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TABLES

No. Title

1 Hours per day in which temperature at depth ?.5 cm exceeded15'C under three site condit ions

2 Percentage probabiliiy of receiving monthly rainfall ...

3 Percentage recovery ol P. cinnat@mi Jrom infected soil samples

exposed in the field for monthly periods ... .. .

4 Soil compositions used io determine influence of soil type onP. cinndmomi activi,ty

5 Mortality of Bo,nksia groTrdis seedlings associated with varia-tion in soil type, watering levels and inoculum levels of

P . cin Lamoltui

6 Rate of spread of P. cinnamorLi infection based on under-

storey systems 1967-19?2

FIGURES

1 Dieback' desiroyed loresi . . .

2 TTpical distribution of jarrah dieback in

JaEah Forest

Page

29

5B

64

65

67

69

the Northern

ihree surlace

10

t7

19

20

2 1

22

23

26

27

28

28

30

3 Soil moisture characierisiic curves of thehorizons of major jarrah forest soil iypes

4 Topographic and soil proflles of the Scarp Road Study Area

5 vegetation. Stratum 4, Scarp Road Study Area

6 Vegetation, Stratum 1, Scalp Road Sbudy Area

? Weekly rainfall and soil moisture levels at the Scarp Iioad

Study Area. 31 August 196? to 18 January 1968 . .

8 Average weekly soil temperatures from 31 August 196? to 4

January 1968 at ?.5 cm under canopy and in an opening

on Stratum 4. Scarp Road Study Area

I Weekly rainiall and soil moisture levels during 1968-1969.

Scaru Road Study Area

10 Number ol hou.rs per week soil temperatures at ?.5 cm were

above 12'C at three locations at the Scarp Road Study Area

11(a) Number of hours per week soi l temperatures at ?.5cm were

above 18'C at ihree locations at the Scarp F,oad Study Area

dudng 1968

11(b) Number of hours per week soil t€mperaiules at 7.5cm were

above 15'C at three locaiions at the Scarp Road Study Area

durins 1968

Number of hours per week soil temperatures ai 15 cm depth

were above 15'C and 18'C under canopy and in an opening

at the Scarp Road Study Area during 1968 .

t2

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No.

32

Title

Spread of P. cinnqm,oni irfection based on symptom develop-ment in the understorey species oyer a 5 year period. Ilighspread rate

Me&n amual linear spread of dieback infections for 19 plotsoYer the period 196?-68 to 19?1-?2

Dead B. grand,is zone, which is formed following disease ex-I€nslon ... ..

Dense canopy coyer from understorey of dense B. aquiroliulnwhich originated after severe fire

Eealthy forest showing typlcal iuegular stand structurewhich results from "group selection" management .... ....

Page

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Figure 2Typical dlstrlbution of jallah dleback in the nolthell1 jarrah fore6t. (Non_timbered areasoccur on lower topoeraphlcar sites aD.l hence merk drainaee rinii. iv,jiJ ct,iie assocls,tioa

between these 6ites and jallah dieback).

Research has shown that there are some exotic eucalypt and coniferousspecies which are resistant to the fungus and capable of producing aneconomic crop. These species, however, grow most successfully on the lowertopographical jarrah forest sites that cauy low quality jarrah. It is possiblethat new economic methods of establishment, cuuen y being investigated,will allow economic replacement of the diseased jarrah forest crop on freelydrained sites with resistant economic species. It would be more desirable toretain the established jarrah crop on these sites.

1d

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SECTION I

ENVIRONMENTAL FACTOR,S AFFECTING PATHOGENICI,IYAND SURVIVAL OF P. CINNAMOMI

Eeptine (1964) has defined the environmental conditions Iequired beloredamage can be expected by p. cinnamalnr as foltows:

Is this often alimiting factor.?

Hosts woody; great l.ange of families.nonoyesyesyesyesyesyes

Characteristic

pH lange for. g1.o\rth and infection pH 4_g.

Survival pool in soi l dry for long periods.Long sulvival usuai ly requi les a l iving host.

Growth slo$' below 1b"C, optimum at 2?"C, ni l above 34.CInfection sl ight below 15"C or over 34.C ..FIee soil moistur.e needed for long periods lor infection.Damage rare on l ighi soi ls, common on we!, heavy :oi ls.

-Eiven a superficial analysis of the jarrah forelt envir.onmeni iD lelationto the disease indicates that several of Hepting,s , , l lmit ing factors,, ar.e uotopelat ing. Thus, a more cri t ical evaluation of factors affect ine survival andpa lhogen i c i Ly o f p . . i nnamt m l i s nece :sa t . v

1. FACTORS AFFECTING PATHOGENTCITYSoil moisture, soil texture anal soil aeration.

SoiI moisture, soil texture and scil aeration ar'e intelrelated anal theirefect on the pathogenicity of p. cinnamami is complex. Numelous authorshave shown that high soil moisture levels are required for pathogenesis(Roth and Kuhlman, 1966; Zak, 1961; Torgeson, 19b4; Hine ei at, 1964).The association of dtseases caused by p. ctltnamomi with conditions of poort irainage and/or excessive Iainfal l have been observed by Anderson (19b1),Campbell and Copeland (1954), Torge on (1954), Copeland and McA]pine(1955), and Newhook (19591. Stotzy et al (196?) shotved the necesslry torhigh noisture conditions for pathogenesis but, also demonstrated tha[ zoosporesul 'vival was inhibited at high soi l moirtule levels. Torgeson (19b4) loundthat soils of moderate texture tvere most favourable for pathogenicity butZak (1961) d€monstrated progressiv€ly greater pathogenicity as soit textur.ewas made heavier ' . Zak sugge:ted that soi l aeration was not an rmporlaurfactor in detehining pathogenicity.

Roth and Kuhlman 0966) summarised cuuent knowledge on the effectof soil moisture on pathogenicity. They postulated that there \rere two thres_holds atrecting pathogenesis: soil saturation which qlovides the stimutus forsporulation, and fleld capacity which enables spore migration anal possibll,influences infection. Data on the effect of soil moistule and associated lactor_son pathogenesis, ho\r'ever, were inad€quate for' critical evaluation of flelclconditions. The moisture chalacteristic curves of tesi soils were not. tpecifiedand there was no information on the periods Iequired for signiflcant diseasedevelopment at moisture levels above the critical minimum. In the abaenceof data on the moisture character.istics of test soils, a moisture conren!OJ 20 per. c€nt may corlespond approximately to . .wi l t ing point,, or. . .neldcapacity", d€p€nding on soil texture and structule. Even if the moisture

.-

Suuds aql Eurrnp olnl€rodlua? Iros ul uot?tsnlcng ltsurup elqeJoptsuoc sr oJaql 'eteurlc addl u"auEtlaXrpey\l € lll .ap€ur oq uEc sel€lurlc "in1"""a*"1 go *a,o11 -EnlB^o Ierrlrlc aloJaq sler\al (c.zl) lunruqdo doloq pu€ (c"BI) umrulldoqns '(ColU) lunrurldo oql uoa]!\loq alnl€laalrual Sur1eu,ra11e lo q"nJ"'"Ul aulurrolop o1 peJrnrjal orE sorpnls r€lruns .lue^oleJ aJour alE 1o"*oot"n"p asEeslp ro] s?rurq Jartol eql a.raqa sol?tull, al€Joatural Ur uorl€nle]Ia "ar"a1p o1 ,ro,1rc --rldd€ arl1rl a^€q €lEp aqt ocueq pu€ -{1rcrua50q?3ci r7.,o147)UUXC d' rOJ s?ru I oJnq€Jedugl Jaddn oql q?rdt paularuoc eJg_{\ Salpnls aseqJ Co6I 1ts dISnO -nutluoc ldo{ era.u slrl€lal eql uaq.& perlncro uollcnJls"p 1ar"" ,"o 961 1nq 'qluoo. auo roJ solur'ol Jnoq zI uo c.gt pu€ J,6I UaOA\1aq paJtsuJaJIE alol\ slu€Id aql uoqrrt uorlcn:4sap loor luac rod Og dluo ""* "i"q.L .Juorudole4-op esEoslp loJ urn.urldo suor?rpuoc lE pleq aJa4{ slu"tcl aq; l€q? srnoqJo roqunu aqq oi pal'Ia.r aq 01 ,{lrla^es ".r",rp p.rnoj ,reqi 1ue*roolaa -ap as"aslp uo soJnle-redural Iros Eurlenlcng Jo lroga er{l p"1""1iaru*"p ."1oo,-eurd Jo loJ ?oo.r pu€ lJ"oq uo suolle:Ilsa^ut "1nqt.r1

j119611 i" 1" n.rr11

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1E sB C"8I lE 4uenbaJJ s€ p.rrqJ auo diuo sE,r puu J.gI ^\olaq Jncco 10u prp uorrcerw punoJ eH .UO4CaJUr JO olqgci€c so.rocrsooz Jo uorlcnpoJd aql pu€ poJnpoJd Er8u€toals Jo sroqrrnu on"^1nq p"q"ra:.,,1a,p

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0^r1ts1e3a^ .roJ soJnl€Jodrual IEcrlu, aql paqsrlq€lso 199611 {ooq^"}.I pue eoqcernlrrodr(Iel IIos

.pessoss€ 3q osl€ lSnIII rncco 01 uollroJur 1u€cu -ru8rs JoJ le,\ol IEcr3rJc eql oloqE sle^ol O.mlsrow 1e peJlnbar elull Jo polJacleq1 {reqlrnd srllfi.ra+rEJEr_rc aJnlsroq lualeglp qlrnd sFos uI srraua.SoqlEclJOI SIalaI Ornlslour IIoS IecrlrJc 3q1 Jo uoll€ulurJalap eal0aur lsnur llrcrua8 -oq1sd uo ornlsroq llos Jo s?caga aql Jo (ror1€nlEr\e elrsuaqarctuoc v .sq?€ap

lqanorp pro^e 01 .uollEnlrs plag aql ur sJncro qrlqd\ leq1 ueql ra?rorls qcnua uoaq altsq lsnu-r But.rele,r uaa,ujaq l€^Jo1qr oqqsnllJ uorltsrnlEs pue ,.dtrIc€d€c- piau,, uoal\?aq sE,\\ IIos aql qrrqr{ aurrnp 'scllslralc'rEqc

lros lEcrs.{qar uo .8ulpuaarep ,sporied .aurd.rea "i"*'"*rq3 a1,r"* -leoJl osoql Jo lsarrp aql ur rIo^E.(196I .IE ja orllH :996I .u€rrlqnx puE qloE :T96I .>lEZ) BulraXB,{r uae,\{loq l€^.ralul or_l? Furlsntp€ dq palsel dllensnarol! ,{lrcua.3oq1tsd uo saull8ar olnlsrou ltos luaJaglp_ Jo slcoJe oqJ

perrddB uaaq lou o^€q ,ron".,o" ","ffito':i,;rtiiil."ltii,,,.il"t"',.il; ""lfi&":;Iros aulfop oJ pa.reprsuoc oq ?snul dlrdrlcnpuoc arlntsJpdq IIos pu€ selJnccllslt3lrE.r€qc arnlslouI oql qloq lsql po1€.rlsuouop (896T ,0961) roupJ€c 'alEnbap€ul s€,{r (lqroat Futllr,rr pu€ dlrr€d€c pIoU) "1"oq- ""n"1""io* y,o" 1o suogducsep rrl€1s Jo esn 3q1 'paqrrcsep 3ro.r sllos lsal Jo se,,in, ,rlsrralcer'llc

months. For the period of the year when soil moisture conditions aresuitable for infection. there may be a total of six hundred hours duringwhich soil temperature is above the minimum for disease development. Theaverage period per day duling which the temperature is above the criticallevel, however, may be only eight hours. It is unlikely that ihe processesinvolved in the production of zoospores, their transfer to the infection"court" and subsequent infection are favoured by disjunctions. Ilence, theassumption that disease development is directly related to the cumulativesum of the number ol hours above the critical minimum temperature pro-bably results in an overestimate of the suitability of a particular environmentfor infection.

Soil microorganisms

Zentmyer (1965) shorved thai bacteria, ]rolably Chromobclcteriumviolaceunt, stimulate initiation of the production of sporangia. Marx andDavey (196?) demonsirated that ectotrophic mycorrhizae act as deterentsto root infection by P. citunamomi. Zentmyer (1963) demonstlated controlof avocado root rot by incorporation of alfalfa meal into the soil andattributed this to the large increase in saplophytic Jungi in the ireatedsoi ls. In a later paper' (zentmyer and Thomson, 196?) i t was shown thatsaponins could be part ial ly responsible for control. Kuhlman (1964) showedinhibition of fungal growth was less in forest soils than in cultivated soils,but found that none of the tested soi ls had a lethal effect oa P. cinnamomi.Roth and Kuhlman (1966) studied pathogenicity in eight differeni soils anddemonstrated that the fungus was more destructive in heaviel soils, buitound litUe evidence that a saprophytic soil microflora was actively influencingdamage by P. cinnamoni.

The influence of host susceptibility on the critical conalitionsrequireal for pafhogenicity,

The host, pathogen and envilonmental factors of the disease complexare not independent. Prediction of the degree of disease severity, in envi-ronments which are optimum, sub-optimum or marginally suitable fo'_ thefungus, cannot be made without considelation of the susceptibility of thehost.

The effect of dlfierent host susceptibililies on the environmentalconditions required for severe disease is illustrated by comparison betweenavocado and citrus. "One Phatophthora infectioD on a. gr-owing avocadorootlet destroys it, whereas in citrus massive infection is necessary for dam-age" (Stolzy et al. 196?).

The ease with which the fungus is able to invade ihe root system, thecapacity of the host to regenerate roots and the host's ability to withstandstresses br_ought about by fungal attacks ai_e ihe plincipal factors deter-mininghost susceptibility. The more susceptible ihe host is in any or aU of theabove respects, the lower are the environmental requirements fol fungal patho-genicity and sedous disease development.

2. FACTORS AFFECTING SUR,VIVAL

Soil miaroorganisms

Conflicting data on the ability of P. cinnamo,nti to survive as a soil sapro-phyte was presented by Kuhlman (1964) and Zentmyer and Mircetich (1965).

