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s ugt.IVHf Table I TAXONOMIC DETAILS FOR INSECT SPECIESIN THE ORDER MENTIONED IN THIS PAPER

VenucularName Letin Binomial Order Family

Jarrahleafminer P erthida glyphopa Lepidoptcra Incurvariidae Gumleafskelctonlzer UrTba lugens Lepidoptera Noctuidae Bullseye borer Tryp hocaria acant ho c er a Coleoptera Cerambycidae Diade gtn sp- Hymenoptera Ichneumonidae Chrysonotomyia sp. C Hymenoptcra Eulophidae

Lcafblistersawfly P h! Iac teo p hagd Jr ogqqtti Hymenoptera Pcrgidae Autumngum molh Mnesampela privat.t Lepidoptera Geomet dae 'Cupmoth' D orattfera quadri guxat a Lepidoptera Limacodidae Lipdretrus spp. Coleoptera Scanbaeidae Chrysophtharta amoena Coleoptera Chrysomelidae P aropsisterna elliptica Coleoptera Chrysomelidae 'Spitfires' Perga spp. Hymenoptera Pergidae

Theextent of infestedforest was mapped by theForests in 20 standswithin a 50-km radiusof Manjimup.This Depafimentin 1964-196'7and in 1983,and by CSIROitt researchwas extendedin November1987 to 20 stands 1968-1976 and 1980.These maps, drawn to a common within a 4O-kmradius of Collie.For eachstand, damage scale.zre included in Abbott(1987). causedto leavesby leafminerisexpressedas apcrcentage of fully expandedleaf area.Multiple regressionanalyses Theinvolvement of theForestsDepartment in leafmincr wereused to conelatethis variablewilh environmental researchhas until recentlybeen minimal. Van Didden factorssuch as stand basal srea, years since previous fire, (1967)showed that an experimentalaerial sprayingof years since previouslogging, averageannual rainfall, jarrahstands was ineffective and uneconomic. Since 1984 percentagoforest coycr and soil texture, researchhas concentratedon measuringdamagc levels nearManjimupand Collie,relating these to environmcntal During 1983-1987J. Hatt of CALM estimatedthe faclors,and quantifyingmedium term impacton crown extentof browningcaused by leafminerin eachjarrah conditionand wood increment (Abbott unpublished). crownin 15plots north-east of Manjimup.Some of these treesexpcrienced crown scorchdudng autumnburning Informationon silvicultumlcontrol andthecause(s) of duringthe studyperiod. thcoutbreak is lesscomprehensivc ard morespeculativc. Theapproach used in theCSIRO study involved sampling Life history (Table2) theabundance of larvaealter they had ceased feeding, and expressingthis variablein relationto a defined ea of Mothsemerge and fly duringthe day. They ire weakfliers. jarrah canopy.The abundanceof mothsemerging fiom At night or duriogcold wet daysthey seek shelter in the soilhas also been measurcd relative to adefinedarea ofthe litter.Almostall eggs are laidon the lower surface ofj arrah forestfloor. Studiesin fhesame stands over seYeral ye s leaves(Wallace 1970), at 14-220Cin diminishinglight or havebeen uscd to assessthe impactof prescribedlow overcastconditions (Mazanec 1986). The longevityof intensityfires, logging, cleariDg, and annual variation in mothsaveruges 10 d:lys in the laboratory. rainfall on relative abundanceof final irstars and/or moths. The crterpillarstage (within the lea0 lasts160 days. After feedingfinishes (instar4), the caterpillar faUs to the The approachused in the CALM researchinvolved groundwithin a case,generally around midnight, to avoid markingnewly developing leaves each November (or as predationby ants(Mazanec 1980). It thenburrows 1-2 cm soonas the petiole is longenough to holdatag) since 1984 6L

