mitchen. 1954: Frissell. 1973:D.), and Woods, 1977)_

In 'he soulhern pollion 01 irs range. on Ihe xeric glacial our""h "'nd plainsof cemral Wiscon,in and lower Michigan. man)' jack pine stand. h.>,c beensubjecled 10 surface fires of moderate inlen,;t) that kill only ponions ofstands. The.. periodic fires h"'e c.eated Ihe open. palch). multi-aged pinestand' Ihat cha.acrerize Ihe 'pine barrens (Hein..lman. 1981).

Rowe .nd ScOlter (1973) in discussing fire in Ihe boreal fNc.t h."esummarized some of Ihe aUI""ological chararteristics of the dominanlconifers. indicaling in a relati"e way Iheir degree of adaptalion for successfulregeneralion follo,,;ng fire. A lOt.1 'index of success' showed thaI jack pine isbest adapted 10 fire, whereas balsam fir (AI>;~ ba/jamto IL.1 MilL) is leastadapled. Jack pine ranked hig./t in relation to ..ed rrlenlion on the " .. ,earline.. of seed production. seedling growlh rate. seedling frost hardiness.and ,.edling grO"lh response 10 full light exposure,

10.2.1 Cone StroHn)'

Wilhout doubl the single most importanl characle",t.C of jack p.ne Ihatmakes the speci.. ideall)' suited 10 regeneration afler fire is Ihe ,.ro"!\Oushabit of irs cones. Scrotinous cones are wmmon O\'er mos, of the range of ,hespecies (Eyre and l..cBarron. 1~) and only in the southern porlion of therange in Ihe LIke SlateS do nOn·serOllnous cones occur 10 an) eXlent(Schoenike. 1976). II may be po>tulated Ihat the serotinous habil isperpelUattd by fires ,,-hich are most pre"alent in norlhem and "eSlern stands(Schoenike, 1976)_ A fire regtme in which fires are spaced to allo,,· fo' Iteemaluration. as occurs in Ihe boreal foresl. should f",'ou, serolin); 100frequent fires could eliminale Ihe speei... "hile less frequenl fires. as in IheLake Stales. would sele<:1 open-cone indi"iduals "hose seed Would cause IhegenOI)'pe 10 increa.. irs ,ep,e,.nla"on in Ihe Stands (Rudolph rI QI., 1959;Hendrickson. 1972).

Jack pine is an early and proficient ..ed produce' and "ees may' begin tobeat wnes at 3 to 5 years of age. wilh significant seed produclion attained by

'"age 10. Lighl 10 medium wne crops are produced annually. "ith good crop'al inleriab 013 or 4 )cm (E)"fe and leBarron. 19~: Roc. 1'KJ3). Seells inthe tightl¥ d""'d oerotinou, coneS accumulate on the tree, for 2S years Orlonger (Ellis. 1911): Ihe cone scales are held IOgelher wilh a bondIng materialthat mells al "oout we (Cameron. 1953). Seed in d~d ,ones remains"iahle for a long period of lime, High '-iability is maintained for al lea't 5~·eal'5. and seed' ZO )e .... of age rna) "'-crage as high as SO% germinallon.Even sce<b from the oldes' roneS pla~' 'he" 'ole in natural regeneration(Schanll·Han,.,n 19-11: Ey'TC and LeBarron. 19o1~).

16.2.2 Co..., Op<>nlng aMI Sttd O;ss..mination

For"" fir... pro,'ide the heat necess.a~ to opt" the large number of cones thaIarc present in jac~ pine ew"'o•. Although a fire may generale ,try h,ghlemperalure•. lhe seed. "'ilhin lhe cone, are well in,ulaled and Can Su ....·'Hbrief period, in flames (Eyre. 1938, Eyre and LeBarron. 1944), In detailedSludie•. Beaufait (l96Oh) found thaI seed ,',ab,loty was unaffecl.d wh.n con.sw.re .xposed to 900'C for 30", al 7lXl'C .'jabilil; did 001 deerea", unlil coneswere exposed for a 3·m,n lOlerva!. E,'en lllough mOSllree. are killed b) fire.cone. arc opened and pro"ide an abundanl "'ed ,uppl;'. Follo,,·,ng a firc. up'0 5 milloon !.Ceds per hcetare may' be disseminated from lhe ne"'I; openedcone, in lhe scorch.d cro"'n, of slanding uees (E)'re. 1938).

10.2.3 Gcrminallon and Seedling 1.Ic,·clo-pmcnt

Requiremenls for ",ed germinalion and .arl; ",edling grO"lh are oflenideall)' crealed b; fire. Fires remov'e loose organk maller such as surface Iolleror moss. and upose mineral soil or deeomposed organic maller. "hich aregenerally' f",'ourable seedbed. for germinalion (Ahlgren. 1959. 1960;Chros<:,ewio~. 1959. 1974, 197&1; Ahlgren and Ahlgren. 1960: Cal'ford.1963). Complel' .xposure of minenll soil is ",Idom required. and a mixlure ofe"posed mineral soil and 'hin humu. rna;' provide Qpumum CQndiuQns(Chrosriewio:. 1970. 19(4). On moiSl .ites. suilable "'edbeds may resuh fromhuming onll' lh. loose surhce Iill.. and lh. fealhermoss. l.a'ing thccompacled. lower organ'e Slrala imaCl (IZ. Chrosri.wic:z. 1979. perwnalcommunicalion)

Fire. nQrmally kill mosl of lhe O'·....10re;· Ir.... bUl dead slanding Ireespermil much Oflh. solar radial;,;,n 10 r.ach lhe ground. Moreo"er. the panialshade erealed b; dead 'land,ng lrees as "ell as by logs Qn lh. ground surfacecreale, suilabl. conditions for jack pine ",cd germination (Fraser and Farrar.1953) which normally occu... promplly alter "'ed dispersal if lemperalur. anJmoiSlure rondiuons arc favourabl., If such condilions continue during lhe

'"'first gro"-;ng season. a sland of iack piM is hkel~' to be regeneraledsuCttssfully (Eyre and UBarron. 1'41),

Fire temporarily eliminates competing 'egelation. and the imensity of thefire affecl> regrowth. While certain herbs and shrubs may provide a shady.cool microcn,-ironmem "hlch is beneficial for germination and early gro"..th.a dense co\'e'. especial1}' of haul (em:!"'.... rom",,, Marsh.). may retardgro,qh of jack pine seedlings, The additional nunient' released b) the fir.mal aid In lhe reestablishment of jack pine. sustain its growth. and stimulatethe growth of herbs which will pro"ide lhe beneficial cool. moisl microclimate(Ahlgren. 1960. 1963. 1970: Ikan!ait. 196O:a).