Kuhlman found that P. cinnanami was shortlived in soil, mycelial growth

1 4

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d Jo uorl?,lrlc€ur Joj rfiBsserau ararlt (Cogt a^oq€) aJnlEladlualqgrq .{Fnonur?uoc Jo spouad €uol 1tsql paleJlsuouap (t96I) Ie la aurH

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uI le]ll^rns xluo,tuDuulc d uo suorlc!.rlser oql Jo uoll€nl€,to a}€lncr€ ,{\oll9 ox.{J?ssarau sI saddJ Iros lua,raJlp ur slalal €rnlsloru Iros tua.reJlp lts spouodlueJaglp JoJ a]rt]lrns o? sn8unJ aq1 Jo dlrc€d€r uo uoIJEurJoJuI .salsurrlc

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e-mlsloul lps'uo$ore pa8qolord fq +noq€

lqSnoJq slusru€E:roorcrlu o^lllladluoc Jo acuosq€ eq1 ol anp s€dr .V.S.n oql Jols€a-q1nos aql Jo slros ur ,{Ielruuepur ?dl^.ms 01 !1uo1uouulc .d aqt Jo ,{lllrq€eql peJaprsuoc (196I) )t€z sar"cads un.nuna B qlr,r palelncour uaeq p"q{JO.{OCaT 4SsrOOd aql 8qt^t5 IIos aq? sE '1u€lrodull aq pFoc susluEsoorcrurpalElcossts 1tsqt peJsasBns osIB sorpnls s,uEurlqny .sn8unj eql pal€ururla J€ollauo roJ sSullpoas snlllr o? Sulddolt .sorpnls u,$o s,qcrlacJrw pue JafwluszuI luEuodwl oq 'lo^a^\oq'plnoc suorl€Indod ,ruE8lootcrur Ilos aq1 ur sacu€-Jeglc anbruqcal uorlcalap Jo dcBnboptsu oql ,{q ]J€d ur paur€Iatxe aq plnot€l€p drolorpEJluoc eq,jl olqrssod otadr ,aJnlsroln

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The major physical environmenial elements controlling tungal pathogenicityare soil moisture and soil temperature. It has been assumed in the evaluationof the environmental conditions of the forest thai ,.field capacity,, is thelower soil moisture level for infection and tha,t 12'C, 1b"C and 1B.C are theinitial soil temperature levels below which mycelial growth, infeciion andsignificant infection, respectively, does not take place.

LI

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lualcuts eql ]o sJoog ,{altEd aqluo 'spu€ldn aql ruol] odolsudroo qldop 1€ seuoz pt ed puE ,{Eic panlolu ,{qur€Irapun scrlozpod cr1r.ra1€[ arB slros puEldn aqJ .[r€Idauad petJrldn ,luercu€

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INflIAINOUIANS ISIUOd HVUUVf SHI

II NOIICSS

and have a range of pore sizes which remain filled with water, and henceavailable fol zoospore tlansport, at relatively high soil potentials. The threesoi ls exhibit favourable ( lateri t ic si l t) , intermediate oater, i t ic gravel) and un_favoulable (sand) moistule retention fol p. cintua.nlami acli.frly -

Field obselvations ol the disease and evidence fiom the litelalure sug_gested the valiation in soil t€mperatule and moisture regimes were the prin_cipal factors affecting variation in disease distr.ibution, intensity and spread.Consequently, an attempt \ias made to measure ihese regimes undel a vanetyof site and stand situations which were leptesentative oI the fotesr envuon_ment-

The principal experimental area was located at Scalp Road in tne llwel_lingup Forest Division. This area provided a r.ange ol site anct vegetationtypes that are typical of the cutol,er jarrah forest. The area $,as disease-freebut was sullounded by diseased forest and thus could be inoculated withoutendangering large areas of healthy folest. The soil moistute and soil tem_pelarure reglmes of stand and site types not Iepresented in this study areawere sampled in other areas of the forest but it was not feasible to moculatethese sites.

1. SOIL MOISTURE AND SOIL TEMPER,ATURE REGIMESUNDER MATUR,E UNEVEN-AGED CUTOVER FOREST IN

DIFFERENT TOPOGRAPHICAL SITUATIONS

The experimental ar'ea was located 8 km north_west ol Dweuingup, ap_proximately 0.6 km east of the Darling Scar,p. Experimental plots were locatedon the south-west side of a shallow valley formed by primary erosion of theancient latel i t ic peneplain (Fig.4). The site types in the experimental areawere representative of the major. types found in the ancient peneplain, exceptthat there were no depositional sancls.

The soils ranged fr.om the freely drained coarse gravels of Str.atum 4 tothe poorly dlained deposit ional si l ts of Stratum 1 (Fig. 4). Stratum 3 wasfreely drained and had soils similar to Stlatum 4, except that the surfacenoruons were formed of colluvial material and massive ironstone was lessprevalent. Stlatum 2 \ias a site intermediate between the freely drained soilsof lightel texture of upper. stlata and the poorly dained, heavy textured soilsof Stratum l_

Jauah was the principal component of the overstoley on the upper !opo_graphical sites (Strata 3 and 4) with occasional malli. Bclnksir- grarrdisWilld. occurr.ed both as a scattered undelstorey tree and in dense thickets.XantlLorrhoea preissii Endl. ^nd Macrozanlia rie(llei (Gaud.) C. A. (.ialdn.occul led prol i f ical ly. There were a number of shrubs but the shrub laye!. wasdiscontinuous. The forest on the upper topographical sites had been manageclaccording to a group selection system and consisted of groups of difielent ageclasses varying from poles to over-mature trees. Jauah saplings and advanceglowth (r 'egeneration) wele absent (Fig. b).

In Stratum 2, mar.ri occutred more flequently in the overstorey and B.grondis lormed a scattered understorey. The shrub species were similar tothose on the upper strata. In Stratum 1, jarrah was absent and marn anaiblackbutt lEuco,lvptus pdtens Benth.) formed an irr'egular ovelstorey. B. gratuaLis

1 8

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Figure 5

Vegetation. Stratum 4, Scarp Road Stuoy Area_

20

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^I{aad\ paplocel sE:t[ IIgJU!€lI "OJ€ I€1U31u -IJodxo aql uo suoillsod uedo aaJql q pallElsul olaa saSn€8 ura,tf-.l.lD!u)D?I

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Fisure 7-Weekly minfau and soli moisture levels at the Scarp Roaal Stucty Area Al Augusr 196T-1g January, 1968.

uI s1u01uoc oJnlsrou ,{\ol aqJ uorlceJur perno^"J qtoq sla^al arnlEJaduel IIosp(I€ arnlsroul Iros ueq.\r sporlad 5uo[ alaa elaql .la^a$oq 'z puE T €l€.rls uI'qqrJlurll

1ou s€r!{ t lunt€lls uf arnlslolu IIos uaqdr sportod Jalrq aql SuIJnpuorlcaJur JoJ lo.\al IEcr?rJc aql arloq€ trtlq8rls,{po elad!,{dorr€c rapun Ea^eloJnlgJadlual Iros sorlllqrldacsns ar\llElaJ Jloql uI socuaJagrp pa{r9rll eJoadslaql 1€q1 paltscrpur EJ?tls aalql aql ut uorlceJur tol alq€1rns 0lo,{1 suoillpuoceJnl€rodlual IIos puE arnlsroul Iros qcrq.{t Suunp spouad aql }o uoslJpdlqoC

'?arv irpnts p?o1l dr?rs F unl?rXs uo Surusdo u€ ur pu€ ,{dou!crapun urgl1€ 896I drEnuet t o1 196I lmanv Ie uoJJ salnluadual Ilos,{I]Ioea a€?ra^v

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'a.rnlEJadluol Fos uo ddou€c Jo lcoJa pa{JBlu

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.qldap urc 0t ol qIeql 18 U€ql reia.ol aq ol papual Bl€rls esaql Jo uozrroq acBJlns eql ul s[3^alaJnlsloru 'sqluoru re1utuns aql SuIJnp "{Ilusnbasuoc sal€r rart\ols .dllqalls 1€paloldep s€d\ a.mlsIou lros t pu9 Z €l€J1S uI 1ErI1 ldacxa'uozlJoq actsJJns oqluI esoql ol JBllruls oro^\ uozlJoq Illcot 01 sI aql ur spuell oJnlsloru Iros

'luatuarns€aur Jo porJed eql lo lsour lnoq5norql dtrcBd€c

pleg e^oqE slartol atnlslou IIos pautBlur€tu 'qros eql Jo sarllr€dtsc Surploqernlsrou qSrq eq1 pue a8€ur€rp algold 01 ocu€pedur qlrdr JaqlaSol rornlsrou.r

Jo acrnos l€uorqlppE sIqJ sa?Is ltsrrqdtsJSodol Jaddn Iuol] o8€dees pu€ II€Jur€lqloq uorJ arnlslou pa^rrop z pus T ElElls lueueJns9aur Jo porrad aq1Sutrnp sl€ ralul JaIJq fluo roJ ,tllcEd€c pIaU oloq€ araa t unlElls Io suozrJoq

I

Stratum 4 duling the summer months suggest that fungal survival in soil oDthese sites could be lestricted. Soil moistule i4 Stlata 1 and 2 never ap_proached levels which could be considered lethal to p. cinnamarrr.

This preliminary data suggests that difierences in the susceptlilility ofdifferent sites in the jalrah forest, appalent from field observation ol jar'rahdieback. could be part ialry accounted for by dif ferent so moisture anar soi lbemperatule regimes. The lack ol sampling intensity in both t ime ancl spacedid not per.mit precise evatuation of the susceptibi l i ty oJ the dinerent si tes.

( i i ] DETAILED MEASUREMENTS, FEBRUAR,Y 1968 TO APRIL 1969.Prccedure

Rainlql l-Weekly r.ainfal l was lecorded at three open posit ions in the exDeri_men l a l a r ea .

SoiI r taisture-Ten 40 m by 20 m plots wer.e located on the experimental area.Three plots were located in Sttatum 1, two plots were located in each ofthe interDediate sit€s (Strata 2 and 3) and three ptots wele located in Stratum4. The plots 1ler 'e selected with iongitudinal axes ol iented along the contour-Four soi l samples wer.e taken for gravimetric determination of moisLul.e conaen!f lom the 0 to 15 cm hor. izon in each plot at randomly located posit ions. The15 to 30 cn horizon was not sampled, as the presenca of massive tr.onsroneboulders on some sites plevented unbjased sampling. Sampling 11,as contrnuedat weekly intervals throughout the measurement period except on ptots Iocatedin Stratult 1, \ ihei-e sampling was discontinucd dur. ing the winter months whenf teF \ a t c r ' occu t . r ec l on t hc su f l ace .

Soil moistur-e content r, 'as determined by oven dr.ying at. 105"C. Whilethe soi ls remained moist, samples wele col lected with a reinforced King tube.When the soi ls were dr 'y a sl lecial ly reinforced soi l auger was usecl. . .Fieldcapacity" of the freely drained soi ls in Sites S and 4 was determined byaveraging the soi l moisture values for the eight_week period in spdng, fronlAugust 22 to October. 10. The ,'fleld capacity,, of the soils of the two towelsites was obtained by deterrnining the soil moisture content aft-.r 48 hoursfree dlainage f lom a saturated condit ion in the laboratory.

The moistur.e characteristic curves of soils fr'om Strata 4 anal 1 weredetelmined using the pressure membrane appalatus.Soil temlrerature, ( cL ) Continuolts meclsurements_Continuous recording doubleplobe soi l thermographs were located in Strata 1 and 4. At each site theprol les were instal led at ?.b cm depth, either under canopy or. in an ol leningin the forest. Standar.d glass soi l thermometers wele inserted at each pt.obeposition to check thermograph accuracy. Check readings were taken weekly at10 a.m., 12 noon,2 p.m., and 4 p.m., except during the winter months whenreadings wer'e tak€n $,eekly at 10 a.m. and 2 p.m. During the spr. lng monthssoil temperatules at 15 cm depth at. the probe positions were measured weekly$ith dial-type metal thermometer s.

At the installation site on Str.atum 4, probes wele located under dilTelentcanopy covers-complete canopy and no canopy. I t was impossible to locatethe tuo pr.obes on Stratum 1 under markedly dif ielent canopies anal so oneprobe was located under pat.t ial canopy and the othel in a palt ial opening.

(b) W eeklA measurement ol sltr|d.ce soil temperature_Soil temperaturesat 7.5 cm depth were measured on six of the moisture content, plots, weeklyat 10 a.m., 12 noon,2 p.m.. and 4 p.m. with diat_type metat thermometers.

24

uorlcaJul lu€Dgrusrs puE (C"gI) uorlraJul '(CqZI) q$\ora ItsIloc.{ul loJ la^alI€crlrl, aq1 a^oq€ s9,{1 arnl€iroduel 3q1 {aoa lad srnoq Jo .Ieqrunu oql altsl-nrl€c 01 posn a'rand. slldBrSouraql Sulprocar snonulluot aql EIorJ €t€o luau.i-erns€arrr Jo polJod oqq rado slE,{ralul fl.rnoq-o,$.1 roJ 'suoltisod aqord aq1 Ioqc€a 19 lqc 0 9I l€ ornl€rodulal oq1 ol€rqrlse 01 pasn o.Ie,{l suoll€nbe asaqJ

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STRATTTM 4 \-'--'\

M I J J A I S o l N D J

PERIOD Of ffEASIJREMFNT

Figure 9'Weekly Iatnfail and soil moiEture levels during 1968_1969. Scalp lioad gtudv

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'uurnlnE uJ ,{lrc€dEc pIeU: or\oqE arond sIaAoI arnlsrou IIosuaqA\ spol.rod alqeJoprsuor roJ cozT a^ocl€ oJorf sarnlEJoallue] (oI srd) se?rseotqq Il€ uI lueruarns€atu Jo por.rod aql +noq8nolqt C"ZI ^\oleq .qsnonull-uoc dlor€r ora,$ satnl€Jedural Ilos-turg,1€ cozT a^oqE {ae^\ laal sJnoH

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SOIL MOISTURE NOIL IMITING STRAI I . ]M,1

SOIL MOSTURE NOT TIM TING STRATUM I

PERIOD OF MEASUREMENT

rigure 11Numlrer of hours per week soil temDeratures at ?.5 cEr were (e) above 18"C and (tr) above15'c, at three locatioas it ttre scarp n6aa Si lav

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.sa?rs Il€ uo

CogI la.olaq dllualsrsuoc a.raid soJnl€.realural ,sqluoru talurad eql gur.rn(

The periods during the year when soi l moisture was not l imit ing and soi ltemperatules were above 18"C on the four strata were markedly different(I'ig. 11a). The higher soil moisture levels of Stratum t had a cooling efiectbut there were still long periods during which soil moisture was not limitlngand soil teml elature was above 18'C. Under canopy on Stratum 4 ther€were no periods when soil moisture was not Iimiting and temperalures wereabove 18'C. However, soi l temperatures in an opening on Stratum 4 wereabove 18'C for brief peliods in autumn and considerable periods rn spnngwhen soil moisture levels were not limiting. Sholt, heavy showers in eariyautumn and late spring depressed soi l tempefatures below 18'C for longerperiods under canopy than in an opening. There $,ere no heavy falls of Iainduring the summer months and hence there $,as no opportunity to examinethe effect of heavy summer sholvers on the number of houls soil temperalules{ 'ere above 1B'C in each of ihe sites.

Houls per week above 15.C and 1B.C at 15cm depth Soil temperatur-esat 15cm under an opening in the canopy commenced to r ise above 1b.C and18'C in the second week of September and the second l l ,eek of Ocrooer. res_pectively (Fis. 12).

Under canopy, holvever, although the soil temperature gradient with depth$as les:-, soil temperatures r.emained continuously below 1B.C throughout theperiod of measuLement a4d nere only above 1b.C for bt. ief periods.

lb) weekla mettsurements dt 7.5 crrl dPpth--_Ihe weekly soil temperaluremeasurement at 10 a.m. and 2 p.m. on the tena.perature plots in Strata 1, 2, 3and 4 provided the same trends as lor the continuous recolcl. Sampling intensitywas not ad€quate to show the significance of diferences during the criticalperiods of the yeal and none of the plots sampled had sufficient carropy coyel:to demonstrate the effect of canopy on soil temperatures.