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urooileN into the soil, whereit remainsin diapausefor about150 Crown scorchduring autumnfire causeda notable days(depending on latitude),until the endof February, decreasein leafminerinfestation (percentage darnage) the whenpupation begins. Theprepupal stage lasts 2 daysand followingspring (Table3). However,by thenext spring, thepupal stage another 35 days. damagelevels could not be distinguishedfrom adjacent fieeswith unscorchedcrowns. Most eggsarc laid in leaves6-months-old (Wallace 1970iM^z rcc 1986;Abbott unpublished)but thereis Drought conflictingeyidence whethcr females prefer crownsof treesor foliage of groundcoppice. Newman and Clark (particularly (1925)noted heayiest infestation nearer the groundalrd Drought duringwinter) results in decreased production thiswas confirmed by Wallace(1970), with theexception of new leavesin the following spring and (Mazanec Thenumber laid of one standlacking abundantadvance growth. There summer 1980,1981). of eggs infestationin thecrowns12 m aboyeground was extremely in fhefollowing autumn is thereforelow,and so the leaves heavy.Mazanec and Justin(1986) found a statistically availableshould receive more oviposition,resulting in significantdif ferencebetween infestation in uppercanopy more damageand thereforeobvious browning of the and lower canopy.CALM studiesin 12 standsnear canopy. Manjimuphave not detected any significant difference in infestation12 m aboveground comparedwith 1-2 m above Loggingand regeneration ground(Abbott unpublished). Nevertheless, where female mothshavea choice,lower levels are used for oviposition Loggingfollowed by regenerationor thinningof siands first (Mazanecr,personal communicalion). results in rejuvenationof remaining foliage, perhaps respondingto increasedavailability of soil moisture Fire (Mazanec1980, 1981); this favours oyiposition. Adyance growth is releasedand continuesto grow and proyide plentiful foliage suitablefor oviposition.Detailed hescribedlowintensityfires in spring,depending on their bchaviouralstudies (Mazanec and Justin 1986) indicate intensity,may kill mostof the larvaediapausing in soil. thatlogging by itselfshould disfavour oviposition, because But adults(moths) can disperse at least1 km andprobably gravid femalesrequire subdued light and cool weather farther passively.The result is that bumt standsare (<20"C).The abundanccof new lcaves(i.e. thosewith recolonizedby moths the following autumn(Mazanec suitableoviposition sites) may be thetrue limiting factor. 1980,1981). For up to 2 yearsfollowing this type of fire, (1970), manymore eggs are depositedthan in adjacentunbumt Wr.llace usingbroad-scale surveys, concluded stands. thatsevere infestations were usually associated with open country,or with naturalor artificial cleaings. He then Springburning rejuvenates the forest crown, resulting suggestedthat loggingand othersilvicultural practices in productionof moreleaves on which femalespret-er to whichtend io openup the forcst may require modification. ovipositthefollowing autumn. These females then have a Mazrnec(1980) examined this hypolhesis moreclosely by widerchoice ofleaves. Damage to individualleaves may measulingstand basal area at42 places within the leafminer not attainthe 40 per centthreshold which the human eye outbreakzone and at 38 placesin non-outbreakareas in perceivesas brownish tinge in theforest canopy (Abbott, November1978. The averageswere 30 and 38 m'ha'' personalobservation). The presenceof abundantnew respectively,astntistically significantdifference. However, foliagethus dilutes the effectof leafminerinfestation. recentresearch by CALM nearManjimup has shown that neither stand basal arer nor time since logging are Regtessionanalyses ofdamage to tagged leaves (ground significantpredictorsof Ieafdamage by leafminer, but that coppice)at 40locationsin thecentral and southernjarah rainfall zoneis significant.These discrepancies can be forest found 26 per cent of variation in damagewas reconciledbecause the dilferencebetween outbreak and yery accountedfor by periodsince fire (negative)and 14 per non-outbreatareasin stand basal area likely describes cent by averageannual rainfall (alsonegative). Perhaps the differencebetween stands experiencing low or high thelatter correlation simply reflects the concentrationof aycrageannual rainfall. Recent surveys ofsiands recently ovipositionsites on few new leavesin siandsin the low loggedand adjacent slands either never logged or logged rainfall zone.Surveys of standsrecently bumt in spring up to 100 yearsago indicateno significantincrease in andadjacentstands notburnt forup to 11yean indicate no leafminerdensity in theformer (Abbott, unpublished). significantincrease of leafminerdensity in the former (Abbott,unpublished).

' Z. Mazanec,CSIRO Division of Entomology,Floreat W.A. I8

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SI}ISNI JStUOd JS:Id AIITIHI dO SII'ICAJ AdI'I z alqBJ Table 3 EFFECT OF CROWN SCORCHDURING AUTIJMN FIRES ON SUBSEQUENTINFESTATION OF LEAFMINER IN JARRAH CROWNS.

Plot No. Recentfire Voof janah crownsinfested' history No nees with leafminerin Spring 1983 1984 t98s 1986 1987

501 Spring1980 15 1,8.7 21.0 16.7 35.0 r7.3 505 Autumnl983 48 22.7 27.5 38.8 68.1 40.6 na ,1 506 Autumn 1985b JI 32.9 2X.3 26.3 27.6 557 Spring1984 l7 86.8 65.3 43.5 70.2 5t.L 558 Spring1984 l5 95.0 76.7 65.7 62.3 60.8 559 Autumn 1986 16 93.8 92.5 69.4 JJ.I 82.7 not scorched 69.2 87.s scorched 2.5 66.3 560 Spring1984 20 92.5 59.8 43.0 50.3 59.0 561 Autumn 1986 25 88.0 &.6 51.6 6.0 LI.6 notscorched 15.5 14.0 scorched 0.0 11.1 562 Spring1981 27 55.0 52.8 57.2 42.4 42.8 5& Spring1984 12 87.5 70.0 64.6 70.8 68.8 565 Autumn 1986 JO 76.7 74.6 t 3.) 7t.0 9.2 not scorched 75.2 68.8 scorched 50.0 40.8 566 Autumn 1986b 27 94.4 60.4 60.7 54.8 6r.9 567 Spring1984 13 96.2 47.3 38.5 55.4 61.2 618 Spring1983 t4 94.6 43.6 63.9 66.8 66.1