Ahlgren (1959). Cal'ford (1%3). and Chrosc:iewic:z (1910) have obse ...·edthat m0'51 germination OCC'ITl; promptly afler fire. Ahlgren reported thaI. inMinn.sota. 57% of lh....dlings originated lhe fim y.ar and 35% the serond)'ear, Cayfotd found in one slud) lhat 99% of more than 1000 seedlingsgerminaled lhe firs! year after ,,'ildfire.

IO.2.~ Post-fi~ Sueemional ranenl5

A number of in""ligato~ h.,'e carried oul romprehensi"e fire erolog)studies ,n areas in ....hich jack pine is an importanl componem of lhe forest.The eafliest comprehensi"e work was undertaken b)' Ahlgren in north­easlern Minnesota (Ahlgren. 1959. 1\l6O; Ahlgren and Ahlgren, l\l6O). Morereceml~·. Day and Woods (1m) have carried Oul a detailed Slud~ innOrthweslern Omario. while Heinselman (1973. 1981) has done simila' workin Minnesota. These in,·.,t,ga,ions ha"e all been ca'ried oul in lhe GrealLakes-Sl. La....r.nce Foresl Region (Rowe. 1972), In lhe Boreal Foresl.Ro....e and Srotte' (1973) ha"e in"esligaled fire ",olog)" in northern Canada.while Alexander (1978) is cu'renlly carrying out sludies in Onlario. In N.,,·Bruns,,·,ck. Maclean and Wein (1976) ha"e in"esligated posl-fire jack pineslands in the Acadian ForeSt Region.Da~ and Wood' (1977) h..'e presenled a detail.d descnption of five

post-fire jack pine Slands ranging in age hom IJ to 160 years. The ,esull'from lhis sludy confirm lhe facl 'ha' 'he Oueliro Pro,;ncial Park forest innorthw.,tern Ontario is romposed of a mosaic of ea,ly and late successionalslands originaling from la'ge "ildfires. The resuhs also indicat. thaI. in lheabsence of ,,;Idfire, jack pine "'ould be replac.d by stands of moreshade'lOlerant species. In lhe sam. pape'. Da" and Wood' presem lhefoUO"'ing h)'P"lhelieal model for post-bu,n succession in the jack p;ne-aspen(Populw Iffmuloide. Mich.) rommunity.

In lb. fi~l phase (15 years). a dense. ,'igorous. e'en-aged jack pine-aspensland dominated lbe burned·over area. Jack pine al 10000 stems per heclarewere lwenty limes as abundanl as aspen; beneath the overslo",)' ....as a blackspruce (Piao ""'tiona [Mill.l B.S.P.l underslor.,., In lh. serond phase (45

'"')'carsJ, the fi,c';m'ialcd Jack pme and aspen s"n formed an e'-cn-aged caoopy'hat dominate>:! a spruce and ~hade-1Qlmmt hardwood [sugar maple jAc",accharum Ma"h.).iwhite birch (Brrula papJ'ifera Marsh.)) underslorey.Neither pine nor 3$p<'n had regener.:l1ed because of 10'" light 1.,ch. severeroot rompclllion. and an increasing depth of pine n«dle Iilter. In Ihe th"dphase (75 years). the jack pine and aspcn Slill formed an e'en-aged canopy.but their crowns had declmed In size and the canopy was begmmng to 'howsigns of decadente. The un<krsll:>rey of black spruce. balsam fir. andshade.lO~rant hardwood' was <!e"elop,ng and ,"'as beginn,ng to become themain 'land canopy In the fOllnh ph.", (105 years). the jack pine andseRucTed aspen were decadent and dymg. Black spruce. fir. and shade·tokrant hardwoo'h ....·N~ ,n lh~ prClCfiS of .~plarins lh~ O"~,ma'ur~ Jackpin~-aspen 'land. In lh~ fiflh and sixlh pha..,s (lJ~ and 165 ~~ars). Iheonginal jack pme and aspen had be~n almosl complelely r~placed b~' Ih~

undN,torey. Al 13~ }'~a!S. only scaltered jack p;n~ r~m.ined. "hile at 16~

yea". only a f~" larg~ anriem and decad~nt sperimens r~main~d. Theoriginal jack pine-npen forest had been ,eplaced by an all-aged spruce­balsam fi'_lolerant hardw<><>d sland.

He;n",lman (1973) has reponed that in Ihe Boundary Wale" Canoe Areaof Minn""'la. jack pine-dominated eommunni.. "'here blaek spruce issignificant ma~' e,enluall)' be su«eeded b) a black spruce-feather moss(Pleurozrum Jehreberl) eommun;ly. Whe'e balsam fit tS sign,ficant In

as5OCialion "ilh jack pine. lhe ,u"""",ion "ill be lo....·.rd' a baJ"m fir-"hilebi,eh-"hne spruce '~pe "'ilh a lall shrub lale!. Hov.ever. Heinsclman found'hal the'" sucr=ions ....·ere n,,'cr entlfeh compleled. and in 1<)70 he found afew scallered Ii'usiack pine in a sland that had burned in 1727. In Ihe "mel'ear he found numerous 21>-lear-<>ld jack pine In a sland burned in 175~.

Thus ,t I. "'idenl lhal indi"du.1 Jack pine can pers'st as a scoueredo"er-slorey for alleaSI 210 10 250 ~'~ars "'ithoUI fir~.

10,3 FIRE En-ECfS

Forest fir.. hO"e 0 profound effeCl on lhe jack pine ecosyslems. The lreecomponent and all orhe, ph),;e"1 and biological p,ocesse, are direetl~

affeCled_ Changes caused 10 an)' one of these processcs "'ill ullimalel) affcclmost olher procrsse<.

Scoller (J'J6.I) noled lhat soil temperalures in burned..,,·.. are.s "'erenormally h,gher lhan ,n ..mila. unburned areas. Th,s lempcralu,e differencedecrea"'d as soil depth increased. The ,ncr..", ,n lemperatu,e ,n Ihcoomed..,,,r area waS auribuled to the blackened surface. "hieh readil)absorbed solar eners). 10 lhe remo,'al of unineOfporated organic maller(duff) "hieh aels as an insulator. and to the absence of shade from lhe foresl.anop). Sconer belie"d Ih.1 m.ny of lhe fire-induced 'PCcies tn lhe earl~

'"... ra! ,t'g~s of succession ....ould Mnefit from the incre.",d 5Oiltcml"",aturc".-hion is eoologic.II)' f,"oura!>le for tl1c germination and growth of someplams

In northeastern Minnesota. the hne. romponent of the ""il "'.... found toincrea.. rapidly within one or two momhs after fire through needle fall fromkilled lree, (Grig.1 and McColl. 1~75). Needle rail decrea>ed .fler ,h" reno<!but oonllnued for up \0 \"" years after the burn Needles in the burned.{werareas rell prematurely w';lh higher nitrogen. phoo;phorus. and potassium andlo"er calcium concemrations than in the unburned areas.