5 0 L M O ] s T U R E N O T L f f I I I N G

t 5 a

t 3 c;]

o

) E R I O D O F M E A S U R E ^ I E N T

Figure 12l iumber of holus !q \ \ eek scr l temrrcratures at t5ch depth wcre above 15!( j and 1B"Crrr locr caDopr aDd [ ' Jn opFtr jng at rc Scarp Roa. i Stualy Area dur i ] rg 1969,

30

'Elerls lse,!\ol aqJ ur 'aJojeraq'I, sarnl€.radual flos uo alnlsiou] Iros Jo 1c3Ja

allssoJdap fulr roJ poltssuaduoc raururns-prur oqm sla,lal elnlsroiu os q8rq JosCualSISJad aqt 1"ql aunsse 04 elq€uos€ar sr 1I

'p01se1 1ou sErt\ T urnlBtls ur

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Suls"aJcur qlr,{r ssal sB^l saJrs o,{1l aq4 uaar$laq acueJallp aq1 q8noqttv

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fI arnSIJ

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'-ddou€t

Japun u€ql uorl€nlrs uado oql uI ralEoJS sE]l{ qldap qlri luorpEtB ornl€roallualeqJ 8T aJn8rd ur po4Erlsnllr oJE t lunleJls Jo rror?Els 5ulpBal ,{Isnon!41uocoq1 ol luac€lpts sol.s uo qldap rur00€ plle g'ZZ 2,e ud t 1€ popJocal salnl€'Jodwal llos ilaeat -4+dop I1Lc 0'0t pup 9 zz ?D sluautarnsoaul n|\aaftl (c)

even under conditions of fuu canopy cover, ther_e would be long periods whensoil moisture and soil tenpera'uure conditions would be expected to be suitablefor infection by P. cinnomonri. The susceptibility of Stratum 2 varies markedly,due to rapid changes in the soil moisture regime ol'er short distances tvithinthe str'atum. In the lower' section of Stratum 2 (see preliminary soil moisturemeasurements), the ]evel of susceptibilty $ould be similar to that ol Stratum1, but in the upper sections, susceptibility vould be similar to that of Strata3 and 4. In Strata 3 and 4 the maintenance of soil moisture levets suitablefor' infection was dependent on the input of moisture from rainfatl. Susceptl-bility in these strata was influenced by the deglee of canopy cover. In allpericds when soil moisture was not limiting, except in late autumn lvhen hea\,:fand consistent rainfall occurred, soil temperatules were below the cdtical levelfor ' infect ion for longer periods in sites with canopy cover. than in sites withoutcanopy cover. A Ielat ivcly greatel volume of roots is susceptible to infect ionon sites without canopy covel because the difielence in soil temperature levelsbetween the two cover types was st i t l appat 'ent at depths of 22_5 ancl 80.0 cm.Stlata 3 and 4 \\,ere relatively mor.e susceptible in spr.ing than autumn becauseautumn rains had a depressive efiect on soil temperature.

Soil moisture values on Stfata 3 and 4 in summer apploached levels whichcould be critical for' survlval of the fungus. SolI moisture values in Strata 1arrd 2, howevet, did not approach levels which could be considered lethal toP. cinndmoni.

2. SOIL MOISTUR,E TR,ENDS UNDER HIGH QUALITY POLE STANDSStudies 1 ( i) and 1 ( i i ) at the Scarp Road exper. imental si te indicated that

moisture in soils located in upper topographical sites falls below ..field capa_city" before the soi l tempelature under caDopy I ises signif lcan y above thecdtical level for. infection. Soil moisture measurements, however, wele tes-t l icted to the 0 to 15.0 cm hodzon. The susceptibi l ty ol forest growing onupper topogtaphical sites is important, because these sites car'r'y the highestquality forest. Therefore, a further study $,'as conducted to provide informationon soi l moisture trends under even-aged high quali ty jarrah stands glowingon upper topographical si tes.

The Study Area

The experinent $as located ?.S km Dorth-east of D$,el l ingup. Tne u,ansec!$'as select€d on the north-east side of a f lat-topped hi l l , which dlolrped to asaddle on the north-east ancl a small stream on the souih_east sicie. Aprofile of the nolth-east slope of the hill showing plot location and soilcharacterist ics is pt.esented as Figure 14. The stand is i l tustrated in Figure 15.Procedure

Four 40m by 20m plot: tvere located along a t lansect from ridge topto the saddle (Fig. 14) and or. ientated so that their toneitudinal axes welepalallel 1\ith the contour. Soil moisturc content was samltlecl at six randomlyiocated positions fron the 0 to 15 cm holizon on each plot ovel the ledodAueust 20, 1968 to December' 1?, 1968. Sampting of the 1b to 30 cm horizon\^,as car'r ' ied out from September 1?, 1968 to December. 1?, 1968. The moisturecontent at both sampling depths on a.ll ptotj was also sampled on February2, 1969 to obtain an estimate of soi l moisture levels dudng the period ofmaximum moistur-e stress. Samples were taken $eekly except, where indlcated.Soil moistule content was detelnined gravimetlically.

.r{r€nrr€f -p u ul dn8uqlar\CIts peprocoJ aroA\ uorlElrdlcatd Jo soJleruqFw uaslxrs .ZU Joqolco tolJs n"J -qr"r 1u€cglu8rs ou s€ad eJeql .raquoloN ur papJocer uIuI tI ruoJJ utsdv

slFsau.flo^rlcadsal .sqldep

_ ulc 0t ot qT pu€ ErcgT01 0 eql JoJ 8 Joqolco o? l,I Jaqrue?das pu€ I Jaqolco 01 6u lsn8nv sporrad€ql Joao uozuoq puB oqrs qc?o JoJ sonl€a olnlsroru fDJea,$ oql Bur€€to^?.tq paunulalap s€id,tlrc€dtsc plag ol BulpsodsaJJoc 1lroluoc oJnltlolrl aqJ

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6utroi))!' qrr^ a^or6 r^.aH.u.ot ^ot) ^PUb. uaorg 196l

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.t:^br6 aebo)'uoot rpuos uaorq tq6a

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ta^or6 l^$N .uool

roP rpuDs uao4-'l!'ppau

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'€0dFuor' tbus touolsor)o

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Petrpu ao|3^ - ^a)6 a od

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slullw

lo' 'l"o

l"o 'lno -

l"Ii2

'lo,i

I'o -

190

II'ol.olon

---+-- e--;f+z_-FrJ

Jarrah pole :iTj: 'J,"0"

^*, ,.

Field capacity tevels for each plot and horizon. are plotted with ihecorresponding moisture content data in Figure 16. Field "upuciw t"vet" inplots 1, 2 and 3 rere approximately the same, lut tne vafue' in"plot 4 wassjenif lcan[Iy higher.Soil moisture levels were consisten y below field ca.llacity in the 0 io lS cmhorizons in aI1 plots, as early as the first week in October fiic. iii. aru.o,r.l1

9e

'69-896r - reluluns pu€ 896r €uirds Furrnp $lrs r€crqar€r€octol raalarn IIo Fur,ldo.rs spu€ls slod q€rr€l r{ul€nb q€Iq lapun uozlroq uic 0€_9r pue tui sr_o eqt ur sF^aI ei.isro_ rros,rrae"u

9I arnAl.{

6II

t

p

{l;-*--n--=*--*.;zi

soil moisture data for the 15 to B0 cm clepth were less variable than at thesulface, the tlends were similar. Moisture levels l.emained at apploximatelyfieId capacity in plots 2 and 4 until the end of Octobel but by the first weekin November they had fatlen significanUy belo$ field capacity. In plots 1 and;1, soil moisture remained at apploximately neld capacity until the first weekiD November. The rapid fall in soil moisture after the nrst week in Novemberin plot 3 suggerts that sarnpting bias $as Iesponsible for the higher soilmoisture levels recorded in that week. The maintenance of soi l motsiure atapproximately fleld capacity levels in p1ot. 1, however, cannot be attributedto sampiing error and probably reflects slight impedance and/ol seepage orwater from upper topoglaphical situations on this site.

F,ainfall in the tveek ending November 19 cau:-ed soil moisture to be]lighel in plots 1, 2 and 3 but the levels were still significan y below fieldcapacity at the t ine of measurement. Soi l moisture levels on February 2,1969 were, in both horizons and in all plots, approximately the same as thoserecorded on December' 1?, indicating that the soils had reached maximumdeplet ion by this date.

Discussion

The results of this study confirm the conclusions on the soil moistur.eregime of upper topographical sites in the jarlah for.est summarised in StudyI (ii) and indicate that valiation in soil moisture over a range of slope posi_tions in a uniform pole stand of high quality is minimal. The maintenanceof soil moisture levels equivalent io field capacity on these sites is dependenton ihe consistent input of moisture from rainfall. The soil moistule tlendsin the 15 to 30 cm depth are particularly signiflcant, since they indicate thatthese holizons fall below field capacity at the same time as, or soon alter.ihe surlace hol izon!.

The soil temperature measurements of Study 1 (ii) are considered ap_plicable to this study, since the rainfall and soil moisture irends in both ex_perimental areas wele pr.actically identical. In the absence of intor'mationon the soi l moisture trends in the lb to 30cm hor. izon in Study 1 ( i i ) , i t wasimpossible to calculate the number of hours during which conditions weresuitable for infection at a depth of 15 cm. The results of the present stucty,however, when integrated with the temperatui.e data at t5cm of Srudy 1 (ii),indicate that it is unlikely that signiflcant infection occurs at this depth inupper topographical sites. The decr.ease in soil temperatule with depth isnot compensated fol by an excessive prolongation of favourable soil moisture1eve1s.

Comparison of the soil moisture levels at the end of December with thoseat the beginning of Febluary shows that minimum soil moisture levels in thesurface horizons are reached eariy in summer in freely drained jarrah foresfsoils. In the absence of precise data on the efect of moisture on survival ofP. cinnolnLolrti in soils of different textures, it is impossible to determine ifthe minimum soil moisture levels reached in this exper.iment .were trelow thecritical level for survival.

3. THE INFLUENCE OF ASPECTRoth and I(uhlman (1966) demonstrated in Douglas_fir lorests rn Oregon

that aspect had an importani etrect on the "oif -oi"tu"u "rrJ i"-O".utu""regimes of difierent sites. They fouhd that sites wltf., "outf.rel" aspects trad

36

palcosslp dldaap ,{{aI € ruoJJ 1r€dv sam}B.radrual lros Sururlurolop ut lu€Jgru8lsllqFlq or€ edols pu" lcads€ ol onp secqoregrp l€!11 papnlcuo, sqocaf ,{lodll-cadsar 'aJloulrluac arEnbs Jod sarrolEc 209 puE IZl, sl .{BJaua IE?o1 €ql .0IJO SedOlS UJaq?nOS pUB Uraq?JOu UO OJlarulluaC arBnbs rad SaIlOt"C t8g SIpunorS Ie^aI uo roqolco uI pa^recar tr8Jaua ,{lrpp I€?o1 aqJ uoll€lp€r r€los Jouorld:rosq€ aql uo adols puB Jcadse Jo ocuenEu! Suud.oqs 'frolIJJaJ

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uorssncq('luourorns€oru

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| !_ . { NORTHERN SI IE

O-- - - O SOLJIHERN SI IE

MOISTURE NOT L IMI I ING INTHE o- l5cm HORIZON

--o- 'a

i F y € i f i i 6 # # * i i i ? , f i B +PERIOD Of MEASIJREMENT

Figure 1?Weekly soil temperature trcnds on northeln enat southenr aspect6.

river valleys, slope a,ngles in the Jarrah forest area rarely exceed 10. and areusually le6s than 6'. Thus it would be expected that differences in soiltemperatures due to aspests in the jarah forest wolrld be somewhat less thanis indicated by Jacobs,

This study demonstrates that soil temperatures are higher on northernaspects and it is Ukely that soil temperatures on these sites would rlse abovethe cdtical level. for infection earlier ln spring than on southern sites.

Itre characteristic pattem of spread of p. cinnartuottui, infections, tha,t isupslope from moist sites, makes tJ:re effect of aspect more important, Onnorthern slopes, because the fungus destroy6 tJle vegetation as it progresses,the sun's lays penetrate ahead of existing infection, uninteryupted by canopy,aDd would produce higher than normal temperatures in the infecdon zone.

- - 1

6€

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lseroJ aI{1 uI arr1tts.rd luaura€€utsru peqsrlq€1so u€ sI SuluJnqpoqFcso.rd esr€ds s€A\ Ja^oc Jallrl aq1 'dlluanbasuoc 'puts qurnq uolldutsold

.{Irtsln8ar uaaq p€rl qclqnd s€oJ€ uI palcnpuot arala salpn1s qlog paJnssau

lou araa sarnl€radrua+ llos 1nq spuels a8€-us^o -dlrl€nb qBIq repun spuarlernlsloul Uos uo €1Ep papr^ord Z npqs spuEls q![o.r8al q€JJ€f pa8E-uo^a

,tlllgnb Jeq€Iq eq1 Jo aAll€luaselda.I lou se'$ 9o:r€ trpnls slql lncl suolllpuoc

aXrs Io o8u"r € ra,!o lsoJoJ .,uollcales dnor8,, o.Inl€lu repun spuar? eJnlE-raaiual Uos pu€ ernlsloul llos rro €l€p papl^ord -{pnls pBoU dl€ts oqJ,

S{INVJS U'IOd HVII'I{V-C AJI'IVOO HCIH HECNNSCNEIIJ, SUNJSIOru ']IOS (INV IdIJOJVTISdTAIfiJ

'rlos No AdoNvc cNV usrJra do sJcs,{ds sHJ }'-{ll^rqc€

IEBunJ .roJ elq€llns eJB olnlslour puE aln?E'radural Iros qclqnd

Suunp spolraci eql lo ql5ual aql Sululllualop uI re^ot lallll pu€ dclouPc 'o!ul8a.I

arnlsloul s€ luElrodrur sE lou sl lolt€J srql lnq .{lllqrldacsns ells Jcagg s3op

Iradsv slcads€ luereglp uo osEaslp aql Jo dllJo^es eql uo lo^ardoq 91Ep ou als

oraqJ (896I 'ro8pod) s1c€dsE II€ uo punoJ eq f€ul uollcaJur ?1.uloTunuuxc d'sal-Is uraql:rou uo sel€l Suldlp prd€r alou fq lasJo

eq plnor lcads€ Jo lcaga aJnJe.realdel oql qEql alqrssod s! 1r pu€ luaulladxaslql qllrrA uollaunluot u! peprocal 1ou arerd sluarualns€aur alnlslou] uos

'lI

It '

1

Litter tayer tn forest unburnt ior approxihately 40 yeam.

layer was discontinuous. Me,croaanzia ried,lei a.Ird pteridiurt esculentunl t,I.orsL..)cccurred abundanfly ar\d Clernatis pubescens t]tsluec), nfnbiriia montana(Steud) and Le1!.capasan rerticiltatus (R.er.) occurred ;"";";;;uii;.Procedure

The experiment had a randomised block design with flve blocks and tworrealments. Ten 24m by 12m paired plots were establ ished in the experi_

Figure 18

urnq 01 alqlssod lou sts,|{ 1I Bulu.rnq Jo 1ta$a aq? alBlnruts 01 pa^o(llel ara!\Jltsd qc€e uI lotd uesoqc ,{Iwopu€J euo uo sqnrqs lle puB la.{tsI rollll 3qJ

'paltstncltsc stsA\ ar€lcaq rod

lqara^l rallq o8€Ja]tE aql pu€ lold qc€o 01 luactsfp€ ^Ial€lpouu! slulod 1€ ua{€1aJana JolM aql Jo salduEs o^rJ slurod pal€col .{ta^ncalqns 1€ €arts lBlua(u

'urnq paqur$rd AulaoIIoI IIos peue{c€Iq 'ore€t

6I ernru

0I

tf

Jar*h pore I'iH: :r"o" ^,"" ".

the area because of the high moisture content of the litter layer. ,I.he litterwas remoyed by raking and all shrubs were cut of at ground level withsecateurs. Each plot was subdivided so that each block consisteA of twoa d jacen l l 2m squa re p lo l s .Rqinfu l-Weekly rainfall was recorded with gauges located at a fixeci positionon one of the subplots of one block ancl with two gauges which were randomty

:":1t_:-g :":n rl€ek on remainine subplots. Rainfati * *"u"u."J L a clearins

u .+ Km t rom tnF expc l im rn ta . l a rea l r om Oc tobe r 23 . l 968 onwarc i s .