al least20 per cent ofcrown brown, causedby leafminer

no crowns were scorched by this firc

Outbreaks recorded around farms near Lake Muir in the 1870s (Mazanec,personal Yet Theoccunence of outbreaksof leafminermay be modelled communication). thesurrounding jarrahforest remained free 1960s. in three ways (Fig. 2; Ridsdill-Smith'? personal of outbreaksuntil the communication).Model l may apply to standsnear Severeleafminer infestation itself promotesfurther Manjimup (infested for more than 20 years) whereas infestation,because severely damaged leaves are shed in model3 ceriainlywould applyto Collie region(infested October instead of the normal January-Februaryperiod for 5-10years). Model 2 is not supportedby CSIROdata. (Mazanec1980). Thispremaiureabscission contributes to Reasonsfor a non-outbreakstand being transformed an abundanceof new leavessuitable for ovinositionthe into an outbreak stand are not clear. Single factor followingautumn. explanationsare not supporiedby the history ofhuman use and managementof the forest.Thus, flue was usedby Site Aborigines; many areasnear Collie and Manjimup were cut-overin the1920s, yet these were free of outbreaksuntil It is commonlyobserved that sandy sites in the soutiern recently.The outbreaks beginning eastof Manjimup were jarrah forest consistentlyhave browner canopiesthan uplandsites. Sandy sites may resultin more stressthan ' J. Ridsdill-Smith,CSIRO Dvision of Entorilology,Floreat W.A.

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Proportion of jarrah with Sping Grand '85 1983 '8i Mean

0 % leafminer infestation " 0 0 0.003 0.13 0.003 0.001 <10 7oleafminer infestation 0.09 0.03 0.05 0.19 o.2l 0.11 <20 7oleafminer infestation 0.15 0.16 0.15 0.30 0.32 0.22 <30 7oleafminer infestation 0.n 0.33 0.29 0.44 0.44 U.JJ 940 7oleafminer infestation 0.27 0.47 o.43 0.53 0.53 0.45

Total No. tsees 323 itJ 375 375 374

' Recognized as bmtning ofjarah crowns,

Table 5 Natural enemies SPATIAL VARIATION IN PROPORTIONOF JARRAH RESISTANT'IN SPRING1985 TO Mazanec(1987) recorded 10 speciesof Hymenopteraas parasiioids immature leafminer.Parasitoids LEAFMINER INFESTATIONAssuming S20 per cent ofthe stagesof weremore common in oneplot within the northemjarrah infestationof eachcrown by leafmineras the criterion 'resistance' forest(non-outbreak) compared with twoplots in southern of jarrah forest (outbreak). Relative mortality of feeding larvae was greater in non-outbreak than in outbreak Plot No. TotalNo. trees No. tees'resistanl' situations. Muchisknown (Mazanec 1988a, b, 1990a,b) aboutthe 501 16 t2 basicbiology of theseparasitoids. Most speciesfluctuate )UJ 49 o markedly in numbers from year to year. A key factor 506 J_t analysis (Mazanec'unpublished) indicates that these 557 3 parasitoidsare unlikely to prevent outbreaks. Most species generationsper year ))6 l5 0 have several and so require an host to April whenleafminer is ))v T7 0 altemative duringOctober within the soil. This is not available,so most of the 560 20 2 parasitoidscould never attain consistently high densities 561 68 4 from yearlo year. 562 28 0 564 l6 0 The two univoltine species arc Diadegna sp. arld 565 37 I Chrysonotomyiaq. C. The former is very rarc but is not specific to leafminer, whereasthe latter can be abundant, s66 29 0 but is specificto leafminer(Mazanec, personal )o/ t4 I communication).Both speciesare intemal parasibids of 618 16 1 larvae.