In the first year aher fire. the \>oood)' componem of the Il11e, la)'cr wasinCTealoed by the hreakdown of SIll.1I branches and t\.,gs of the Iree< killed bythe fire (G.;gol and McColl. 19751. During the ..rood \ear. the component"'as augmented b) the breakdo"" of tree er",..ns and the fellIng "f completetrces. Sloughing of large quantitIes of lree bark "hich accumulated al thebase of Ihe dead !tus ""as e'iden!. By the third year. "'nody liuer fallreturned to unburned le'·els. Litler buildup by cones and seeds "'M reducedb) fire because of loss of reproductiw organs, Liner Increases caused by thc"nod) components of trees. frullS. and seeds after fire did not aff", numentconcentration' of thc btter component of the soil.

Fire also has an effecl on soil nUlrients. Ahlgren (1960. 1963. 1970)obse"e<J higher concentration of nl1rates. am,TWn,a. pota,h. phosphateS.calcium. and magnesium in upper soil levels than on comparable unburnedareaS. The higher concentrations of nitrates and ammonia diminished rapidl,afler fire; those of potash. pllosphates. calcium. and magnesium "'ereundoubted I)' caused by Ihc residual a,h. and they prevailed 10 SOme exlent 'nthe soil for at leasl fi\"( years aflcr the fire, Howe"er. the actual soillype is ake' factor in delermlning thc dfect of firc on nutrlcnt cycliog. M<:Coll andGrigal (1977) o!l<;e"'e<J Ihal ol1rogen fixatIon. I1rter decomposition. andmineralization ...·cre enhanced by cenain t)'pe< of fire, The) felt that Iheincreased soil pH was Ihe reaSOn, In Saskatchc"'an. ScOller (1%4) found thaIcxchangeable hydrogen decreased and .xchangeable cakium increasedfollo",ng fife. On Ihese same burns he fouod that "'il pH In.rcased hecauseof the fire but thaI increases werc Ics, at grealer "'il depths, He consldcrcdthaI increased pH WaS due to Ihe d«lruction of unincorporotcd organICmatenal and the addition of "'nod ash 10 the soil, Ihe addition ...·a' naluralt,grea,ell at the soil surface.

Vegetation response 10 fire is a complcx matter in,olYing man) factors(Ahlgren. 1960) including 'preburn condition of the land. season of bum.seed suppl)'. (fire) inlensity. a,h concenlralions. 'u!l<;equent miner"l­nUlrilion. soil moISture. rainfall. humidity, "'il and all temperalures. animal

populations and plant competition'.In Mtnnc""ta. Ahlgren (1960) obserwd three calegories of plants in

fire-kIlled stands: those Illat "'ere found on unburned land. those thaI "cre

found onll on burn~d-<»'~rland, and tJ>o.c that occurr~d on both burned andunburned land Fell species occurred onl:' on unburned land. MOSI ,peci~s

found onl}' on burn~d-(),~r land rerTOduc~d b~ seed and th~ maJO"t>' w~rewInd or animal dl~semlnat~d. The third cat~go..,'. occurring on both burn~d

and unburned ar~a~. contain~d a numbf'r of "egetatively reproduced specie~

"hich su!'\'ived the fire, Vegetati"e pans of se"eral speci", were remarkablyfire toleram and "ere equally distributed on unburn~d. lightll' burned. and.....' erell burned ar~as. In general. th~ occurr~nce of "egetati, ell reproduced~peClCs depended primarill' on the resistance of the reproductiw pam to fireor their escape from it

In the boreal forest Ro"e and Seot1er (1973) ha,e noted that a numbf'rof,ubordinate plant 'pecies commonll associated "ith jack pine are able toregenerate prompt I}' from basal or underground sproulS. Still other ,peciesfrequently in"ade burned areas. These are fa't-gro"'ing ,pecic,. adept atrapid ,",·.,;on b>' seed or ~' "egetame means. "hich dC\'elop In full light.Slmilarl~'. various bryophl t~s and lichen, appear soon after an area is burned,In nonhern Albena. Carroll (1978) has studied the successional trendsfollo"'ing fire in jack pine-lichen woodlands. noting in parucular lhe changesin occurrence of lichen ,peCles o,er time.

Caribou shO\. a prefer~nce for stands O\'er 50 years of age "hich rontainthei' preferred food source fSro1ter. l~). Fires "hieh alter the jack pineforcst from matur~ to Immatur~ reduce th~ food ",urc~ for caribou, Seo1t~r

found that a mature forest can produce 61~ kg-ha of high·"alue lichen loragefor the caribou "int~r 'Uppll' ,,'hile an Immature forest 1-10 lears of agesupported only 2 kg ha. In terms of total forage available 10 c."bou. recentburns prmid~d onll a third of the forage pro'ided by mature forest. Peek.1ai, (1976). in discussing the need to reintroduce fire into the Bounda.., WatersCanoe Area of northern Mlnne"'ta for ,uccessful mlX>SC managem~nt. foundthat vegetalion locared on 'holler" burns was hIgher '" pmtem' th.n th.t o~

'cookr" hurn~. I"ut"e~t qua lit) of plants follov. 109 fir~ appcar~d to depe~d ona'ailable "'il moistur~ as ,,'ell as o~ the 'intensn{ ofth~ burn,

IO.~ WIlOHRECHARM:rERIS1"lCS

Thc role of fir~ as a faCtO' in thc ecology of [he boreal forcs. has been welld'>cum~ntcd b) numerous In"cstlgators (Kalil. 1968: Komarek. 1968: Kiiland Chroscie.. iel. 1970; Ro..'c and Srott~r. 1973: Meth"en rI al.. 1975:Burge" and M~th..n. 1977: Woods and Dal. 1977), The iner~ased buildupof forcst fucl~ .ft~r ~arl) log,g;ng .cm'nics resulted In ""Idfir~, rhat Increasedthe ~'tenr of jack p,n~ in tbe logging sectors of tb~ boreal forest (Benli~_

1977; Burgess and M~th'~n. 1977), Thc major sour"" of ignition lor ,,'iIMir..in the boreal lorest before th~ arr;"al of man .... lightning and ~v~n afterman', .rri"al i[ contributed signifie.nlll to WIldfire ignitions (Koum. 1967.