42

tt

slualul€rl lurnq pu€ luinqun rsBvn ra,t€I u, 9I-0 aq1 fi sF^aI 3rn?srou Iros ,{I{aal^IZ arn3ld

tNtw;Sfsvlw .lo oorurJzL,_u ltltz ltreL oLroE ol,r9l ol/z a/3t 6/, vta

a

s

;

q

I€lorx adfl-Igtp qlr^\ papJocoJ sts,{r qldap urc 0.9I 4" alnl€radua? IIos .urJ g, Jo

qldep 9 lE sloldqns oq1 Jo qc"a ul uo4rsod pa?€col ,{IuroprrEJ ? 1E pallElsur sB^\ilelordoruroql Ifos pJ€pu€ls ouo sGlauourJaql Ielaur ad.4l-l€rp paltsrqrlEc qlr^\ricotq qc€o Jo sloldqns l€rluac o^\1 aql uI sqolqrsod peltsrol { ropu€J o^\1 ?€popJocoJ s€Ar qldap lllc g l, 1€ oJnlEJaduel l\os-slualuanLsnaw EqaaA4 (q)

'srala(uoruJaql lglaur ad,{l-IBtp pal€rqrl€r qll,{ oqord qc€a 1D ua{E?

ara,\! sSurpEal {caqc trDlaadrqldap luc al, lts ,{la^rlcadseJ lros e.r€q puE JaIIIJopun palErol saqord qlri[ s:lcolq oql Jo ouo uo pallBlsur s€,{\ qdgJEorxraqlIros eqord olqnop Sqap.rocar flsnonurluoc V-s1ua1].Larnspan snonu!?uoC (p)

.{colq qr€a Jo slold-qns I€rlqac o,{11 eq4 ut palns€ew arer[ seJnl€radtual lrog-alnxprodulal .lxos

.(Z'6I) IIEI-ISJtsIAI

pu€ sur€Ifird\ ]o poqlau oq1 .{q paurriJol€p sEr$ lurod €(IrlIUA oq Surpuod-seJJoc lualuoc eJnlslolu os eqJ .896I '9T toqolco 01 896I

.TU lsn8nv uolJ

pouad {oa.u qq8ra aql JoJ suEaru fDlaa^! oql BuISEJa^€ ,tq pouruuaqap s",u,qrttsd€c plard spoqleur cltlallll^tsrS fcl peuruJalap sBAl lualuoo oJnlslou lros'{colq qc€o lo sloldqns ourarlxa o^\1 aql Jo r_IcEa ur suotlBod pal"col flruopuBroA11 1€ ua{4 er€At suozlroq luc 0 9I ol 0 aql Jo saldru€s lros-alnlsxotg lxos

fIil

lnermometers at points immediately adjacent to the standard soil thermometers.SoiI temperatures at 22.5 cm depth were recorded with "t""iri"ut "".l.turr""blocks at two .andomly rocated nxed posltions in "u"r. "r,toro1. -i.'o

t""r.r"n""were dug immediately adjacent to the five blocks. Tne iltter wus removecifrom around one of the trenches. Thick polystyrene st eetlng was useo tocover the trenches between readings. Temperatur", "t- :O"O-irn alepth wer'elecorded by inserting metal thermometers horizontally into the sides of thetrenches. AII temperature measurements were carried out weekly at 1 p.m.Results

L_it!:r -

The aver.age litter weight measured was 16 tonnes per hectare.SoiI Moisture-The avelase weekly soil ,""i.tu"" ;;i;;; rrr*'oori ,."ur_"l,r"are plotted, in relation to the determined n"fa "uoulttv, -irl "i,"nu""

,r. "omoNture in both treatments was below fi:ld capacity lr"i"n" ui the end ofseptember and beginning of october and consistenfly after october 2J. Litterremovai caused soil moisture to fluctuate rr1o." in l""por.r"- io-"""urrrru,r. Uo.the first weeks of measurement the moistur.e values in plots with litter removedwere higher that ihose with l i t ter present (Fig.21). During drier perrocts soi lmoistule levels were slighfly higher in plots with titter intl,ci. Oinerences inthe moisture trends of the two rreatme;t" "u" b" ;;;lu;;;; tv 1n" rr,.,n""irrgefiect of the litter and shrub layers on soil moisture. Rainfall interceptron isgr€ater and evaporation less from surface horizons in ttae pfo* with litterpresent. Aithough soil moisture levels were consisten y leto'w neiO capacity

1iL,,i.::n"*"x'.;litr'Jil:l'*1l"TH1n#._::l*il1*J:ff ?*:fi ;;until November 6. Soil moisture teached wilting point-ly miO iecember inboth treatments.

S O I I M O I S I I J R € N O T L I M I I / N G -t

,/auartP

- 8 ,

tr/9 Bb 25/9 -

2nO- 9/lO r o l r r 3 I ? o l t 2 / t t 4 , 2 t t t l , 8 2 2 5 , 2 t l

*"-o"To3*ttsi"".ir",,$*iliir *.."jfl+'#t*-i,*. *r *****.*.,.,*,,".

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o. '..3

uosl.r€duroC slold ,,luJnqun,, u€ql EoId .,turnq,, ?qt uI uotlce]ul roJ alq€llnsspolJed rotuol oroia. aJaql a?BtlpuI €1€p arnlEradrual snonuquoc oqJ

'Jad?I qn.rqs pu€ra1M oq1 fq uollclaJ.rolur Jo asn€caq stold ,,lurnq,, uI u€ql slold ,,Ju.rnqun,,ur lrlurl l€crlrrD oql o,!oci" eJnlsroru Ios SuISIEJ ul e^IlcaJa ss]I aq pFonduolqcaJul roj EaAaI ltscrlrJc aqt e,roq€ a.I€ sarnltsradual fos uaq,{l Sulrlntco'Jo^oA{oq 'II€JuIE.r

IB(oslasuIl -tqlcluaSoqqEd I€8unJ uo lcaJa Jlaql Jo surreluI aIqEJ"dUOC aro^! Sluaur€arl oia.l Aql u! SIaAaI Ornlsrotu IIOS

'UOll€rOd€,{A

pu€ IleluI€J ol a4llrsuas e.roru o.rand s1uau4tsarl .,1urnq,, aql q;noqllvuorssncs!o

'uo!lJajul ol Surlrrutt 'Jou se\arnlsloM IIos uaq^! poiJad eql aurrnp sluorul€a4 .,lurnqun,, pu€ ,,lulnq,, uIfirr 0 0€ pu€ urD g ZZ 1E sarnlErodural aq1 uoand.loq ocua.r3Jlp lugculu8ls ou sts.{taraqJ suols€cco aorql trIuo uo solls ,.1uJnq,, lspun laq8lq ,{Ilu€cgluBls eJeaqldap slql 1s saJnl€Jodual '8uIlruIII

1ou ss^{ aJnlsrour IIos uoql\ polJod aqqEurrnc salls ,.lurnqun,, oqg ul roqSlq ala,t\ tuc 0'gI 1€ sa:Inl€ladrllal Ilos'trutslllq qtdop qll^r pasearcap ,{aql lnq luEcgluS(s dlletllsllEls aJa$ salls

,,lurnqun,, puE ,,lurnq,, ropun slo^al rrn]EJadua] I(os aq1 ur saJualsJlc'pallo4uoc dlt€rrlsrl€ls lou arorll ruroot 1€ paprocar saJnl€Jodlual IIos.{Dlaa,\,tuEatrI luo[r1€aJ1 qt€o uo sluaurorns€aur uropu?r 0I Jo sutseul aql luasa.Iderqldap lucal l€ a,inlEJadural Iros rol €1tsC .{Ia^I?cadsal 'C.t0 pu€ C"t0'J,tr

0 oreA qldap qc€e 1€ EInsoJ roj sltsAralul ocuapuuot lua, rad 96 oqJ,'€e arnBIJ uI patlold ?JE tJJc a ZZ puts 09T

'9, 1€

sa:rnl€radlual $os fDlaar$ u€our eqJ,-s?rauarnspaw arnlDraduial los n'Ltoa&'Euaur?€aJl,,luJnqun,,

eql u! d?IJoAo qcns ou s€itr oJoql lnq 'SuIllruJI

lou s€r\ aJnlsloul Iros ueql\olrs ,rquJnq., oql u! sJnoq ZI Jo pouad E roJ CogI aloq€ aJoa{ soJnlEJadural IroS'(ze 8Id) -{Io^Ilcodsal 'salts

,,1urnqun,, pu€ ,,1urnq,, aql uI t Jaquracsd puE

eI raqru?AoN €ulpua sleei.. aq? tlun C.ST a^oqE ,{Itu€culu8ls esu o? aculu-uroc 4ou prp ornl€redural IIoS-urcal1B C.8T aloqE {aar!!. rad srnoH (rrr)

'I raqruiJac Sulpua laa,!\ eq1 .{q s1ueurltserl qloq q o9I a^oqts ,{tsno-nulluor stsar pu€ ,{lplder asoJ eJnltsJodurel IIos

'1no -{.rp ol petuauwoc sllosoq1 uaql\ '( sJqtz) 1u0!u1eoJ4,,lurnqun,, aql uI utsq1 (sJq0rT) lualultso;l,,lurnq,, 6q1 ur (CcaI o^oq€ aJod. sarnq€rodrual IIos qcrq.{\ Sulrnp) 8ur1rlurl

lolr s"dt eJnlsloul uaql\ spor.red ra8uol ,{Ilu€curusrs aJelll\ er3qI slueu-leart qloq !4 6 Jaqo4co Illun C.SI a^oqts ,{Ilu€cgru8rs osrJ 01 atuaruuroc

?ou prp a.rn1€radluel lros-ruc q l 1€ CogI o^oqE Iaat. .rad sJnoH (II)'Surllrurl

lou s€.$ a.lnlslolu Ilos pu€ CoZT a^oqe aJa$ soJnlEJodual lros qcrq.{\Suunp squarulEorl rlloq ur spolred Buo[ ora,$ aJaqJ raqlrl ropun spouedJeBuoI roJ CoeT e^oq€ aroiA sarnl€radural 'Sulrds aqt uI JalEl 'qnq qualul9arlpoioural rallrl aql ul spolJod .ra8uol .roJ C"Z] ?Aoq€ s€,$ arnlE.radual oqltrIt€rqrul (UZ €!d) Eurrds dIJEa uI C.ZI o^oqs fllu"culu8ls aslr ol pacuaut-uroc sa.rnl?rad'"ual IloS ulc a I lE C.ZI a,rocl€ >laa.{l. raal srnoH (r)

'(c.BI) rror?celul lu€cutu8ls puts (c.9I) uortcoJur '(coel) q1,\oJ3

I€rlordw .roJ slatel lEcrlrJr edllradsal eql e^oq€ s9dr. ornl9Jadlrle+ 0q1 :Ioaa\ .rodsJnoq Jo .roqurnu 3ql sE patElncl€c ara.$ qdEJSourJoq] IIos 0q1 woJ] €lEcr

'-{r?ssacequn paJaplsuoo sEla sSuIp€aJ qderSoureql eql Jo lu3ruqsntp€puE C"to u€q1 ssal s€,$ s8ulp€ar lcaqc aql uroJJ aJnF9dap aSB]a^E or-IJ'lUeIIeCXa S€rld. 'S8ulp€er

lcaqr trploeia. oql fq pel€crpul sE 'qd€r8ouuaq1 IIos

eql Jo dcErntc€ e\\J-Iuc 9 l lD lualualTtsDow arnlpJaduroJ lzos snonu?luoc

5 0 T M O S I U R F N O T T M I N G

9AO )610 23AO 3aAO 20Il 27 ) 4n2 nn2 )aA2

!

|19 lA19 15t9 2IO 9/\O t6AO 23/lO 3c't:o 6/rPERIOD OI MFASI]REMENT

Figure 2j

20nt 27111 A1l2 |[2 )8n2

Mean week ly so i l tempera tures a t ? .5 .m, ts .Ocm,22Jcm and 30 .0cm depth under unb mtand trurnr trearments du ns spdns rsoe ant su;;,!i r*&-8lie6i'

lipnls aq? ur saJrs paurElp,{IaalJ Jo snBunJ oql o1 dllFqrldacsns aql uo Ja^oa.(dou€c urnlurx€ur Jo lcoga aql eruel{ salts dpn+s aq? ur palEcol .{nuerua^-uor .roloc osuap pEq qclqrll. s€al€ lsetoJ q€.u€! ou aJor\ aJoqJ ra^oc ^douBtasuep Jo s€e.rE apnlcul 1ou prp 'trarg

lsatoJ oq1 Jo,{}rro[€ur eq1 Jo adr]"Juos-ordar qSnoqll€ 'sarpnls asaql ur paldur€s ra^or Jaqlq pu" fdouBc ]o aSuBraqJ. salrs paurEJp dlaatJ uo uorscaJur JoJ alqellns aie sIeAeI aln?sroul FospuB ernlEJedual Iros qcrq,{ SuIJnp J€ad aql Jo spouad aql uo ladoc Jallrl pu€Idou€c Jo lcega po>ll€ru eLil psleJlsuouap a]!oq€ paqucsop sluaqrrlaalxe aqJ

scNvJs Hvtl€vr QNv vJvlova d,-{o ss] IISg€S2fNIVIISdIAISI CNV qIIOISIOW 'IIOS SAI.IV?IVdIAIOC '9