Total 375 Themostpromising species, C hry sonotomyd sp. C, is apparentlyadversely affected by fire. Mazanec (1988a) found it present at high densities in the Meribup fire

84 s8

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Themostrecent outbreak in thesouthern jarrah forest ma.rriprcscnt and distanceto nearestnon-regenerated beganin 1982,resulting in widespreadbrowning ofcrowns krrri forestwere also recordcd for the stand. in January1983. Some 90 000 haofjarrah forest bclwcen Nannup,Greenbushcs and Walpolewere affected.One Most bullseyeborer damagcis associatedwith yearlater the infested area covered 230 000ha.In January proximity to maturekarri stands,sites where karri is a 1985and 1986 the extent ofoutbreak was 300 000ha ard minor componentbefore regeneration, small coupe size 240 000 ha respectively.Maps showingthe maximum andsite susceptibility to drought(Abbott e/ 4/. in prcss). annualextent of the outbreak(in January)have been producedfor 1983,1984, 1985 and 1986(see Abbotl 1987).The outbreak has since declined (Abbott 1990). ECOLOGICAL IMPLICATIONS

In January 1986 the seyerityof infestationby The impactofpest inscctspecies on the ecologyofjarah skeletonizerwas quantifiedby crownometerin two stands andkarri foresthasbeenmeasured foronly afew attributes, nearYomup, based on 200random obscrvations in each namely biomass of othcr crown-dwelliDg invertebrates, stand.Crowncoverwas66 percent in theoutbrcaland 54 condition ofjarlah crowns,wood increment,and mortality per cent in the non-outbreakstand. The propo ion of ofjarrah. brown jarrah foliage was 60 per cent ard 2 per cent marrifoliagc was 7 percent respeclivcly,and thatofbrown It was fortunatethat leafminer outbreaksdeveloped in per and0 cent. W.A. decadesafter methodsoforganic chemical control becamecommerciaUy availablc in thc 1940sand had been Bullseye borer erthusiirstically adoptedin North Amcrica and Europe to control irsect oulbreaks. By thc 1960s it was widely The life history(Table 2) of thisinsect was elucidated by apprccialcdthat insecticideshave adverseervironmenfal Clark (1925) in a study donc mainly on marri near andevolutionffy consequences,including watcrpollution, Mundaring.Interestingly,although recording six cucalypt contaminationofbirds and m nmals feeding on affectcd speciesas hosts,Clark did not include karri. Voutier insccts, extensive mortality of non-hrget insects, and (unpublished)studiedaspeclsof thebiologyof thisspecies evolulionofresistcnce by pestinsects to chemicals,which in karri from 1983to 1985,and found several differences countemctstheireflicacy. Insecticideshavenotbeen used from Clark's study, prrticuhrly the presenceof'frass to reduce leafminer outbreaks. ejectionvents'along the bole, whcrc tho laryaempties out the contentsof its tunnel.Infestation was found to be Conservationvalues widespreadfhmughout the kalri forestand have the caprcity to inflict severestructural damage to youngvigorous karri It is reasonableto hypothcsizethat stands with insect- regeneration. drmagedcrowns would offer fewer feeding sites, less food andfewer suitable oviposition sites lorothcr invertebrates Borerdamage is alsoassociated with brownwood, i.e. comparedwith standsin which most of the foliage heartwood,which developsrapidly into decayedwood comprisingthe canopy is grccn. (Donnelly,unpublished), and with kino exudationfrom (Voutier, frassejection vents unpublished).Interestingly, In the southernjarrah forcst, sampleshave been (1925) wls Clark vigorouslydisputed that this borer the collected(9 samplingdates over 2 yearsx 45 hees)from maincause of kino Yeinsin marri. j:trrahboth unaffected and affected to varyingdegrees by gumleaf skeletonizercatcrpillars. These samples have In 1986R. Smith(CALM ResearchDiyision) studied beensorted to spccieslevel, and the biomassof each theassociation between internal borer damage and cxtental specieshas been estimated from a body length/oven-dry symptomson kari trees,and statisticalanalysis showed massregression based on a reprcscnlalivesample of that the associationwas significant.These data revealed species.Biomass of eachsample is expressedrelative to that infestationof kari regenerationbegan sometime oven-drymass of foliagein the sample. between10 and l4 yearsofage.In 1987, Smith surveyed regeneration,and recorded on a sample 30 standsofkafi A graph(Fig.3) showing thebiomassofall invertebrates in five indicationsof cxtemal of 50 trees each stand (exceptingskeletonizer) in relation to the biomassof status,bole damagecaused by bullseyeborer, dominance skeletonizercaterpillars in thesame sampleclearly shows length,stem diameter and bark thickness at breastheight. that thereis no significantrelationship. Similar graphs years the last firc, coupesize, site type, Standage, since werepreparedfor Coleoptcra,Araneae, Hemiptera, other annualrainfall, standbasal area, proportion of average Lepidoptra,Hymenoptera, Diptera, other leaf chewers, L8