'"Komarek. 1968), Komarek (l%S) '-;ewed the rok pla)'cd b) lightnIng in lheoorcal forell ecolog)' as ,'cry significant. and refe ... to the boreal forest as alightning fire bio-dim.r;c region. Boreal foreSl species such as jack pine.lodgepole pine. black sprucc, trembling aspen. and whne birch arc all welladapted 10 reproducing afte' fire. Heinsdman (1971) found thalll6-90% oflhe virgin fo'c<l in lhe Boundar)' Waters Canoe Area in northern MinnesOlahas resulted from fire.

Before mao', inten.mion. internls as shon as 5 years and as long a, 100years between fires ",ere documented for lhe borul forest (lieinsciman.1%9. 1971- 1973; Shafi and Yarranton. 1973: Swain. 1973; Burge" andMoth.n. 1977; Cwynar. 1977: Woods and D.},. 1977). Heinselman (1971)found an imen-'al of 300 yea.. m some instances. H~in~lman (1973)eSl,mal~d that th~ fir~ qcl~. i,~ .. lh~ number olyears r~quired 10 burn O'eran area equalro lhe ,,'hole area of lhe forelt (Van Wagner. 1978). ,n lh~

Bounda!) Waters Canoe Area was belw~~n 70 and 100 y~ars. Van WagnN(1978). analysing H~in>elman's (1973) dala. th~orized lhat 50 j'~ars waS amore r~alistic figure Woods and Day (1977) showed Ihal. in weSlernOmario. fires now occur on the a'erage e"ery 870 years becaule of impro,'.dfiTe suppression lechniques

As d~lermin~d with lhe aid of lhe Canadian ForeS! Fire Weal her Index. lhemajor proponion of lh~ jack pine range hes in a 10", to moderate fiTe wealherzone while the remainder is c1as>ed as a high fire weal her ZOne (Simard.1973). Fire ,n most of lh,s jack pin~ r~gion will occur al Ih~ rale of 6-3.9fire"'l000 km' year Wilh small~r portion. of the region ha"ing up ro 7.7fir""'IOOO km',-'year (Simard. 19;5), In are.s ne.r urban centreS. fires mayoccur al a rate of 27/1000 km")ear

Increa"'d imere" in quantilati"e dara has r~sulted in lhe in·depthdocumemalion of wildfires in lhe jack pine 10Tesl (Van Wagner. 1965,Walker and Slocks. 1971. Slocks and Walker. 1973: Slock•. 1975). Descnp­tion of lhe aerial fuel components of the jack pine forest (Bro"'n. 1965:Oumtilho <I g/.. 1977: Walker and Slocks. 1975) has enabled in"esligalors 10eSlimal~ cr""n fu~1 consumplion during wildfires. The ~SI,malion of emwnfuel consumplion mak~s il possible 10 pr~diet th~ fronlal fir~ intensity(k W m). which is synonymous with Byram's fire intensity (B) ram. 1959). andInial h~at reka>e (kJ/m').

Fire research~rs alth~ Greal Lakes Forest Research C~mre in Omario arein,'e"igating fiTe beha"iour in a malure jack pin~ stand ",'ilh a black sprueeunderstorey originaling from a fiT~ in 1899 and in an immalure jack pine standongmaltng from. fire in 1'f.l8. Since lh~ in""plion of Ihis Slud). 13 and 17burns. r~sreCli"e1y. ha". be.n oondueted under "arious fir~ w~alh~r «lOdt·lions. [k..riplions of lhe IWO fuel types are oomained in Walk~r and Slocks(1975). fire: rales of spread in Ihe malur. ,tand ha,. ranged from 0.5 to 15m'min ,,'ilh fronlal file intensili~s ofI~ kW m. Th~ immalure stand

192 Th~ Rof~ of FiN' on Sorrhrm Circ'''''I'o{ar Ecosplems

ha~ had rale of spread measurements a'·eraglng II-<>fl mlmin .. ilh a,,,,,,ialedfrontal fir. intemiti., a'eraging 672-59596 kW{m. Les< intense lire.ocrurred in the malure 'land because it "'as difficult for lhe fire 10 cro"n,,'ithout lhe aid of an) ladder fuels. Onl)' a1 a "'nd .peed of 32 km h ..... thefire able 10 IOrch in lhis mature sland (8. J. Siocks. 1979. personalcommunlcalion)

Ouinlilho rI al, (1977) conducted .."en experimental burn, In open.malure jack pine .Iand. in non hem Alberta in whIch fire ,pread ralcSa,·craged 0.6-6.1 m min. They ol)sel\ed thaI fuel mO;'lure .. a, theconlrolling factor in lhe fire·, rate of ,pread ..'hen Ihe wind wa, con,tanl.Olher faclOrs which were found 10 affect the rale of ,pread were wind ,peedand dircction. ground ,'egClalion. and quan1l1y and conllnu")' of ~urface fucLIn 'hese bums. as a reflection of fire .. ealher condilions. fuel consumptIon(kg,m l) w"' J3..·57% for woody fuel. and 36--76% for the duff, The duffdeplh wa. reduced by ~2'l'o.

The sludy in Ontario suppons remarks b) Van WagJ1er (1977) who foundthai )'()Ung to sem,-malure Slands were more .uSttplible 10 crowning lhanwcre older stands "hose crowns are highcr and Ihinner, Such malure Slandsgenerally' support onl)' ,urface fires. Normal daily con'·ection acti";li..produce "regular "'Inds Wilh gu,ts inlerspersed "'llh lulls. Van Wagner (1977)obsel\'ed lhat ,·.,ial;on, in "'nd .pecd in combination wilh 'land structureand topograph)' can produce intermitlenl ac,i,'. crown"'g, Such interminentero.. ning Can be readil) ol)sel\ed from lhe air where;1 p.-odu= a nbooneffecl of burned and unburned cro"'ns in the landscape.