'luAurlEoll ,,1ulnqun,, 3q1 JO 1Bql Ueql

Jaq6lq s€,r qnq luarqqEall ..1u.rnq,, puEls alod oql lo 1€q1 ol JEInuE s€,$,{pn!spEotI dJ€cS aq1 Jo t unlerqs uo olls ddouBr eql Jo arql8ar arnlEJaatruol IIosoqJ Jaqolco dlr€a Itlun slodal J€I urs qcEar lou prp sJuaurlgall ,!lurnqun,,puE .,?uJnq,, qloq Japun putsls olod oq? Japun sa,rnl€ledulol lros aq1 lnclraqualdos {lJEo ul spolrad +u€cgiuBls JoJ (c"gI) uo!?caJul roJ Ie^oI I€crlrrc0q1 oloqe asol alls p€olI drecs eql JE uor?tsod uado aq1 uI se.rnlEladrual Iros6qJ lsaJol lurolrlrn puE JEInSallr Jo sarulSal oJnleJadurol IIos a]\rl€Iar 6ql Jouoll€crpur euos sadlS (€e pu€ ZZ s8ld) dpnls srql w paprocar saJnl€laalruolIros eql qlr.{l (tI pue ZI TT 0T SEI,{-I trpnts) alrs p€o?I dJEcS aql l€salrs l€crqdEr8odol reddn aql Jo soulSor ernlEJodulal Iros erll Jo uosr.r€duroC'lsoro] pu€ls elod poEE-ua^o Jopun pu€ (I ,{pn1s) e?rs pBo2{ dl€cs aqt lElilsnoau€llnu4s poprocar aJal{ salnlgradruol Iros

'sqluou Squds eql Surmc'IBAIdlns

I€5unJ uo lcaJa s1I ]o srxrol ui 1u€+rodrul oqpino,!r\ scuaJaglp aql q€ql .{io>lrlun sr 1I 1nq

'.salrs ,,lu.rnq,, aq1 uo o?EJ ralsej B

1€ sn8unJ oql 01 Surl|Lrrrl eq pFoc qcrql\ sle4el paqreoJdd€ oJnqsrour [IoS'sdlJdds qn.rqs Jo

uollBraua5or eql pu" ra,(El la1lrt € Jo uorlaJccB aq1 ]o osntscaq 8utuJnq Jaqles.r€od 6ulpaaccns uI osEaJcap plnond sa.rnlEladwal IIos uo SuruJnq ]o lraga aqJ'uorlroJul JoJ lEcrllrc oluBcaq sle4el elnlsrour IIos raqJE Jo 01 loud ?no paur€ts?Ar Sulurnq roqlaqd\ uo 'aeJ8ep E 01 'puodep plnor\\ uorlraJur lujounuue

.d

uo Sulurnq AuIrdS Jo lcaJa oqJ, Suuds pu€ Suru.rnq uumln€ uea,{116q I€l.raXuIarll ur Jadel qnrqs aq1 Jo uorl€rauo8at auos oq plno.{t aJaql pu€ suleJ .r3lulra.trq pa,{oluor aq plnoid ?roJa Suluo{a€Iq aqt ?nq

,sqluour Euuds lEctlr.rc oq?SuIJnp Jaltli ou ur lFsar plnord SuruJnq uunlnv qcrel I ol laqur:c3o portaalaq4 uI dlr,rPurrrd srnrco IIBI raqli1 ulunln€ ro Sulrds ul Eurlg qlr,\\ .uorl€qoJ

r"ef xls ol rno] € uo lsaJo} aql lnoqgnotql lno polrr€c sr Eutulnq peqrJcsord'1ueur_riedxe srql ut peptccar asoql ueql:ralBair8 eq plnoa sso.rB lulnqun u€lurnq IEntc€ uaaa\1aq sacuoraJrp ornlErodwal IIos 1€ql patrcadxa oq pnorll 1r'pauolrelq

lou o]aid slold ,,lurnq,, aql acurs 'snqJ (61 8rd) acBJrns IIos aq1Jo Euuo>lc€Iq sosn€c puB ra,{€I re4lrl pue qnrqs oq? se4ourar ,{lrr€:roduel 3u!-u.rncl pccllJcsaJd dll€rrullrB po^oruar eroa Jotr€I qnrqs puE Ja1lII er{l acuaqpue 'paqErlrul s€,r lueuruedxa 0q1 olrlll aql lE urnq oJ alqrssodutr s€r$. lI

'Ja1€at8 uaaq o,\Eq plno,( solrs ,,lurnqun,, pu",,lulncl,, roJ€}Ep snonulluot aql uaaluaq sotuaroglp eql stsrq srqt lnoqlr,rA t€ql €Icllssod sr1r

'aruoH slold luoluqEo.rl ,,lu.mq,, aql IIe ro] a8€ro^E aql nA.olaq doqSrls aJealold ,,lurnq,, eq1 Jepun palEcol oqotd aql lo] saJnlE,radlllol 1€rll sr!\oqs qoldII€ rOAo SBurp€a.r rtlrleaa 3q,l qlrrra 'ru-d

I +E s8ulp€ar qd€.rSouraql aql Jo

afea was determined by comparing soi l moisture and temperature resimes ofadjacent Piruis rad,iatq D. Don and jar.rah stands.

Measurement of the soil moisture and temllelature r.egimes of a p. re,d.iat(Lstand plovid.d some data on the poiential susceptibility of plantations of thisspecies. Newhook (1959) showed t:nat p. cinnamolni was responsible for wide-splead P. rdd.iota dealhs in lhelterbelts in New Zealand. The fungus alsoocculs in plantations but appar.ently does noi cause widespread moltality.Newhook suggested that the difierence in evapotranspilation rates offull and reduced crowns in :helter.belts and plantations. r.espectively,cculd be one of the factors responsible for the rapidity oI disease developmentLn the shelterbeit si tuation. Hart igan (1964) has reported dieback and mortal i tyin P. radiatq plantations in New South Wales associated wi.f,h p. cinnaiLomi.Although mortality was low, tlte afiected trees were salvaged as the orowmng,which occurs after fungal attack, ,,opens the way for. blue stain and teads todegrade timber.,, Both the New Zealand and New South Wales attacks wereassociated with unusually high r.ainfall in autumn and spdng, respectively.

Batini and Podcer (1968) have shown tinal p. rad,iata. and pinus pznasterAit. deaths in shelterbelts on the Swan Coastal plain are caused by p. cin_namatui. To this date, ther.e have been no reporis ol p. rad,iatI, deaths attdb_utable to the pathogen in Western Australian plantations.

The ability of two to five-year-old p. rl.diatd d\d p. pinaster io sulvrve onold jauah dieback sites suggests that, at least on the freely drained jarrahforest soils, P. cinnamomi infections are not severe enough to cause significantmortality in these species. Following Nelvhook's hypothesis. mortality is evenless likely to occur in matule plantaiions because of the decr-easedevapotranspiration rates of individual forest trees. However, althoughmoltalities have not occurred, and are not likely to occur in mature planta_tjons, i t is posi ible that the fungus could cause severe economic loss bvdepressing tree growth and timber degrade.

The Stualy Area

The experimental area was located 21 km east of Dwellingup where atdal plot of P. rad,iata $'as situated immediately adjacent to poor qualityjanah forest. Both pine and jarrah stands were situated on ; sand flat,sl ightly concave in pronle, which sloped gently (1.b") to the north. The soi lswere of colluvial origin.

The jarrah stands $ere of ir.regular distribution with trees of pile (topheight 22.6m), pole (top height 12.2m) and sapl ing (top height 1.8m) size(Fig.24). The understor.ey component was made up predominan y of jarrahsaplings. B{r7rlasio grri,nd.is arLd persoonia IotLgiJoIicL Fl.Br. occurred infrequently.There lvas no taII shlub layel and the rmall shrub layel was disconllnuous.Litter was spalse and, al|hou]h DrAanclrd niDed. R.Br formed a dense carper lnsome places, thele were consialerable ar.eas of bale soil.

The pine stand was seven years o1d with a basal area of b0.5 squarennetres per hectare and a top height of 9.8 metres (Fig. 2b). Completecanopy closure had occurred and the trees had been pruned to 2.5 m stem height.The litter layer \ras sparse and there were areas of bare roil

Procealure

One plot (40m x 20m) was establ ished on each of the pine and jauahsttes.

48

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'fttsIu!tsr A8tsra,{E JO SpOUAd SUIlnp 1UA1UOC eJnlSIOlU IIOS

a^llcoalsoJ eq1 8ul8€rallE fq poulluJolop oJo^r uozlJoq puE alls q'€a Jo sllos

aq1 Jo ^lrcrrd€c plag oqJ .{fltsclJlaurl.r€r8 pauluJalap 3ro,{'r slualuoc oJnqslol I'lold qctso ul suorlrsod pal€col .tlulopueJ xls ls paldurEs s€x| suozlJoq tut 0 0t01 urcogI puts uc09I 01 0 aq1 JO lualuoc alnlsrour IIos aqa-aln?s?ow l?og

'salpnls snol^aJd Jo! sv-nnl '?oA

'q ,tpnts pu?1s alod qerl€f TEInFaTJI

tz arn5u

pitLlrs r.tdidtti, stand_ StLrdv 5.

30iI Temperature-((D weekla Meas,remellts-soil temperature at ?.5 cm and15.0cm depths was measured weekty over the period ;;;;;'; -;"

December13, 1968 at 2.30 u.m. \rith calibratecl dial_type metal tfr-"r*o_1t"", ", u" r.r."Iandomly located positions in each plot. fl1" t"-p""ui*""ui- a., "_ ur.rO300cm was determined bv inse't ing diar-type metar the'raometers nori-zontally inlo the sldes of a sma.ll pit dug in the "a"tr".f "o"f_r-if.t. ,r,ne pits

50

I9

'ocuBIJ€,,|. Jo sIs,tIBu?

01 polraqqns are.\\ ?l€p alnq"Jadulal eqJ, uI d t 01 'ur'€ 0I uorJ E€lJaluIflrnoq 1e suolllsod pal€col i(Iruopu€J xls 1€ paplocor araa sqldop luc 0'gIputs rxc g L 1" sarnl€rodwal IIos gZ raqopo uo-slualaalftspow nIJrwH (q)

'sanbluqcel etu€lJ€a Josls.tl€u€ 3u6n posfl€u€ erea sqldep luc 0 qI pu€ rrrc 9 r,oql roJ €1€p amltsrad-wal. s?uauraJnseau uoerlqeq Sulloaqs ouaJltlsflod {clql qlIA para,\oc eJe,!'\

'896I ral[rlns puE Al4rds EEIrnp sptr€+s ,?tz16touL s pugottwot'a +uacelpd-iapun FJ3lti Ec ot-gl pu€ uc gI-0 3q+ EI slar!3l amlqoE llos dll3ei$

9Z arnalq

1Nt!'\,ltnsvtw lo 00t3ld

avat ztll av? rrua lvcz lvtl ll/9 ovoe oyez olpl ov6 ova 6/cz 6/at A/tt 6/l a/aa

I

6

o

I

u,a€-4

vlvNtcuvw I )J

(c) Cozttittuous Measuremcnt_Soil temperatures at ?.scm depth wererecorded continuously wiih soil thermoeraphs over the perioJ octotrer g tooctober 16 and October 16 to october 23, 1968 in the jariah and iine stands,respectively. Soil temperatures at ?.5cm and 15.0cm depth we're recordedcontinuously over the pedod November 6, 1968 to .lanuary t, 19?1 in the pineplot with a doubre probe soil thermograph. continuous measurement of soirtemperatures over the whole period could not be obtained because ot malfurrc_tioning instrumenis.

t r 2 5

sor l MoTSTURE NOT LMTtNc i l

{ . 5 . 9 - ,o-

_ E,nargiiata- _ _ _ P . c d i s l a

23i10

OF MEASUREMENT

fiA:1"?"T""iI;{ ;:'}"::?'","?jH'3 ?l,li,)"J;l %.1f"" #;?""fl";,l*" (},]",L',if J}* jli8ft:

(b)

5 z o

t 5

l0

5

'.radoc -ddou"c esuap € dq palEururla

aq ptno^\ salls lseroj q€rr€t puElaln uo ! trtolrlDuulc d dq uollcaJu! 1tluJadplnora. sle^el oJnlslour puB slo^ol alnlErsalulol Ios qclqr!\ Eulrnp spolJad €ql

l€q1 lsaS8ns ^pnqs srql Jo sllnsa.r el{J l-lldJp luJ 0 gI !D luEcgluSls llrls oraa\

slold uaealaq sacuoJaglp o.Inl€.lodulel Iros eql esn€coq Jel€el8 qcnur sr?ia. elrs

qErr€t aql uo pa{cEl1e eq pFoc qtrq^\ slool }o aurnlo^ eql osiE lnq 'a1ls

1s3loJqtsr.r€f aq! uI uollceJu! JoJ alq€llns spollad le8uol qcnd 0laq1 are'\\ '{luo

loN .{llllqrlciacsns uo lalot .{douEt lo qcaJjo paIJBul E salE'rlsuowap uoilcalu!

roJ alq€rnol€J a.reiA oJnlslolu llos puE alnlEreciulel Iros qclqr!\ Suunp spolrsd Jo

srs€q aql uo spuBls q€Jl€t puE auld aql Jo nlIIIqIldOCSns oql Jo uosu€dluoc'lroqs drar aq pFori\ spor.Iaal osaql lso8Ens (Ir) I ,{pnls Jo slFsal aq} lncl'suorllpuoc arnl€.realruoJ IIos pu€ arnlslou Iros elq€.rno^€J JO etuapr'ulot €

sr e.roql uaqra. urunln€ oql uI spollad olE olaql teql alqlssod sI 1I ?7uc1uo1r-u?c a [.q uorlceJur roJ alqelrnsun oia,\\ pu€Js DlDlpDJ d oql ropun suolllpuoc

l€luauruor!^ue 'sqluolu Eullds I€crllrc aql Sulrnp 1Bql OlEtlpur SlTnSa?{

uolssncqo

'sqldap asaql 1E Sulllrrlll aru€'aq suollrpuoc eJnlsroru

Iros ralj€ s{aen xrs puE edg lllun coBT o^oqe asrl }ou plp so'rnleladural Iros'dla^Ilredsai 'q1dop luc0gI pue rutgl, eql JoJ s.moq Bt pu€ t9 AIuo s€ltr o.qT

a,\oq€ aJor! puEls auld aqq uI sa]nlB.raalulal qclq.{{ €ulrnp eurll Jo polrad eql

Sulllrurl aurEDocl slodal alnlslou IIos ralJ€ s>loa,$ o^g ua,!o t€ql sr$oqs (BZ Sldl

I trr€nu?f o] I lequa^oN pol.rod oql loJ €q€p o:rnl€ladulol snonurluoc oqJ

'urc 0 qT pu' ruc q r 1€ srnoq .LnoJ puE ua^os .roJ Ie^aI l.crllrt ""r:l"fil"X::isoJnl€redwol lros