peqsllqBlse 'uorlsrouelep lEuosred)sauBzBI,{.{q slolduo 8 g6I urposscsss srqlJo lualxeeql ,roqs s€,4{solod qeuef po33e13o su,no-ro cql;o suoqrpuoaeqJ lsoq(t'Erd) su.ror3qeuef palcclas;o sqd?J8oloqd.srue,( .suoDqseJur 0l-g ulqll,r\fprdu perueddeuo{eroueleq '(r86I .,y\srlspurg).(1qenb ro uorlrsod plo Jo &ue^os oql psqJ?eJlou p?q [orpJouelepu^rorc 'ezrs 'un8aq 1o sselpre?arpoulqar eJPse34 luqsrsor Ip l?ql d ua$r .(1uopeq uorleqo;apereq^r 1eql pelou srspuqs q€lJufJo Suruurql SurllIe3uo3 dc11od 11193 f,eqJ 'uot1u1saJu!raurur;eay qeueI uor3 pElnsereEeu.rep pe,terlaqpezr$lnJnf, ',roleq luou?uradlBql sJalsaJoJlsotr^I ro 0I -6 Jo en[€^B seqxepur uollI puoJ u^\olJ Jql uJqf oJJzJqptno^|uJurJJJUr (qn p) .fluo ry?q rapun reletuBrp leql et?crpu|I elq?I ur osoql qlrl{ s"alPpolseJur JsururJsel qer?f ol pourJuoJsE^1. eSueqJ peurqruosueq,,('elBp eseql'(tuoJ rcd 6I) lseolpetsJoualap elqrsr,rduu Joqleq,rpue ,{?^1alqer^ duu ur mo8r,rasrl '(luec:ed satodluBurruopqns lo su.r,ror3 SE)lseloJ ,{lrpnb pw ,{doueJ oql pelJeJp taulul"sl '(1uec IBrouJA lseroJ {JellB qErq ur selod tu?urruopo3 ,{q lxau pa,rolloJ :ed roqloq,,r dnur[UBII l? s]olseJoJpelse (paqsqqndun ^\o[ 'OUISJ 'eJslleld 'Ir'{) ?t) lsoJoJ.{lJIBnb ur solodtueururopoJ ur lseleo]As€,r err?uuorlsenbe 0t6I uI JeururJ?elf,qpesn€3 uorlrpuoJ u^roJJ ur uorlsJouclcpeqI '(9 alq€J) uoltrpuoJ u/y\orJJo sosrpur(ZI=) oAeJo^Be^oq? rqtprra ,{lrsuapJsel pu? rlldap ur p?rltsoJoJ rl?lJ?i,{lrlenb,tolpue qSrqur sotod]u€lsrse{ 3u;1e.ror.ra1aparesuraoJJ qBJJBf Jaqlaq,lrJo luaurssassy '(xpueddy aes)7g ot (p?epeall) 0 ruo{ sc8wr xopureqJ 'soqJu?Jq pBJpJo JJuJprJUrJql pup JBB|loJol sJqJuprq ,{1r,lrpnpo,rdpus sJlrutufp pusls 'flrsuep JnuoJIdoJo uoqnqr.guoJ JBaIJosel€tllqsa pnsr^ 'sut\oJ3 uo posBq (8t61 sorurJC) xopul rm Jo uorsre^ porJrporx qeuEI 'rsunxJsel P pasn(uorsl^rq qJrBesedI l'IvJ) pJel|J ol ur SurArlserJods el?Jqolfe^ur cruepue/snoua8rpur JeqloJo elqqdecsnsro luelsrserqurreIluzunuopqns pue lu"unuopoJ sperpunq eql Jo plrqsqorJru eql ur uolerouEsp elqemsBeu 'eJoJeJer{J'uouslaroJ palselss Jo u,rorJ eql Jo uorlrpuoJ aql 3u4BJ slold f,ue pasnec1ou sar{Jazruolela{s fuoluo,\uruosarpnls dq pauoddns3lE suorsnlJuoJaseql luuJuru8rs ,{uBpo,roqs euoN s:o1ype-rdpue 'sprolrsered

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iiii 'JSJB ls.roJ {trpnb Aro.I ruo{ ueesu.rl^r snot^go sr su^rotr qE-uEfuo lf,edur Iresw lsed Jo luetxe [nJ t, ornilJ

'J?JB tseroJ^flpnb qBlH ruo{l ueesueqa snor^qosr su^rorcrlsJjef uo lcBduntJesu! lsed Jo lu.lre IInJ J} ,rntu (Table in Table6 mostdeterioration in crowncondition 9). Poles lowqualityforesthadsimilarcrownconditioninespective MEAN INDEX OF CROWNCONDITION FOR of locatici with respectto outbreaks,perhaps a legacyof JARRAH CODOMINANTAND SUBDOMINANT pastwildftes.Sim ilarrelationships were found for diameter POLESIN HIGH AND LOW QUALITY STANDSIN underbark (d.u.b.) increment (Table 9). TI{E SOUTHERN JARRAH FOREST Tabl€ 8 IN 1988 Crown cond;tion MEAN INDEX OF CROWNCONDITION FOR JARRAH POLES IN HIGH AND LOW Stand. quality Codomiaant Subtlominant STANDS IN THE SOI]TIIERN JARRAH Resistant Susceptible Resistanl Susceplible QUALITY FOREST. AIL TREESIN 1967HAD INDEX High (9 plots) l'1.5 11.3 r 5.6 12.'l VALUE = 24. Low (6 plots) 17.O 9.5 t6.2 13.1