Fuel moi'lure. an important faclOr in fire boh",'iour. i, strongl)' affecled byalmosph.ric rond"ions such as lCmpcrature, rdalive humidit)". "·lnd speed.and precipilation. Van Wagner (1%7) found foliar moiSlure conlenl ofeaslern Canadian conifers including jack pine 10 be at their loweS! in lhe,pring. Slocks (1970) obsel\'ed lhal ",'aporalion from the 'urface of lhe fore'lfloor is important in lh. duff-<lr)"lng pallern of ,·arious duff, including jackp,ne

Van WagJ1er (1973) has predicted fire spread rates for standing jack pineand for jack pine ,Ia,h ming lhe fuel moisture codes and fir. behaviourindices of the Canadian Forest Fir. Weather Index, On Ihe basi' of theirexperimentai burns. Ouintilho rl uf. (1977) predicled lhe rales of ,pread for~im,lar jack pine ~lands ,n norlhern Albella. USing a simplified "enion of VanWagner·s model.

10.5 PRESCRIBED FIRE

A fore't manager may employ eilher wildfire or prcscrilxd fire as amanagemenl 100110 meel a specific objecti"e, Wildfire' have seldom beenused in tbe pasl because of lhe inherenl d,ffocuhy of comro\ling lhem and

Ja<:k /,,,,. Fort'" '"o.cau~ of the common altilUdes IOwards 1(}(al fire suppression by' Ihep"rsonnel r"ponsible for rhe,r eonrroL While prescrio.d fire has!>cen used inNonh America. 'I has nor found "-idespread applieallon

Jack plne regeneration ,il\'iculture o.g.n in the 1930s ,,'hen il o.camee, ,denl that the ,pecies would nol regenerate adequately after han'eS! ,uningunle...pecial silyicuhural practice, were undert.ken Rea",n, for failure ofjack pine to regenerdtc on [he cutover site' hay'e included non.reeeptiY·e"'edbeds. insufficient moiSiure. inadequate seed suppl). lnsufficient ligh!.and ..cessi",e planl comp"tilion (Eyre. 193!l: E)re and ~Barron, 19~4:

Rudolf. 19~6: Farrar n al" 195~: Beau!ait. I~59. 196Oa: Chrosciewicr. 1959.1%7. 1974. 1978<1: Adam,. 1966: Ca)ford. 1966b: Cayford elal.. 1967, Fosterel al.. 1967). Researen into Ine use of prescribed burning as a managementtool In jack p,ne regeneration ,ilviculture began In Ontario and Michigan In

Ihe 1940s and 1950s (Beaufan. 1959: Chrosciewicl. 1959) and inilial result'indicated that the lechnique had polential, Further researrh wa< conducted inIhe l%Os and early 1970, in 1I-llnnesota. Omario. Manitoba. andSa'karche"an (Adam,. 1966: Ca;ford 1966a: Ca)'ford el al .. 1967:Chr=iewicz. 1967. 1970. 1974 19780. b, c. d: Fo'ter.' al .. 1967: Ahlgren.1%3. 1970). The resulls are still being evaluated ,n lerms of jack pineregeneration (Chr=iewicz. 1978<1). bUI practicall' no new ",pcnmenlalwork has o.on underlaken in recenl years.

At [he same lime operational prescribed burning has found relativelv linlesupport largely because of tlie anilability of ahernah"e mechani,ed methodsof 'ite preparation (Cayford. 1971). Howe'·er. by the late 1970s. an ineTeas,ngbacklog of non-regenerated jack pine land. inereas,ng costs of meehanizedsite preparation, and an increasing awareness of fire management h",'e placedpreseribed burning in a more fa"ourable light as a 'il"icultural practice(McRae, 1979b).

The major use of pres<:ribood fire,n jack pine ,Ia'h toda}' is for sih'iculturalpurp"'es. namely ';te preparation for plaOling and seeding. Other minor usesof pres<:rio.d fire are for fire hazard redue\lon and wildlife managemeOl.

D;rect seeding has booen shown 10 be succe"ful on s;te, that h",'eundergone pres<:ribed burning (Ahlgren. 1~70: Chf=icwie~. 1970. 1978<1:John"'n. 1974). Following a successful seneS of burn,ng and seedingtreatmeOl'. Chroseiewie~ (1970) compared results from scarified and un·seanfied seedbeds and reported 'hat pre-buro scarification was unnecessaryfor burning and seeding treatmeOl' !O 0. effective. He also stroSM:d thatdetermination of the amOuOl of seed required for broadcast seeding dep"ndson man) factors indudlng "'ed ,-;ability, quality of fire-produced seedbeds.mineral soil mmerials presem. and amicipatcd losses 10 bllds and rodents.

Chrosctcwicl (1970) oOOc,,'ed that. 10 secure the d"ired number ofseedlings per he~tare at more or le~~ uniform spacing. the inten~ilyof ~eed[ngmust be in"e,sely adjusted in relation !O the qual it)' of poSt·hum ""edbed

,... Th. Rol. of Fort '" Sank,,,, Cin:umpolo, £("05""<""

conditions as defined by the 3wrage depth of residual duff and the e'po,ureand texture of minual soil malerial•. In hi, oplmon. pro>'ided that seed ismore than 7()% viable and IS tr.aled lOUt> bird. and rodenHepdlcnlchemicals_ 'h" Ill" of adj,mmrnt would probably mnge from 70 to 7oo! ofseed per hectare, Ahlgren (l'I7U) found thaI 385 g of ,oiahlc \.Cod per beetarega'-. a well-dlSlrIhuled >land of trees if~dingwas~ in the aUlumn ,,-henmouse act,vit) wa, low.

Sp"ng plantation' on areas that ha'-. undergone prescribed bu,mng wcreobst-"ed to hO\c a higher su"ival rale and a grUter height gro,,~h thanaulumo pla,lla1lon<. II "a. also found that plantations s.,,,i'-cd §C'"C"drought bener than diroet ",.ded ue", iCa~·ford. 1%6a: Walker and Dobbs.19611). Chr=iewia (197lld) cons,dered lha, lhe duff dep'h need be redueedonll modualdy dunng a pre""ribed burn for plan'ing. Planlers can easily gel,he seedling rOQ1' into direcl conlaCi w;lh mineral "'il e'en through amoderate duff ~plh. This obse"'alion is lrue onll' on 'ites "'here hardwood,pecie,. including deciduous shrubs. afe absent and will !lO1 re,proul lOrompele wlfh plan'ed seedJingi.

Prescribed burning condu<1ed under cerrecl condnion. ean reduce oompe·"lion for jack p,ne seedlings (Adam,_ 1966: Dlelerich. 1964: Fo<1er tI ul..1967; Chr=iewic:z, 1970, Anon .. 1975, Bro"-n. 1977). Prescribed burningcan al'" contribule 10 Ihe control of damping·off (Beauf.n. 1959).Sekrodern, canker (G.mfmcmcIlQ ubicllnQ Lagerb.) (Brown. 1977). andd"'arf mistleloe (A'UUlhoblum pwillum Peck) (Alennder .nd Ha"'h""onh.19751.