'1old qErl€i eql uI IeAoI IPCrJrrt aq1 aloq€ asor lo,{au lalnl-EJadurel qldep urt09I ]€ pu€ srnoq or!r1 JoJ lo^al ltsrltllc aql a^oqe asor fluo

slolal aurd eql ur qldap tuc a l 1e se.rntelaalual llos luoluaJnsEalu Jo pollod

aql Suirnp rnoq qt€e 1E luEeuru€ls ere.\{ Jaqolto qlgz eq} uo sle^ai ult 0 9Ipue lur g L lB slold orlA.l oq! uo samltsJsdr al IIos ueolUaq atueroglp aqi

'luBrgluErs,{Iq8rq e]alA

uoilraJu!.roJ pa,rlnbar suorlrpuoa e.rnl€rodulal l€trlllc oql 01 uollEler ur soauo-reJrp aqJ, luaula]ns€ew Jo poFad 3q? lnoqanolq? lualsl uo, a.IaM ruJ 0 gl

pu€ tucal1ll uBql ssol qanoqll€'sqtdap asaqJ 1e sacuaragrp arnl€radua] lnqpol€Incl€t aq 1ou plnot ElEp ornl€Iaduel luc00t pu€ lqtqZe aql Jo acu€cg-ru8rs aqJ Juaurorns€3r{ Jo poued aq1 lnoqanolql qldap ulto9I pu€ gl oql

1€ luaroglp dllu€tuluBls alaA\ ould putr q€IIEI ,Iapun salnlg.raalulal lros'IZ arnFrd ur pallold arE sqldap tqc 0 0t pu€

lrlc g ze 'rIlc

0 qI '1!Ic q l lE solnle.raduel u€aru trDlaalll aJnlDJadluoJ I'los

'pu€X.s surd oql uI rer\ol ,{ilqalls aq orpoJEaald€ suozlrol{ qloq uI uollaldop arnlsroru Jo o1€r otIJ ll€Ju!€.r r o.IJ Jnduluo dla,rllue luapuadop sl sle,{.al alnlsloru lros q8lq,{la.u}Elal Jo ecu€uolul€lueqq l€q] puE sollit€dEc a8€rols r$ol o^Eq sllos 0tI1 1l'ql sel€clpul pasEacpEq U€JuIsr a^llrago ralJ€ suozlJoq pu9 solls qloq ul uolloldap a,Inlsroru JoolE,r pral€r oqJ lurod Eurllrrd ol luol€^lnba .{Io1€unxo:rdd€ s€,!\ Mqun dlpIdBJourlcop 01 panulluot ornlslolx I1o: puB alqt8rl€au lsourl? s€nl1. II€Jui€l

'0t

raqolro .ralJv (9Z BI.{) 896I '0€ .Ioclolco 04 dn dlrt€d€c plag ,{laleruFoldde

1€ peur€lureu s€r,\ suozlrorl puE salrs qloq ul aJnlsrou Fos-oJn7slotr4J lxos

sllnsell

HOURS PEF WEEK ABOVE

l5 'C tHr 2 .5 .m DEprH H l5 .m DEPTH

l8 'C . . - -a 2 .5 .o DEPTH O - -O t5cm DEPTH

2utt 27it 4h2 t1/12 :tr .A2 25112PERIOD OF MEASUREMENT

Figure 28Numbc - o I hou rs so r t . cmpFra .u r4 a ? . i .m and 15 .0 .m deD ln undc r ad la r4 p f o4 taLosn.l E. ho'o.aato siorcis were abovF rs"c and r8 c dL.:, e "pi i"e ;r i i i i i l f "u_-.- rrr,u.

Although mosi p. ra(Tiat.L plantations in W€stern Australia are located onthe fertile krasnozemic soils of the river valleys, depositional sands of theiauah forest may eventually be used for p. fttd,iatd, or p. tr)inctster establish,ment. The assumtltion that, the results of this experiment are equally ap_plicable to Nbe P. rad.ic|tct plantations growing on klasnozemic soil would appear'to be valid. The river valley soils, aithough of heavier texture than sand,are freely dlained and usually occur on steep slopes. Jauah dieback has notcaused serious damage on these sites and it is llossible that they are inherenflyunfavourable to p- cinnalrlami. The data derived from this experiment areonly applicable to unthinned stands on which canopy closure has occulreal.It is possible that heavy thinning could result in the ci-eation ol lavourabletemperaiule conditions in spring when soil moisture is not limirrng. tt isreasonable to conclude that p. citund.mami is unlikely to cause damage inunthinned or moderately thinnecl p. rI.d.icrtd, plar.taLians in Westeh Australiagrowing on freely drained soils in which canopy closure has occulred.

The micr,o-envir,onment of unthinned p. pinaster stands duling the spr.ingmcrths would be even less suitable fol p. cinnanlomi infection because thelitter layer .ilndelf P. pinaster stands is much heavier. (2S tonnes per hectareas opposed to 12.5 tonnes per hectale for ten_year_old p. pinc&ter and p-r/tdi@rq, respectively.)

'lcaloj L-{Erlef ulJLllloJ Jql ol rurJelpe puE

ulq1l.\\ suollBls poltoles snolrB^ ro] rEo,{ oqJ Jo l-lluour qt€a uI

sla^el paglceds €u-Ipaacxo IIEJUI€r .roJ s?rllllqEqord 1no slas Z eIq€I (q)

'pouad TL6I-'86I orll JoJ a8€Ja,{€ aql ucrJ s€.rnJrEdap Jo srula} uI

posserclxo sI ElEp aqI 116I ol tt6l polJad aqt r3^o (I'rdV pu"

qcJ€Il) uunqnv pu€ (Joqula^oN puE Jaqolto) Sullds '(dlBnlqod

01 roquacacl) ,roruruns '(.Iaqulaldos 01 ,{etr I) .Ia1ur$ uI (urur9z0

papaacxa IIBJu,r€J qtlqr{ uI s{Ep) sdBp ul€J Jo 'raqulnu eql pu€

po^I3ca,r II"JuIel aql Jo uollEluasarc{a] l€tlqdEr5 € sI 0t arnSld (€)

-i{e.\ oA\t uI pJssa-cl\o Jq ueJ lqaloJ qBr'tEt

uraql.rou aql ui pa^Iota.r IIEJui€r lEnuuB pu€ IEUOSEaS aql uI uorl€Il€A

oltrlJ ul uo-rl€lrea trI€JuI€lI

'as€as-Ip oq1 Jo uollnqlJlslp oql qlIJ,\ pol€lello' osl€ s! s€a'I€ lsaroJ

lo uorl€qlolclxe Jo ooJEop aql 'alalru€xa .rod s.Iolt€] roqlo Jo esnEc3q oleru{c

uI uorl€lre^ qllic. sn8unl aq1 lo uollnqlrlslp pa^Jasqo aql al€Itossl -{IlcaJIp

o1 aiqlssodurl sI 11 lnq'uollenl-rs aql rlc€J uI 'sl s-IqJ lsaroJ aq1 Jo lsoA\

aql u-r s€ar€ ll€lul€r Joqalq oql ur ralBo.lE aq plnoqs as€aslp aql Jo lualxo eql

"{IBUIpTOCCV SEarE IIBJUTEI .raq€lq aql uI uE!I} ssal aq plnor' s"a]ts IIEJU!€I

raao[ aql uI uollce] roJ alq€llns sraa suolllpuot qclqtd Sulrnp spouad a3€

-rale aq1 'st€a.{ Jo pouaci € Ja^o 'l€ql '.ra^all\oq 'palcadxa eq plnor$ lI lsaJoI

qEr.r€i urals€a oql uI lueruaJnsEad lo.IEatr aq1 uI il€Jul€r eBEJaA€ u€ql laq8lq

ol onp s9r. sIqJ sauoz IlEJur€l lualaJlp uI pal€col oJaii naql q€noql ua^o'stsa.rE lpnls eql loJ 4uoJeJIp ^Ilu€cglu8rs 1cu s€rA uolltaJul ?u]culDuulc d totr

elq€lrns sIaIoI 1€ paulelul€lu ser!\ a.rnlsloul llos qclq^\ SuIJnp pouJd rtII

'aln€urllonAc 4B s8ulp.Iocol aql uEqt ssal sr lsai pus ts€e aql ol suollEls uI

IIEJuI€i urunlnts pu€ .Sullds 'I€nuuE aq1 qtlqrd uI ('96I 'qctEII) dn8ulilendc ol

luac€lp€ uorlnqFlslp llEJur€r eq1 Jo elnltld palr€lap e'roul € saoqs 6Z ern8ltr'uroll€aI lEnuus eq1 saoIIoJ,{losolt ac€ds uI uoll€ItBA IIE}UI€-r ralu!'{\ pu€

raurums aqJ, e.rteuoM Jaal luul gT dlal€ullxordde Jo alEJ eql 1€ sasEaJcap

IIEJuI€I IEnuuE 'drets aql Jo clol aqq ruorj sp.I€i\lsBg qsadoals sI apnlllt€

ur osEolcut eql a.raqr!\ Intco e.rleuoll)i 'Iad ulru 00T Jo 'lapro aq1 uI sas€olcul'.{lp]al?r sas€aJcul I!€Jur€J

'do? aql ol dl€cs aql Jo looJ aq} ruol'd dr€cs

SurlrEC aql ol sp.r€r\l!€a 1s€ot oq1 lllo.rJ fll€np€r8 sas€altul II€JuIer TBnuuV

aDtsds ul lroll"Ir€A [I€Ju!?U

'I€AI^rns pu? dllcluaSoql€d IeBunJ ro]

slodel IEOTIIJC 01 UOrl€Ia.l UI Salrs ISAJOJ Jualegrp Jo suorl-Ipuoc ltsluauluoJl^-uo oql uo uorll€lr€a IltsJuIEl JO ?t3J3 eql SSaSS€ 01 pasn eie'{! lsaro] qE'uef

uraq+rou eql uI suoll€Js polcalas 1uoJl €lBp II€JurBJ Jo spJocal Iulel-8uoT

,,8urrds lrlun Sulnulluoc suorllpuoc laiAqlrr\r. uunln€ uI IIOS IO €Ullla'\\al ,{A€Oq .{IFru]ouqE s?,\{ araq1" uaq'[ sresf uI

panncco fluo sll?q.ralleqs D?t)xtrtoJ d uI flllElloul a^Issacxa ltsql punoj (696I)

>{ooqidoN 'eldlu€xa roL{ s}uoluarns€oru o.rnl€radlual pu€ ornlslolu lros uIoI,

pessass€ salrs 1uar3!Ip Jo ftrllqlldatsns aql Jo suollBlnclE' snoauolla qI llnsalplnoc ac€ds pu€ aurll ur stslq €ulldur€s olqeplodEun l€ql elqrssod sI lI

INOI,4IVNN]C 'd

OJ SI4JIS JNSIIS'{'{IC

,{O IJI'IISIJdSCSNS SII,I NO SIAILL CINV ECVdS NI

NOrLOSrtflSrc 'nv,{Nrvl{ Nr Norrvr?rv^ '{o Jcs'{ds sHr '9

s,:

=

s3

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M I L T

3.5 271 105 305 t l 8 339

22.5 4t.8 113.2

Figurc Z0Rainfalt distdbutton at ststions east and west of Dweulnsup_

L9

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(Note: Eff€ctiye rainfall is that rainfaU which is required to ilritiate andmairtein plant gxowth aboye wilting point. fne mean -effeciiv"

"rrrrr*U to"any ore month was determined by prescott,s formula I x 0.? : 0.b4 whereP: efiectiye rainfall a,nd E is evaporation from a tr* water surt&ce,)

6S

€ql l€ql saoqs s.r€ad oArl eql uI sarnl€radrual Ilos aql u3a$1ocl uosrr€druoJ'aEBla,{€ saoqE tilqB.lep-rsuoc s€.{\ II€Jur€r (r3.lolco) Eulrds ,{lr€0 896I ur elrq,\\'33€raAB r\oloq sErlr. l96j Jo (raqotrO) Sul.rds ,{I.rtsa 0q1 Sul.rnp II€Jur€2f

'696I pu€ 896I lO uwnlnts aql Ul 1nO parrr€J S?Uaul-o.rnsEolu eJnlsJadrxal lros pu€ oJnlslou llos urorJ u^{E.rp suorsnlJuo, .{J!po!uol tilelrlun q u[rnlne ul s.r€ad uoadqaq IIEJUIE.T ul uor?erJE4 l€ql slse88ns'sernlE.radrqal Ilos uo II€JuIEJ utunlns Jo ItaJJa o^Issardop padresqo o!{1 qlrd\Jaqlaaol 'dlrlrqEls sJr.IJ '(696I) >looq.aaN dq u^\oqs osoql o1 par€duroc uaqr]a

lIEr s erB IudV pur LiJ.rel I ur podlocal IIEJUI€.I oB€JadE oql ruo.rJ sa.rn?rBdac'eIqEls flodll€JEdruor sI lsaroJ rl€rr€! uJoqtrou eql ur II€Jur€.r uunlnv

'Ja^oc .{douBJ qtIA\ salrs ur luopr^3 aJolu srs3rnl€.radlual IIos Jo uorsss.rdap sIrIJ uollcalq ro] IEcrllrt l€ql ^\oloq la^alE 01 Sern?€.r3drual Iros Jo uolssardap aql dq roJ pslEsuadruoc,{IFrlr€d aJ€sn€un} oql .roJ pal€arc suoiJlpuo, aJnlsrour IIos elq€.rnoleJ ar]1 'rnrJo sro,uoqspol€losr ddEaq ueq1!|sroia.oqs palElosl d^€aq Jo s,{€p Jo .raqunu € Ja/!os.reaoqs 4q5!l raqlle s€ srrlcto dll€nsn (ll.Idv dlr€a pu? qcir?lN) urunlns ,{lleaur II€JUr€1I alE.r aurBs eql dlalErulxorclcle 1€ .re^o, r{douEc lnoqlr.{l pu€qlrr\{ salrs u! possordap eq ol sa.rn}Erldrual IIos pasnEr uumln€ e?€I ur II€J-u_t€J trAEaq puE IUJ1SISUOC S.rE3,{ Jeqlo ur papaarxa aq plnord 896I lrr ulun?nEur uol?reJur roJ alqBlrns ara[\ suolllpuoc o.rnqEJadlual lros puE o.tnlslout Irosqcrqa Sulrnp sJnoq Jo .raqurnu l€1ol oql tEql acu€qc tqBJis € ,{Iuo sr aJaql?€ql apnlruoc ol olqe(rDs"ar sr 1r

'snqJ T16T-'86I porrad eql tnoq8norql

papJorar lsaq8rq oql sEra 896I ul (IIJdV pu€ qr.rBtrAtr] ll€JurlJ u(unlnv'1. eJurE.r !8961 , JSeld^E

aloqe puB Q,96I) aa€Ja^€ r[olaq Sur.roIIoJ sr€ef oq] ur (II) pu€ (r) I sorpnls

Jo solrs 5u!ul€3 a,rnlslouri eql ur sIaAaI ernlsloul lros Jo sluaurolnsBoul aqlruo,rJ luor€dd€ sl s(IJ salrs IecrqdEr8odol roddn 0q.] uro.rJ e8€ur€rp ssEutu-I sosEaJtul lrranbasqns Jo asn€caq o8€.redB a,\oqE sE^\ II€JurtsJ Jolutr\ u?q.{4sr€ad uI 'ia^a^\oq 'pa1tadro aq pFora sslrs SuIuIEE alntsloru uI uollcaJurpa,€arcuI uosEas slql Eulrnp qorlcaJul lrrull sainleradttlal IIos aauls solrslse.ro] rl€r.r€ll paur€.rp fIooJJ Jo.{lrlrclrldacsns aql uo Jcaga lu€llodull u" a,{eqlOU plno,\\ IIeJUIET Jelur.{L a^lssecxg rood sr raErurns pu? uuln}n€ 'Surrds utIlEJulBr oEE.radE s^oq€ Flre IIEJUIEJ I€nuus aBEradE uea,ulaq uolJ€latroc oql'lcal uI srlluoru .rrqu!\\ eql 8!n.mp IIEI plnoc II€JurEir a8€rodB a^oclB qltd\srEad ul pa.uncDo qcrq,!\ IIBIUIEJ aql Jo lsolu 1€ql $.oqs 0t orn8rd ut €?€o

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C UO pssEq 'aurl q1r.{1 II€]-u!?J uI uoll€lr€A Suluracuoc sacuaraJu! 1€ql elEarpur z aiqPI Jo BlEp oqJ.