Crcrvn condiion Table7 Srandqualiry Resistant Susceptible INCREMENT(cm y'') SHOWNBY MEAN D.U.B. High AND SUBDOMINANTJARRAH CODOMINANT Diamond 18.2(13) 16.7(12) STANDSIN POLESIN HIGH AND LOW QUALITY Marranup 16.9(14) 14.1(15) THE SOUTHERN JARRAH FOREST Low Boyicup 18.0(2) 12.9(9) Cessna 17.2 (r0) 13.8(10) Stand qualitJ Codominant Sublominant NOTE: (1) All but6 of the85 treesassessed in 1988had suMoninant High (9 plots) 0.23 0.11 0.19 0.15 (Spring 1973),Marranup t,ow (6 plots) 0.16 0.04 0.14 0.06 (2) Yearwhenlist bumt: Diamond (Spring 1985),Boyicup (Auiurnn l98l), Cessna(Spring communication)in 1967.The two plots in high quality 1962)- forestwere free of leafmineroulbreaks in 1968whercas thetwo plots in low qualityforest had experienced outbreaks for al least5 years(Mazanec, personal communication). Table9 Poleswere selected in 1968on lhebasis of havinga perfect MEAN INDEX OF CROWNCONDITION AND crown(Mazanec, personal communication), equivalent to INCREMENT(cm FOR 24 in therating scheme used in 1988(Appendix). MEAN D.U.B. Y'r) CODOMINANTAND SUBDOMINANTJARRAH Resistanttrees (as designrted by M^zanec)in all plots POLESIN HIGH AND LOW QUALITY STANDSIN showedremarkablysimilar declines in crownindex C24.2 COLLIE DISTRICT(NOVEMBER 1987), -29.6per -26.8per cent) over the 21-year to ceDt,ayerage All jarrahwere classified as susceptibleto leafminer. period(Table 8). This presumablyis causedmainly by drought.Fire wasnot a factoras one plot hadexperienced no fire since1962 (Table 8). Codominant Subdominant Stand quality Crown d.n.b. Crown d.u.b. indet increment index increment Susceptibletrecs (as designated by Mazanec)showcd High crowndecline of-40.5 per cent (rangc -30.4 to anaverage - in outbreak zone -46.3 21-yearperiod (Table 8). per cent) during the (4 plots) 13.6 0.1l t4.3 0.15 and Comparisonof the crown conditionof susceptible - not in oulbreak zone resislantpolesin eachplot shows that the average dif ference (3 plots) 19.4 0.40 19.3 0.51 is 18.2per cent (range 8.2 - 28.3per cent), i.e. leafminer Low outbreaksoyer two decadesare responsible for nearly20 - in ouibroak zone per centof the deteriorationin jarrahcrown condition. (3 plots) 14.2 0.11 16.7 0.07 - not in oulbrcak zone 0.16 Outbreaksof lea[minerreachedCollie dislrict between (3 plots) 14.2 0.07 \7.4 in 1987of jrrrah plots 1975and 1983.A comparison NOTE: Outbreaksof leafminer reachedthese plots belween 1975 and within the outbreakzone, wilh thoseto the north of the 1983. outbreakshows thatpoles in highquality forestexperienced