The behaviour of prescribed lires muSl be underslood by both lire and.ih,cultural 'peci.li,1S if lhe obje",\'e' of burning are 10 be aHa,ned. Eachpr....ribed fire i' unique and its beh,,'iour depends on man) faclors includingfuel condilions. "ealher. and ,opograph). The need for more quamnali'efire beha"iour dala i' no'" recognized (McRae 19'Na.b) and a handbook hasbecn produced 10 enable the adequale documenlalion of fuel and lirebeha"uur eharaClensllC> on prescribed burn, \n norrhern Omano sla.hcomplexes (McRae., al.. 1979).

RelulIom;h,ps bel"een (ire beh.\,our in Jaek pin. slash and lire wea,herconditions m On,ano as expre>sed b}' lhe Canadian foresl fire \\ ealherIn~~ Tables (Anon.. 1978) ""ere dcseloped by' Slocks and Walker (1972).Their ""ork on producing a Fire Beha\'iour ln~~ lor lhe jack pine log~png

sla,h fud 1)1JC has a'ded ,he planner of pr.cribed burns in choosing codes or'ndices of lhe Canadian Foresl Fire Wealher lnde~ 10 meel burn obJeet",.s.Fronlal lire inlen,it)". rale of ,pread, and fud consumplion paramerer-s (lolalfuel eu'liumplion. §la§h COnSumplion. dePlh of burn) "-ere found to bewongly related 10 lhe Fi'e Weal her Index. In",al Spread In~x. and Builduplnde~. re,pc<1i'·el}'. Cllr=iewicz (1978a.b) found lbal lhe Duff MoislureCode was .uflie,.n, for pred'ClIng duff·depth and duff..:o..r reduClion, by

Jack Pi'" Forrs!s '"burning when used wIlh Iht pre·bum duff depth and co"er in Manitoba andSasbtchewan.

10.6 CO:-:ClUSIO>;S

Tht unique ecological role played by wildfire in the jack pine fore't mu,t becon'idered carefully when fire ecology and ,ilviculturt in thi' fore" eco­'y'tem art being studied. 11 mu,t al'" be recognized that one cannotgeneralize from the ecological change< brought aoout by fire in one situationand expect the same result in anothN 'ituation. Each fire may create adifferent fore't ,ucee"ion. the ..i'tenee of which depend, largel) oncondition' prC\'aihng before lhe fire and lhe eharaC!eri\ti~of that I"'rticularfire, Because of the variabilily whICh may be encountered. funh.r re",arch i'needed 10 fill gaps in our knoocledge. particularl)' alx>ul the specific fi,eeffects o"el the emire range of jack pine ecmys,.m,. One must not neglectGrigal and McColl's (1975) ,,'arning that: 'E"ery fire i' an individual ."em.and umil a number of the J'O',ibl. t)'pe'S of fires are studIed. extrapolationmuSt be done cautIously.'

10.1 REFERENCES

Ad,m<. J.L (1966) Pr=:ribed burning t«hniqu« for site prepar.tion in cut-o"e, Jackpino in "",uthoa>!em ManilOba, Palp Pap. Mag, Calt" 67. ~7-l----~,

Ahlgren. CE, (1959) Some effects of fire on fo,..t reproduction In notthe.sternMinne""... J. Fo,., 57. 19-1--2ill.

Ahlgren. CE. (1%0) Some df«" of fire On r.produ"ion ond gro""th of "egetotion Innorlh.a"crn MinneSOla, E<olog;-. Sl. 431---4S5,

Ahl;ren. CE, (1963) Some basic eeo10;ieal 10<10.. ,n pf.""ril>ed bU,"I"11 In""rlh••stern Minnesota, Pro<'. Toll TimlHr.• fire Erol. Conf-. 2. IH_lSO.

Ahlgron. CE (1970) Some effem 01 pre..ribcd burning on jack pine reprodnction innortho"'t<m M;nnc>ota. Uni,'. M'nn. Agric. Exp, Sm,. Mr.sc, R,p. 9J, Fo,. Mr,.l, 14 pr,

Ahl~ren. CE, (l97s) The <ff.m of fir.. "" tempe.... forem; north «"1Ial U.S .. In

T.T. KOllow,kL and C E. Ahll1r<n (.d,), fire and Eco,>,,,,",, Aoademlc Pre",.N.w York, pr. 195-223

Ahlgren. IT. and Ahlgr.n. C E. (19(,0) Ecolollicol eff.c" 01 fore't fire,. BOI. R,,'26 . .>8:>-533

AIe,and". ~t ,E, (1978) Reoon"ru<1ing the firc hi>!or} 01 Pnh.kwa :o.:ational Palk. inDE, Dub<' (ed,). Fi" Ecology'" R"OUfC' ,~""ag,m,nr Wo,hhopI'.oatdings, Can, For. se,,·,. [nt, Rep, NOR·X·21O. pr. 4-11.

Alex.nder. ME. and H.""k'~Orlh. FG, (1975) Wildland fire< and dwarf mi"let"":a hte,ature ,e,'iew of ecoloR" and pr=:ribcd bum;ng, US Fo•. Sm'., Gm. T"hRtp. RM-14, 12 pr.

Anon. (1975) Pr~(rilHd 8a'",ng G"id~lint" US For se[\-'. NOrlh Rog.. M,,>oni.Montana, ';6 pp.

Anon. (1978) Canadian fore>! fire wo..hor index tab I... Con. fo, . .\<0"',, for. T~(h

R~p 15 -IS pp,

'" Thr Roir "/ Tiff i~ \"orlhrrn O,rumpol", £roS)'Slrm.

8<ufal1. W.R. (l9.s9) S"od><1 of tiff aDd ",be, facton ,da'od '0 l><k pone'''f<ncmion '0 """lItrn ~, Mo<l"S"'. I'IJ.D lbna. U"', ~lIch.. AnnArboI. :'>b<h. 11'0 PI'

lkaafDt. W R (l96Otllnn__ of~ lJ"tlsMJ s;,.. T _. utel....", Fiff."" O'-"-ws..ro·.."'........ ba.ohlana. \1JdI. Oq> C " For [)O, '.Lamol1J. l>11CIl "9 PI'

Bea.,aJI ..... II /l96Ob) s.oa.. d,"" of hi'" t.m~ ...lgre> OIl lilt 0lIIIn and ...m ofJXl< 1""'". F~ Sa. , 19-1-199

8<...... Jy, (l91JI Ja<k po......~ S,__ jo#dw .!,f.",. FOffSI T.pn of• ,;~ s-,., l:S Fo< Sen _Apx. Haa<I> ,. Pl'-~

1k...... J ..... n<r7l\h a jl· ..._foIiod_ tbo~""'''lIl;wco>_l:S

To<. Sm. T«II. Rq. \C·l,I. II PI'o.--a.J.K"I~) F",........~.... hod.ap.solml_andjKt _. LS F.,.