'salpnls

plag luasard aql uI lno pali.I€c aJar!\ s?uerueJns€aur arnl€rad-uo1 puE a.rnlslour IIos ueq,\{ sr€3,{ aql ul pa,\-IaceJ IIEJUIET oqJ, (t)

'sqluour ralutuns aql Suunp sol€r uorl€.Iod",la qBIq aqJ (€)'II€JIII€j pUB arnlBradluol IlOs Uaei\laq uollt€ralul parresqo oqJ, (z)

'uorltalul JoJ ladel lEcrlrrteql o^oq€ a.r€ sernle.rsctural IIos uaqa .r€ad 3ql Jo sporrad eqJ (I)

ro1 uoll€Iat ur paraplsuoc aq 1sn!u salls lseroJ qEr]e!paur€.rp dlaarJ uo uortrcaJul I€8unJ uo olllrl u! uollerrE,r IIEJUTET Jo lceJe aqJ

more lavourable moisture conditions in the spring of 1968 w€re partialty com_pensated for by the lower soil temperatures in that year. Thus, althoughOctober rainfall in 1968 was exceeded in three other years during the period1934-19?1, i t is unl ikely that there vere signincanlr ly longer periods whensoil moisture and soil temperatur€s were favoulable for the fungus m theseyears. Nov€mber r.ainfau in both 196? and 1968 was below average anct torthis month rainfall is generally stable. When beavy lainfali does occur lnNovember, it is usually in short, heavy downpours. Soil tempelature data for'November 1968 showed that when heavy r.ainfall did occur., soit temperalufesivere depressed, particulally in sites with canopy cover.

R,ainfall during the summeI of 1968_69 was beiow aver.age ancl hencethe efiect of heavy summer r.ainfall on soil temperatures could;ot, be deter-mined. It is unlikely, however', that summer rainfall would significantiydepless soil temperatutes below the cdtical level for infecticn, even on s1!eswith canopy. The average lainfall received over. December, January 516iFebruary in the nolthern jar,rah forest is very low (S4mm) and evaporartonrates during thesg months is very high. Departutes from the averagehave been low except in the year. 19bb, when Febnrar-,/ rainfall was thehigheatt evel recor.ded (Fig. B0). The low rainfal l . high evapor.aLron l .aiesand small d€partures fr.om the avelage between yeals suggest firat summerrainfall is not important in cl.eating lavourable mojsture conditions lo1. ,hefungus.

Summary(1) The soil moisture and soil temperature experinents cautect out in

196?, 1968 and 1969 were locat€d rn areas representing above average, avel,ageand below average rainfal l zones in the nor.thern jarrah forest. Thus, fesultsobtained apply to the genelal forest area.

(2) In year's $'hen above avelage rainfall occurred at Dwellingup, theexcess rainfall usually felt during the winter months when soil tempelalulesrvere be16$, the critical level for' infection. Thus, although the periods whensoil moisture aDd soil tempelature were suitable lor infection are likely tobe incleased on moisture-gaining sites in those years because ol increasedmass drainage. the susceptibit i ty of freety drained siNes is not l ikely to beaffected.

(3) The compalative stabi l i ty of autumn and spriog rainfal i at Dweit ingup,the deplession of soil temperatules by heavy rainfall. aud the above avelagerainfall in the yeat- of measurement make it unlikely that the conctusiondra$,n from these measulements \\,ould be malkediy modined by year toyeal variation in autumn and spling rainfall.

(4) The low rainfal l Ieceived during the sumlner months, the l imitedvariation in rainfall froln year to year and the high evapotation rates duringthese months make it unlikely that variation in sunlrler rainfall woutd alterresults and conclusions made lrom the study.

7. CONCLUSIONSSoil moisture and soil temperatur.e are the major envil.onmental factors

afiect ing P. cintuamomi activi ty. Direct measurements of these two tactorsover a representative range of site types in the forcsi provides an index ofrelative susc€ptibility. These studies have shown that the soil envrronmenisof lower topographicar jar'ah forest sites are very suitarlle for fungal activityand survival but uppel topographical sites have a lelatively unsirtable soil

60

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Jo I€ArAJns pug uorlcaJul JoJ qloq luouruoJr ua

SECTION IIIDIR,ECT DETER,MINATIONS OF PATHOGENICITY AND SUR,VIVALMeasurement of the environmental factors which influence p. c\nnatlomi

activity and suryival provides an index of susceptibility of different sites. Theetrect of differing site conditions can only be positively conirmed, however.by direct studies. Unfortunately, the fungus and the disease situation makethese studies dimcult to implement and long time periods are required beforeresults are meaningful.

The results of four preliminar.y studies aimed at determining the influenceof environmental iactors on the iuneus are summadsed in this seclron.

1. TIIE EFFECT OF SITE, SEASON AND INOCULUM POTENTIALON THE ASILITY OF P. CINNAMOMI TO ESTABLISH INFEC,IIONSA major large-scale field inoculation trial was established at the Scarp

Road study area (see Section 2) in August 196?, Three metre by three metleplots with one mature B. grdnd,is located in each were established on Strata 1and 2 and Strata 3 and 4.

Thlee inoculum levels t0, 2 infected tupin (Lupinus orLgustiJolizs L.) rootsand 6 infected lupin rootsl and two dispersal rates t3 spots per plot and 9spots per plot) were useal to provicle a range oJ inoculum intensiiy. Inocutationof both site types was carried out in each season of the year. The experimentwas designed as a 3 x 2 x 4 factorial.

Assessments were carried out at 6_monthly intervals following inoculation_In Februar.y 1969, 28 of the highest inoculum Ievel plots from each site thathad been inoculated in October 196? were sampled for the presence ot thefungus using the lupin baiting technique (Chee and Newhook, 1966). A singleinoculation point \ras withdrawn from each plot and each sample was baitedtwice. Positive recoveties were obtained from only 4 of the samples. InOctober 1969, two inoculat ion points in each of b0 plots were sampled andbaited twice. No posit ive recoveries were obtained.

Dudng autumn 19?0, dieback symptoms appeared in the experimental area.The experiment was assessed in July 19?1 and mortalities were recordecl ininoculated plots, control plots and areas adjacent to the plots. There wasevidence that there had been invasions of the disease from outsicie, eventhough at the time of establishment of the experiment the diseas:d area was30-50 metres dos'nhill from the expedmental area.

Although the or'iginal objectives of thls expedment were not achievedbecause of contamination, it has been reported here in summary lorm toillustrate the difficulties of obtaining, from field tliats, informatron on thefactols affecting disease sevedty. The pdncipal difficulty in trials of this typeis to obtain an index of fungal activity. A sedes of experiments was caui€dout in an attempt to quantify the amount of fungus in the soil, using a vanetyof techniques, but these were unsatisfactory_ Even simple detection techniquesaimed only at determining the presence or absence of the fungus iailed,although large numbers of sampleslvere taken. Thus, assessment of fungalactivity is dependent on the recognition of disease symptoms in the hostspecies. In August 19?1, no symptoms could be detected in the jauah over_slorey, and understorey deaths did not occur until two and a half years alterinoclration. In this period of time it is practically impossible to prevent

t9

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'suorsol Jo acuosq€ ro acuaso,rd aq1 Jo ssalpJ€8ar pel€Id aje.r[ suralnrl .palrEqe.r aJa.u lsal

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'slerrl plog elnlnJuBId o+ posn €uroq sr I€trq srql uroJJ peurElclo uorl€ruroJur aqJ, .,{lr[I(lrlaleCsnsaullu,re.lep ol pasn aq lsnul,{Ial€urr1F uorl"lncow plau ?Bqt snor^qo sr 1r 'po^Io^ur sarllncgrp oqt al]dsac .IBu1 a€tel B ur slolal uea^\laq uoll€ultu€Xuoc

Resull,s

The percentage recoveries from each site in eacll month and the per_centage recovery irom the test samples, sampled prior to inocutation, areFresented in Table B.

TABLtr 3Perccnl,agc recorcr-r of P. .inn&nxni I: ,11\ infi:clert soii samplcs o-xtrose(L h the fiet{l tor norrthlv

iJtfttum 1.1 Nov.LO

l , Dec .

6l 'cb. a0 I r 'eL.

6 Mar. 20 Mar.

Exposnre pe od

:3 Nov. 12 De.. 9 Jan. 23 Jan.ro to to to

26 Dec. 9 Jan. 6 r,.eb. 20 leb.

1 , ' 0 i -_ l l0 0

40 2f)

Tot{l

".:13"'Lrpper Slopc :i

e0 t 8L.,wcr Slope 2 fi) 0 .10 1tl)

1 r ) t 00 80

20 ,to rio 100 100

r'Jl) 90 ,18

.; ;; 60

rot -t -

\:rllciv Botto]n I , 0

(]ontrol 90

In Stratum 4, apart from one sample in orle of the sampling periods(January 23 to February 20), ar sampres gave negative ,.u"ou"ii"" ou"'' trr"pedod November 14 to March 6. Simitarly, in Stratum e r_ro "umpte. earepositive recovelies over the periods December So to March el "ll,ne

funeuswas recovered consistenuy ftom Strata 2 and 1 thloughout the summermonths except for one period (November 12 to Januat./ tl wherr tne funguswas not recovered from Stratum 2.There is an obvious correlation between the soil moisture tlends in eachstlatum and the recovery rate (see Study 1 (i) and f tiil L siratun z is aLransitional zone betw€en the ireety drained and moisture_gaini; srtes analthere is considerable variation in the soil moisture levels in thrs stratum.This accounts for the discrepancies in the ."futio""nip- fr"ir"""n the soilmolsture values and the observed high recoyeries from this ptot. iwenty-nvemillimetres of rain felt durins the final sampting p""i"J Cr"i.-";"v 20 io March20) and this was apparan y sumclen! to cause soil moisture levels on Strata3 and 4 to be raised above the lethal tevel.

DiscussionThe lesults of this experiment indicate thal sulvival of p. cintuamomi lnsoil on freely dlained jarrah forest site3 during the aummer months isrestrrcted by low soil moisture levels. Soil moisture levels on the IowertcFographical sites do not reach lethal le.i,els dudng tt e summer.Soil temperature ii not a limiting tactor. sJt t"-o"rluio."", urano*nleaching comparatively high levets (e.g. 30'C) are "ut:""t io- wlo" oi. ,"rrutfluctuation_ Hine et al (1964) have shown that continuousiy ii*f, "or, t"rno"ru_ture levels are required before the fungus is inactivated.The fungus is able to sxrvive the summer months on freely dlainedjarrah forest sites that have estabtished infections, "irr"" ir]f"*iiir. o., .,_,"n

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t-

Watering*(a) . Lou treatment: During the first six weeks following inocula_tron, l50ml of water were added to the pot saucers *fr""rviiiirrg symplornswere observed in the pra.nts. After six weeks the rrigh moisture treatmentwas adopted.(b) High treattuent: The pots were maintained between fie1d capacityand saturation by watering {rom above and belovr in alternalrve z4_hourperiods.

Procealure

Ten seedlings of Bo,nksia grand,is wete planted in each pot (22.5 cmdiameter) in May 196?. The pots were inoculated in November 1SOZ Uy eitnermixing the sand mycelium in the sudace 2.bcm of "oit- orj t v-ir_r"""rirrg tfr"lupin. roots randomly to a depth of 2.5cm. pots were_ mainrarneal in ashadehouse and assessed for modalities t$dce weekly. soli t"rrio""ur,r"". ,"r"recorded during the experiment.The experiment \ras termjnated on March 28, 1968. Whole plant dryleight, shoot dry weight and seedling height were _"u.r.r."d.

-iiu" soil cores(1.2cm diameter) wele taken from each pot and sampled for the presenceoI P. cinnamomi by the method of Chee and N"*f]oof. f fgoOti.

Results

Dr! WeiSht-Difrerences in height and dr,y weight were not signincant.Measurement was complicated by lne presence of dead plants irl some pots.MortalitA-Thrce weeks aftel inoculation, significan y more deatns occurrealin the pots with high moisture tevels, inocutaled ,ttf, t* l"pii"i""rs. Therewas no significant difference betw€en soil types.

. _ - Six weeks after tleatment signiflcan y more deaths occurred in pots wiihhigh moisture levels inoculated with ten lupin """t" tnun-_ tn" l_rigt_r moisturetreatment inoculated with two lupin roots. ffri" tr""i_""i

-i" tur_n haatsignificantly more deaths than the remaining treatments. DifferencesDetween soil types again were not significant.

Twelve weeks after inoculation the interaction between inoculum level,soil Jype and moisture regime was signiflcant. tfr"." *"." "ig;i"anily mor.edeaths in the hish moisture level. pors, i"o"urutJ-rl,iirr - i";r'"i;;;, .oo.", ,oall

.Jour soils and in the high water level lateritic soit, inoculatect with twolupin roots, than in an oth€r. treaiments (Table 5).

Sai.l .Temperdture-Soil temperatures in pots during the trial were consrsier ywithin the range favourable for zcospore production (15. .uo 32.C)

Recaaer! oJ P. cinndlnomi_At threcovered r'om any "r th. ;;;h'ofi"iTi:"Jj"lT,$i,ll";T"-T#:i#:Positive recoveries wele obtained Jrom

^sO per "J ", in"'l""ri inoculatedpots in the wet tleatment and from five of the Ary tr""t^i"i pots eventhough these tatter contained no mortalities. At the time the experimentwas sampled, it vas believed that unlesioned r"pi" -oil--i",,# the baitingprocess wele uninfected and did not_ war.rant plating on agar. f]his assu-ptionhas since been prov€d incouect and ir un r"pi" ,oit"'iriii."rr.oiu*o, """ou_elies could have been higher.