90 I6

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Thepreferred site of puprtionis beingstudied as a possible means ofpopulation Theinrermediate (900-1100 mm p.a.)and low (<900 control. mm p.a.)rainfall zones cover the parts ofthejarrah forest mostfavouredby leafminer. The mostrelevant watercourses The impactof an autumnfte on populationdensities includethe Helena, Canning, Serpentine, Harvey, Collie, is being assessedin an area of 100ha north-westof and Donnelly Rivers and Bell Brook, BinghamRiver, Manjimup. Harris River, BerakingBrook, Da.rkinRiver, Nockine BannisterRiver. Thoughseveral of these Brook, and The effectof sizeof the femalemoth on fecundityis feed into the reservoirsserving metropolitan sfreams beingmeasured. Perth,recentrelated research has indicatcd that any forest 20yerrs shouldnot causeappreciable declineoverthenext The northernlimit of distributionin jarrahforest has or increasesin watersalinity (Stokes risesin watertable recentlybeen determined. andBatini 1986). The occurrenceof bivoltine populationsand their CURRENTRESEARCH taxonomicstatus is underinvestigation. Detailsofcurrentresearch activities are setout in CALM's ResearchDivision Research Plan,znd - 4th editions. Bullseyeborer Theseactivities are only summarizedhere. Studicsof ovipositionpreferences are underway. Insectecology Monitoringlevels of herbivoryin jarrah forest Leafminer In 1984,an unbiasedmethod of samplingleaYes was developcd(Abboft 1987). A herbariumof damaged leaf,it Whenleafminer completes its fcedingin thejrrrah foliage,consisting ofvoucher specimens on which insects in a cell, which cutsitself out of the leaf whileremaining were obseNcdfeeding, has been compiled.It is now 1986,the presenceof falls to the ground.Since October possibleto reliably assignmost examplesof damaged 'cutouts' predetermined in leaveshas beenrecorded at foliageto a particularinsect species or family. samplingpoints in forestnoflh of the Collie outbreak. Thesedatr allow accuratemeasurcmellt of the rate of Damagelevels in groundcoppice janah in Manjimup spreadof leafminerpopulations north of the outbreal. regionhavebeen quantified annually since 1984, andsince 1987in Collieregion. Damage levels in polecrowns have laryae in mines The level of parasitismof leafminer beenmeasured less frequently. Damage levels in ground outbreaknear within, and to the north of, the leafminer coppicein theJarahdate-Dwcllingup region were measured Collie is beingassessed. on the1985, 1987 and 1989 cohorts of leaves. leafminerand Theeffect of moderateintensity fte on Vadationin damagelevels in termsofsite andseason in 240ha ofjarrah conditionof crownsis beingassessed arenowwell understood.Multiple regression analyses are 1989.It forestnorth of Collie whichwas burnt in autumn being used to relate damagelevels to enYironmental reduce mteof is hopedthat these fires could be usedto the factors.The complexity of interactingfactors makes northernjrfiah spreadof lhe outbreakfarther into the factorialexperimentation impracticable at this stage. forest, Reseffchinto economicinjury levelshas tbcused on of other The effect of leafmincrdamage on biomass manualdefolirtion of groundcoppice jarrah. Itis reasonable jarrah investigatedat l5 invertebratesin crownsis being to expectthat a thresholdexists, below which damage has sitesin Collie district. no impacton jarah growth.An experimentusing ground coppicein HolmesBlock wasset up in December1987 to Skeletonizer assessthe interactionbetween proportion of foliage removed(0,25,50,75,100 per cent) and number of annual Studiesof the populationdynarnics of caterpillarshave defoliations(0,1,2,3). commenced.These will indicatebetween which instffs mostmortality occurs. E6

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A physiologistcould cxamine small numbers oftrees, 4. Virological,bacteiologicalandlormycological options within and without outbreak areas, and resistant or for t:ontrollingleafminer ancl skeletonizer susceptibleto defoliators, and draw precise conclusions about impacts of defoliators on lhe well-being of jarrah Ifreseffchon pansitoids yields little ofvalue, it may trees.Very early waflling ofany impcnding masscollilpse be necessuyto studythe naturalimportance of viruses, of stands would be gaincd. lt would also be valuable to blcteriailnd fungi in killing pestinsects, and to evaluate evaluate the physiological responsc of jarrah to a whether other viruses, bactcria and fungi should be combination of experimentalintensities and repetitionsof introducedto control pest insect populationswithout defoliation. detrimentto non-pestinsect populations.

In pa,rticul lhereis r needto examinewhether aeri: 2. Reasonsfor theinitiation and conti uaion ofoutbreaks sprayingo[ foliagewith commerciallyavaila.ble Bacillus thuringiensiswould contributeto control of leafminer, The outbreak of leafminer that bcgan more than two withoutprcjudicing conservation valucs by reducingthe decadesago in the southernjarrah forest hrd no historical abundrnccof otherinvertebrates (particularlyLepidoptera) precedent,since European settlemcnt. Newman and Clark living in jarrahcrowns. (1925)recordcd that noticeabledamage was confinedto nrhgrowing oo theplains country', mainly the'coaslalj 5. Selectiottof jarrah resistant to leafminer and near Fremantle, Bunbury, Busselton irnd Albany, but ' skeletorizer mdiating inland for several miles to the base of the no was foothills'. They noted explicitly that damage Aworkabledefinition ofresistance nccds tobe devised. Torbay and Busscltonor recordedin the countly bctween Treesand regeneration immune to pestinsect infestation nearPerth (Kings in the Da.rlingRange forests. Outbreaks couldthen be identifiedand rctained throughout thejarah Park) were first recorded in 1914. forcst.The feasibility ofreadily increasingthe proportion of resislilnttrees in standsrequires investigation. This historical perspective invites the question 'What initiated the 1960s outbreak in forest east o[ It is necessaryto testtrccs with resistantphenotypes fire (a Manjimup?' Obvious perturbating factors include (i.e. crownsunaffected in standssevcrely affected) and chalge in inlcnsity, seasonof burning, frequencyof cstablisha geneticbrsis by collecting seed,Srowing factors),logging, burning or combinationsof all thrcc seedlingsand testing them in cultureand wilhin forest. deforestation (for farming and settlemcnt) and subtle changes in climate (rainfall, temperature, soil dryness 6. Compurisonof bird populationswithin a d outside index). Mazanec(1988b) argues thatrmy factorthatleads oulbre0kareqs to the production of new leaveswill lead to an increasein population of leafminer. Exiensivebrowning ofjlrrah crownsmay lessen their suitabilityas habitat for insectivorousbirds. Expansion of Associationsbctween some ofthese vrriables atd outbrealareas may resultin someof thesebird species cxpansion of leafminer and skeletonizeroutbreaks could declining in distributionand abundanccwithin jarrah (1987) besought by modelling.The mrps in Abbott would forest.The suitability for birds inhabiting crowns of seryeas a start. Then thc more accuratemaps that would unaffectedspecies such as [rarri or yarri alsoneeds to be beproduced using rcmote sensirg techniqucs would allow assessed. refinement of thesemodels. It is hoped that thesemodels could explainwhy lcafminerandskeletonizcr have become 7. Taxonomicexpertise forestpestsonlywithin thc pi]st25and 5 yearsrespectively. The taxonomyof many potentiallyeconomically 3. Searchfor parasitoids of leafininer and skelelonizer impo ant insectgroups is out-of-dateor confused.The elsewherein Australia major taxa are Chrysomelidae,Cerambycidae, Curculionidae,Doratifera, Lyctus ^r'td Oc hro gaster. This would complement the research complcled by