Snr.. lin. I'q. L.S~"b, 12 PI'Dr"""•. G. (19'" S". ""."""aliOO laboltx<j.... PI_,.,_ C".I>J"h_.,

S.--, ea.. For So... _S).., I'roc 0-'-5. PI' .16-17.81UJn". Dl>L ..... ' <1 t R (Irllbt Ili<IoricoI iluonrooro of IU.~

and JIUI<: • ~ eI••, ClIaIa RJ",,- O.ulrio. C_. F«. Stn /,,( R'J'PS-X-«J 18 PI'

D}ram. G.M (l~, Combv>'- of lor... twlo. '0 ...r. D.." Cod I. fOl't'Sl FwC~_ t'.... 'kCn..-II,U. 'On.- "ork. Pl'. 61-39

Ca~"",. H (1~31 ".1"", pooo, of ,I>< boodmg "",,.na! ,q lo<IJ<'polc and lad< PO'"conn. C.... Dq.. R.-, Dn-no".. For. BNllCk.. StI, ..... Ltorfl· 80 J PI'

Carroll. S. (1978. n.. 10k 01. fi,. ," ,I>< jad< po....--licbo" ..ondlanoh or ,I>< I\,b.ba"",Plai", RcJi"" of Can.acla. ,n DE, D"b< (cd). Fi... F.roIh,.' ,~ Rn<1O'=.If~'" lI'o,kJhop 1T«Nd"" •• Can Foo-. Sc" . Inf. R.p. "OR·X·~IO, ppa7-l9

Ca"ford. J.lI (1963) Som< lac'o" ,n~..nani I""k p;no «~c ...r."on .f'<I f". ,nsou,h...t.,n M.n,tobl. Con, INp, Fo,.• for, Rd. Bron<:k. Publ. U}/o. I~ pp,

Ca"foro. J.1t (l966al. Ja<k P'" "i.n....,on on burne<l arca>. Con o.p Fo,. Rurol[>".,,!op.. Bi·.~"n R.., ,\'OIrs. U (~). 7,

Cayford. HI. CJ966b) Opo..,)"n,1 ,ri.l, of r.~.n.r>1ion ",c'ho'b lor j.ck p,n. ,n""",hn".,n Ma""obI. Con, f),p. fo, .. For. Rn. B"",rh Publ, /l1>j, n pp,

C.}iord. J 11. (1971) Th. 'ole of Ii.. '" tho .rolog) .rt<! "I,·kullur. of jlCk pin•. hor,Tall T,mbo.. F"•. £roJ. C",,!-. 10. ~21-2-l-l

Ca~f"r<I, J,ll. Ch""""..,C1. Z,. and SI,",. H P (1%7)" .. , .." of ;<1",,,1'0,.1'.""ICh 1n Jxk pin•. Con. CHI'. Fe>. R"",'On·,lop.. For R". BrwKk Pub!1/73.:155 pp

Ch",......ICI. Z. 119591 Con,rolled bu,n,,,. .,pemt'IOnt> on jad pono >I'~. ('... (XI'\onIt. Aff..,. \"","R.-., Fo<. B"'ndI. Fo< R". D" 1..:11. \(lft'~ 19P1'

Ch..,.......ICI. Z. l196Jl n.. .flett> of >I" oa lad< poIX puootb ,n _born Ont.no.C.... Dq.. FO'. For Rrs B,..dt PtJ>/. IO/j. ~ pp

CIlrmao....,.. Z. 1196") bpo~ buml"l for hum... <hl.po>aI "" .....,"',,actpm< ",.. III <em.... 0.-. C.... DqJ. For. __ DoT-dop. For B_II PtJ>///&/ 13 PI'

Chrnociow1I:l. Z. (19"\11~ '" "",t pm< ~ boo""", -.I_. " ..........."" <k...<vt " ..... "'n"a1 Qauono, C_. For. So" . lot[ R.,. O·X·/P Il PI'

~1CZ. Z lira. E,-.-- "''''''.1'''........0 >«chd>fu<,...k P"'" «F.....bOll ... CC1lUIII Oeu:ano. C.. J. For Rn ~. ~-'5--lS,

~.z.Il~ISlatllaDddull.--~·__ .. do._1I"k po_....... ""'tlooftU'" Munol>o. C_ For SIn· I..,. R,.,. \OR·.\··/W II""

'"CIl"'.........oa, 7.. (1973b1 Sla!Jl and duff .fll""'l011 It} b,m"n. on c1.."...... Jad< puI<'

.. ,ts 1ft DO",raJ Sal.k~""""":m. e_. F"". 5m _1"'_ R<p_ \OR·X·](J(} I~ Pl'~oa. Z 119"'8c)~';:"iar.-l ........ ""oJm.bdi>powol_mnlf'"

,"poodut_ III """If" 1lIl. e... F•. Sm- l'fj, Rtp. '-OR·"·YJI"w

Cl••c»c......a, Z- (191'&11 Slh~ __ alii" ... _nkrn Cuad&. on 0 £Dubtledl, FIIr UoI<>c:<'"~" ,'_OiI ll""bAop~_ C...To<. xn , ...... Rq>. ,"OR_X·110. lIP _"-'6

c.~"11U. LC U9'TIIn..~~ oI8aml11 T....-notup. AI"""",,,, ....k e.. JBot, 55 1~:.l-1~18

0.). R L and Iloood>. GT (1"-:-) Tk toItoiWlldf,t< ... 'bt~ "'jack."" fC'dpt... 1of"",. '" Ou<IOCO I''''''nctal Park. Oou " .... 'or. Rnoo.,_. Qwn<-o /'",.f'd,l FIf. £~ Siwf. ~p. 5. "'9 pp

o..und>. J. II 1191>1) U~ 01 fi,. in plan"". 5<10 ~P'''''IDII_ Pnx, Lilk' SMrn f""Tu.lmpr-o'- C""f., 6, 22-n