I)iscussion

"..^^::." ttl"* oJ the sand mycelium inocutum could have been due toexcesslve maceration and ol dilution

.of trre mvcetiri,n,'litf, subsequentattack of viable macerations by other miclo_orgu,l iaI lra. -___' "-"" ,

66

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Measurement of environmeltal factors in relation to known limits forfungal pathogenicity and survival provides some index of site susceptibity. Anacculate assessment of disease susceptibility, however, can only be made byobservations of fungal activity under specific environmental situations in thefield. Direct inoculations ol specific sites, followed by observations of diseasesymptoms, would appear to be the best method of assessing site susceptibility.There are, however, considerable ptactical diffculties in establishing, main_taining and assessing these types of experiments. M3asurement of ,,natural,,rate of spread provides an alternative method of assessing the influence ofdifierent environmental factors on fungal activity.

A series of plots was establ ished in 196? and 1968 to measure the rateof spread of the disease upslope from existing infections.

Procealure

Plots in which the position and condition of understorey and oversloreyspecies in respect to a 66-110 metre baseline located at the boundary between"healthy" and dying forest, were establish€d on 18 difierent locations in iheforest in 196? and 1968. Changes in the health status of the undersioley anatovelstorey species in advance of the baseline were recorded annually. Noairempt was made to observe changes in the shrub layer because of thedimculty of discriminating between deaths caused by natural seasonal mortali_ties and those resulting from fungal activity.

Soil, slope, aspect and position in the landform were recorded for eachplot. one of two plots located adjacent to one another was treated to reducethe density oI Banksi(r grcLndis to determine if this would reduce disease spreact.AII of the plots were iocated in positions above the moisture-gaining sites.

The annual increase in area of Jorest infected at each plot was measuredby assuming that a line joining the most r-ecent understorey deaths to theneafest d€ath of the previous year defined the area of newly infected soil_The average increase in area, of hfected soil iinfected area divided by lengthof plot) and the maximum linear extension for each plot wele recorded eachyeat.

Results

The annual spread of the disgase, based on understorey symproms, overfive yeals and 18 plots is shown in Table 6.

The annual extensions of the disease over a five year period in two plotsthat have contrasting rates of spread, are shown in Figures 81 and 32. Theaverage annual disease spread over a1I plots is shown in Figure 38. The wallof dead B. grarrdfu, which occurs aJter extension of the disease, is shown in_t' tsure ;t4

Plot 15 (Table 6) was the only plot in which a significant change in ihecondition of the jarrah overstorey that could be attributed to p. cinn.otuorniwas detected. Three jarrah over-storey trees died and three exhibited severecrown detelioration in ihe five year period.

Discussion

There is little objective data available on the rate of spread of disease fromexisting infections. Air photo interpretation techniques were useai to obtainan average percentage increase in the area of infected forests of four per cent

68

69

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YEAN OF i,IEASUREMENT

Mean annuat unear spreatl .t u,"or"oitfl"""L'"t"" for ls plots over fte pedo.r 1e6?_68

The ayerage spread of the disease onto the upper topographical sites,however, is not insigrriflcant to management if the larye perimeter ot diseasedareas is considered. Relatively la,rge extensions of the disease at isolated pointsalong the plots indicates that under optimum conditions tne exienston ot tnedisease can be rapid. This type of extension is likely to cause more rapialdisease spread because on sites which slope in two directions it provides theopportunity for passive dovinhill spread.

In one of the plots (Fig. 32), the average spread wa6 extenslye and muchgreater than in any oi the 1? other plots. Itrere rras no appareni sit€ orvegetation characteristic which could explain the rapid extension of the disea.sein this plot. Curren y, attempts are being made to determlne if the chemicalor microbiological characteristics of the soil on this site are difierent fromlhose oI the other sites,

72

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the probability of contamination is high. The results of neld saprophytic su.r-vival trials and measurement of the rate of disease spread on to upland slte$supports the conclusion, formed on the basis of direct mea,surements of thesoil moisture ar]d temperature regimes of these sites, that they are relativelyunfavourable for pathogenicity and saprophytic survival, 'fhere is, however.some eyidence that factars other than soil moisture and 6oil temDera,tureafiect the spread of the disease on some upland sites. preliminarT triats in-dicate soil type does not affect fungal pathogenicity. except by its efiect onsoil moisture regime.

74

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SSVSSIO UOd fVIINSIO'I Cl\Iv INITINOUIANII flHIAI NOIJCflS

(10) The overall dDerdge tate of unaided spread upslope from existing

infections is low. On some sites, however, average and maximum

extension Iates are high. Linear extensions of the disease of 20 to30 metles per year, 4, poizrs along the perimeter of diseased areas,

are common. 'Ihis type of spread suggests that the principal

mode of unaided spread is bv way of verv susceptible host root

svstems.

Measurements of soil moisture and temperature regimes of a range ofsites within the forest have demonstrated ihat there are two major zones that

have markedly different susceptibilities to the dlsease. Lower topographicalsites have long periods during which P. cinnd.momi infection can take place,

but on upper topographical sites the environment for both infection and

sulvival is only malginally suitable. The broad-scale pattern of distributionof the disease. that is its predominance on lower topographical sites (Fic 2),

is explained by this study. The fungus, however, can survive and causemortalities on upper topographical sites to a degree which i.s difficult to explain

by their calculated susceptibility. Hence it is necessary to consider ihe effectof othel factors of the environment to explain the occuuence and activity

of the disease on upland, freely drained sltes.

1. THE IMPOR,TANCE OF OTHER, ENVIR,ONMENTAL FACTORS

The Effeat of Host Susceptibility

The degree of host susceptibility appears to be a major factor influencingthe speed and intensification of the disease on sites {hich have environmentsthat are only marginally suitable for P. crnnqnxomi activity. The large numbels

of highly susceptible species in the understorey and shrub layer of the forestprovide a dense mat of susceptible roots $hich act as channels for fungalspleaal. The presence of a dense mat ol susceptible roots possibly reduces iheperiods during which the Jungus $ould have to survive and sporulate in the

soil- The obseNed pattern ol spread of the disease and the direct measure-ments of the soil environment external to the host l'oois, which indicated

that it was only nargiDally suitable for fungal activity and survrval, supportthe hypothesis that the existence of a highly susceptible species in the under-storey and shrub layer of thg folest is a maior factor permitting the disease

to spread and cause mortality on upland jarrah Jorest sites.

Although a numbel of the shlub and understoley species of the folest

ar'e highly susceptlbte and hal'e i-ooi systems which can be completelyinvaded by the fungus, there is circumstantial evidence that the commercial

species, jarlah, does not have the same degree of susceptibilitv The ability oljar lah t lees to survive fol long periods after the lnderstorey and shlubIayer species have died indicates that mortalitv results flom repeated infec-tion of the flne loots. rathel than invasion of the total root system

Artificial SpreadPriot to the discovery of the causal agent responsible for jauah die-

back, the fungus *,as spread artificially during normal folest operationsFor example, road makrng material $'as frequently obtained from jarrah

dieback areas. These plactices, particularly when they cause infections onupper topoglaphical sites, resulted in large extensions of the disease. Massive

artificial spread of the disease has undoubtedlv been a major factor contd-buting to its spread and intensification on upland sites

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magnitude of the disease situation and its potential for more serious develop-ment make it necessary to make intedm recommendations to managementthat, in part, result from logical extrapolation of the preliminary investiga-tions lather than from direct expelimental evidence.

Recognitrion of Variation in Site Susceptibility

This study has demonstrated that ihere are t$/o markedly difierentsusceptibility zones in the forest. Lower topographical sites have conditionsthat are suita.ble for flrngal pathoeenicity and survival for long periods du ngthe year. Upland sites are only marginally suitable for the fungus. Man-agement aDd research approaches to disease control will difier according tothe susceptible zone.

Thele is no possibility that environmental modification of lower topo-graphical sites will significantly affect their susceptibility. Thus the onlymethod of restoring ihese sites to productivity is by planting resj.stant species-Fortunately, these sites do not support high quality jarrah stands and iheyare \rel1 suiied for the growth of exotic species.

The relatively unfavourable conditions Jor Jungal activity on upland sites,the existence of high value stands and the diffculties of replacing ja$ah withan alternative. productive crop justify research and management approachesprimarily aimed at protecting the existing crop from the disease rather thanreplacing it.

Plevention of Disease Spreaal by Artificial Ino(rulation

The data fr'om rate of spread studies of unaided disease spread indicateLhat, if "man-carried" disease extension can be eliminated, the progression ofthe disease into healthy stands $'il1 be greatly leduced. Information on thejarr.ah forest environment. and the envir-onmental factors influencing the fun-gus, form a basis for ' simple hygiene ploposals. which should have a large im-pact on the spread and intensiflcation of the disease. In brief, restriction ofmovement of vehicles between infected and healthy forest, washing of vehi-cles to rernove infected soil and the lestriction of logging activities in hiehvalue folest ar_eas to the surnmel months $'ill greatly reduce the plobability

that new infections rn' i l l be introduced.

Manipulation of the Forest Canopy to Reduce Site Susceptibility

Measurements of the soil moi'ture and temperature regimes of uplandjarrah sites, under condltions of maximum canopy and litter ccver, indicatethat unde-!- these conditions fungal pathogenicity should be reduced to insig-nificant levels. The ploduction of a r.miform even-aged forest, as opposedto a "selection type" uneven-aged forest, would improve canopy cover andhence reduce soil temperature during the critical spring months. The funguscan cause death of jal lah glowing in even-aged stands, so i t is evident that,even though disease severity wiII be reduced, the conversion io riniform stands$ri l l not el iminate the disease. The pool shading characterist ics of jarrah

clowns make it unlikely that the deglee of canopy cover necessaly to eliminatepericds of coincidence of molsture and temperature conditions, suitable foriuneal inf€ction, will be achieved by silviculturai treatment of the jarrah

overstorey. Heavy litter will reduce disease sevediy by reducing soil tem-peratures in spdng but it is unlikely that conditions {'hich conpletelu prevent

fungal activity will be achieved and the maintenance of large unbulnt areasin the forest is implactical. The degree of canopy cover estimated necessary

?8

6l

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Reduction of Density of Highly Susceptible Host Slt€ciesThe structule and composition of the jarrah forest prior to exploitation

was ver-y diflerent to that of the existing forest. Reports from early lorestsurveyors and examination of remnant virgin stands confirm that the under-storey was sparse and the forest was "park like,'. This contrasts markedlywith the current, structule of a large area of the forest (Fig.36), which hasa dense understorey component comprising principally B. grandis. The causeof the change in stlucture of the forest is unknown. Reduction in competitionfrom ihe overstorey, disturbance and ihe absence of high intensity flres,however, are probably three major factors which have favoured the develop-ment of the B. grandis under--torey. The lapid symptom development in thisunderstorey species foilowing introduction of the fungus confirms that it ishighly susceptible to the fungus. Its r.emoval would reduce disease sprea.d.

In the short term, poisoning $ith herbicides in small areas could removethe undersiorey. The cost of this treatment over a large area, and thepossibility that re-treatment, of poisoned areas would be necessaty, ma.ke thismethod impractical on a Ialge scale. There is evidence that hich intensityfires cause marked changes in the compo ition and structure of the under-storey and shrub layer. Further research is necessary to determine theprescription of burning intensity and frequency which would bring about thedesired changes without damaging the jarrah overstorey.

Eealthy foresi shovlDg typlcal irregular stald structure, which rcsults Jrom ..groupsetection manaeemeDt.

BO

I8

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F

ACKNOWLEDGEMENTS

The study was sarried out under the direction oJ Dr' E R" Hopkins Mr'

P. c. Kimber, Mr. J. J Havel and Mr. A. L. Clifton provided advice and

assistance at vadous stages of the study

The stafr of the Forest Resealch Institute, Kelmscott' in particular Mr' D'

Darling, provided assistance \tith the preparation of inoculum and media plates'

Mr. M. J. Dilion and Mr. R. J Kitt provided technical assistance in the

fleld and laboratory.

Miss L. R. Haves provided invaluable technical assistance and ivped the

manuscrlpr.

LITERATURE CITED

Anderson, E. J. (1951) Tl1e P cinnamomi problem in pineapple f ields ofr{awaii. PhytopathologJ. 41, 1-2

Batini. F. and Podger, F. D. t1968). Shelterbelt mortal i t ies on the SwanCoastal Plain Aust For. Res. 3. 39-45

Campbeil , W. A. and Copeland. O. L. Jr ' (1954). Lit t teleaf ' l isease of short--- ' - tai i ,na loblol lv pines US. Dept. Agl ic. circ No 940'

Chee. K. IL and Newhook. F. J (1965). Improved methods Jor use ir-r studies-----ii-biitointnora cintualnami Rands and other Phvtophihola species. N'z'

J. Agric. Res. 8, BB-95.Chee. K. I{ . and Newhook, F. J (1965) Variabi l i ty la Phutophthol '(r ci l tna'

t om i Rands . N .Z . J . Ag r i c . Res 8 96 -103 .

chee. K. H. and Newhook, F J. (1966). Relat ionship of microorganisms to'^-' "'""i"futio" of Phutophthora cinnanlonLi Rands N'2. J Agric, F'es 9,32-8.

CoDeland, o. L. Jr. and McAlpine, R, G (1955). The inteuelai ions of l i t t le--"- ' f* i i , " i t" index, soi l and ground cover in Piedmont short leaf pine stands

i in's. carol ina and Georgia). Ecolocv 36, 635-640

cardner. W. R. (1960). Dvnamic aspects of watel a''/ailebility to plants

S o i l S c i . 8 9 , 6 3 - 7 3car'dner. W. R. (1968) Availabilitv and measurement of soil water' 1?' Water--

ii".n"it" and Plant Growth. Vol. I Ed Kozlowski'

Hatch. A. B. (1964). The interaction between forest f loor dvnamics and soi l--*-ir"b"""ti"" ir' Eucaluptus marginatd Sm forests. M Sc Tllesis, University

bt Syaney.Hart isan. C. (1964). Some observations on the effect of Phl l tapl l thora iD

Monierey Pine. I 'or. & Timb svdney, Aug, 5-6'

HeDbins. G. H. (1964r. Cl imate and forest diseases. In Annual R'evlew ofPh.ytopathology. Vol. 1

Hine. F,. 8.. Alaban, C. and Klemmer. H. I1964t lniuence of-soi l tempela-i,rrre on root and heart rot of pineapple caused Dy fnlfopt"ttrol 't cxtunQ-iomi and PhAtophthora paro siticq Phytopaihology. 54, 1287-1289 '

Jacobs. M. R. (1955). Growth habiis of the Eucalypts For' & Timb Bur'commonwealth Government Printer' Canberra

Kimber, P. C. (19?4). The root system of ja'rtajf--(Euc' uplus. mdrginatL) '--- neiiarcn Patter No 10. Forests Department of western Australia

Kuhlman, E. G. (1964). Survival and pathogeniciiy ot, -PlrEtopll'thor'L cinnd'-o-i i tr several western orcgon soi ls. For- Sci 10, 151-l5B Publishedalso as Tech. Pap. Ore Agrlc Exp Sta. No. 3839

Leeper, G. W. (1960). Climates. ln The Australian Envilonment, ihirdedit ion (revised).C.S.I.R.O. in association \\'ith Melbourne Universitv Press.

Marx. D. H. and Davey, C. B. r196?). Ectoirophic mvconhizae as deterrentsto pathogenic rooL inlect ions Nature. Lond. 213. 1139

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