Ell". L. \\ (1911) Sorne nolo, "" lad, pont '" .'<11.,. On'ario. For. Q"~'" 9. 1·[ ~E"•. F.1l (1938) ClIO jad pm. bt r<grner",d .,,1'>0., Ii",? 1. Fo,., 36. 10t0'·lVi'E'f<. F.lJ . Ind UB.,,,,n, R. K. (19-14) Manag.",n, "I j.ck pin. >land, ,n 'he I.".,

S'al". liS Iffp. A~ri._. T«h. Bull. /l¢.l, 1>6 PI'Farra,. J.l.. Gf~l. O.W . and A,·or,. D. (19~) h<l:. pine roprodu'h"n. I'ulp Pop,

Mo~. Con 55. 13()...1-16F<»ler. \\ T .. It"boon. G.A .. Ca~f<>rd, J II . O"k""" ItA. and ~lut>c~n, A I'

(19671 S~''''""",u,,, on ",,,,,,"bed bIlrn,n,. I'ul" P"" \I~ <<>n.. " ..... I~F",,~II>, II R 119M) S,"'" of f<wu T...... of!h<' l'n/ltd Su>-It:l. L:S F'" Mn A'....'

Ita""" ~'I 701 PI'F~,- J \\ and famo•. J.l. (Jill) Etk<:r of;/lade "" j.ad: ""'" __. e...

Dq> Rt<L>rU DtwI.. TO'. &--*. TO' R... On . SJ'r. U<If1. 8 PI'F"""U. S 5 J. om),.. ....~ of~ .. _ Ulunl ewk>p:aoI taeror '" 1ta1a

Slal< Park. "l' "n. fl-. R.. l Yn-4lnGnp. 0 F . arid \kColI. J.G. 09""51 lAw- fd afw...tldi... lD "'p forflu of.................... ~"n ,0"'. C_. J. TO'. R.... ! ~~lfkiDo.r_. \1 L(1'169)~ oftbr ...... ...-n·'bd , . \ __, ~ :-IJHcms<_. \1 L CI97lllln1onDl fir< to "'" """l.." d th< ~.n \\.I<n

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Quon,illio. D.. F.h \,,,,, •• G R,. and Dubt. D E, (19171 Fi,. b<ha,.,ar ,n upl.nd Ja<:kpi"", ,'''' D on uk. p<OJ«I. ("QQ. Fo,. .5<"',. In! FI,p, ";OFl·X·I1~ 09 pp,

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US F",. Sm , FIrs. I'Qp. L.~·I Ia ppRQlh F OM). 11>< j..,k "''''' pl .. n, 0/ M,chiga". fo,. I",~ .. S, 013-116,Ro~c. I.s. (19721 f"'NI FI,~iQJIl of Canada, C.n. For. 5<", .. Publ um In ppRo~e. J.5.. aod Soot,... (; W (1973l For. ,n ,"" bo:>rt:.1 fOT<". Qua' Rts. J. ~aa-at>a

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s.-.l. AJ. 11"<' "u..t Flit 0«""",",,, ,w C_, C... Fot So" .00..... a.ODunol_l.

Jack Pme Fore5/5 ""Stock,. B.J. (1970) Moi"ure in the fo,e', f!oo'----it, d,,,,,butoon and mowmen1. Can

For. Sm'. Olla~'o, On,or;o, Pub/, 1271, 20 1'1"S'ock.. B.l. (l97S) The 1974 wildf". ,,,"",Ion in north".,tern On'ario. Con for.

Sm . In! Rep. O-X-2J2. 27 I'P.StOC~', B J .•nd W.lke,. J.D. (1972) FifO b<n3"ior and lutl ron,umption in Jack ptn.

,lash in On,."o. Can. For. Sm'. Inl- R,I'. 0·X·169, 191'1"Stock,. B.J.. and Walker. J.D. (1973) Gim.u. oondition> b<fore .nd du""g lour

signlfican' fi,.. in Ontario. Con. for . .<;.,,,,.. Inl- R,p. O·X·I8i. 37 pp,S"ain. A.~1 (1973) A h,,'o" of fire and "egetation in nonn.a".rn Minn.""."

'e<o,dcd In lake «dimen". Qual. ReL 3. J.83-396.Van Wagner. CE. (t96S) S,O,)' of.n intense crO~'n fire", Pe,."."a. Pulp Pap. Mag.

Can .. 66. 3S8-.l6LVan Wagner. CE. (1967) Season.1 ,'ari.,ion, In moi"u'e ronlent 01 .."ern C.n.da

Ire. foli.ge .nd p<»sible effect of crown fi,... Can. INp. FOF. Ru,al D,\',lop..For. BFanch Pabl, 1024, 151'1'.

Van W.gner. CE, (1973) Rough p'ed"l1On of fir< .pread Ta'" by lucltyf>C" ("anFor, Sm', 101- Rep. PS-X·42. 8 Pr·

Van Wagner. CE. (1977) Condi'Io", for tli. >lnn and ,pre.d of c",,", n fire. Can. jfor. R.,.. 7.23-3-<.

V.n 'l',agner. CE. (1978) Ago-<I""" di"ribu,ion and the fore" fire cyde. Can. J. fo,.R,.'.. 8. 220-227.

Ihlke'- J D.•nd S'ock'. B.1. (1971) Analy'" 01 '''0 1971 ",Idflfe, In On'ario'Thacker",' .nd Whl"lo L.ke. Can, 1'0'. Sm'" Inf. R~p, O-X-I66, 13 pp.

Walk<r. J.D .. and S'ock>. B.L (1975) The fud campi.. olmarur••nd Immature pckpine "and, in On,.,io, Can. For, Sm'. Inl- Rtp, V-X-2N. 19 Pr·

Walk.r. :S R...nd Dobb,. R C. (l'!6ll1 The use 01 pre",rib<d burmng in jack pmemanaooment in """hea".rn Mani'oba. Con. Dtp. for. Ru,al DeI',lnp.. ForBra"'-h. 1m, Rtp, MS-?4. 15 pp.

Woo.j,. G.T .. and Day. R.J, (977) A ,umma'" of the fi'e erolo~)' .rudy of OUellooPro. ioci.1 Park. 0111. Min. \'a'- Rm>h'.. Qher;oo Pro\'lnClal PaFk fire Eool. S'ud,R~p 8. 39 Pr·

Ye>lman. C W (1979) S'atu. of jock pine geneti.. and brcodin~ In On<3rio. in TruImpm)',m'lII S",nf'O';hm, C.n, For. Se"'" Sj'mp Proc ()'P_7. pp. 127-1Y>.