94 96

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Abbott,I. (1987).Review ofpast and currentreseffch into Mazancc,Z. (1980). The jarrah Leafminer,Perthida insect problems in the jarrah forest, with gly p hop a (Lepidoptera:Incurvariidae) (Common), a recommendationsabout the future dhection ofresearch. pestof jarrahforest in WestemAustralia. M.Sc. thesis, UnpublishedConfidential Report. CALM. Departmentof Zoology, University of Western Australia. Abbott,I. (1990). Insectoutbreats in forestsof Western Australia.In:Population Dynamics of ForestInsects. Mizanec,Z. (1981).Environmenialconditionsinfluencing Eds.A.D. Watt, S.R.Leather, M.D. Hunterand N.A. thepopulation density of thejarrah leafminer. In: Old, Kidd. Intrercept,Andover, Hampshirc. pp 9-5-103. K.M., Kile, G.A. and Ohmart,C.P. (eds),Eucalypt diebackin forestsond woodlunds'.proceedings of a Abbott,I., Smith,R., Williams, M. andVoutier, R. (in conferenceheld at CSIRO Canberra,August 1980. press). Infestationof rcgeneratedstands of kari CSIRO,Melbourne. pp. 140-14. (Eucalyptustiiversicolor) by bullseyeborcr (Tryp hocaria acantlrccera, Ccrambycidae) in Western Mxzanec,Z. (1983).The immature stages and life history Auslralia.Austral ian F or es try. of thejarrah leafminer, Perthida glyphopa Common (Lepidoptera:Incurv ar\idxe). Jo ur nal of th e Aus tr aLi an Behn,G.A., Wallace, J.F. and Hick, P.T. (1990). Airborne EntomologicalSoctsry 22, 101-108. multispeclralscanner data for mappinginsect damage in jarah forestof WesternAustralia. Paperpresented Mrzanec,Z.(1984a). The useof gonadsin larvalsexing at RemoteSensing Conference, Octobcr 1990, Perlh. of thejarrah leafmircr, Perthidaglypropo Common (Lepidoptera:Incurv ariidae). I o ur nal of the Austr al i o n Bradshaw,F.J. (1987). JarI. h thinningand regeneration. Entomolo gical Soc ie tf 23, 11 - 12. Silviculturespecificalions l/87. CALM. Mazanec,Z. (1984b).Studies or thesex ratio in thejanah Clark,J. (1925).The marri borer(Tryphocaria hamnta). leafminer Perthida gLyp ho pa (Lepidoptera: WesternAustrulia Journal of Agriculture,2(2) ,513- Incurvr-rii dae). Jour. nal of t he Australian Entomolo gic al 5r7. Society23,69-74.

Curry,S.J. (1981). Native wasps hit introducedfree pest. WesternAustralia JournaLof Agriculture22 (4),6l- Mazanec,Z. ( 1985).Resistance of tucalyptusrrarginata 62. to Per t ht d a gly p ho p a (LepkJoptera:Incurvariidae). Journalof the AuslraliqnEntomological Society 24, Grimes,R.F. (1978). Crown usessment ofnlfural Spolted 209-22r. Gljm (Eucalyptus maculata), llo'rbi:rk (.Eucalyptus fi bro s a, E uc aly p tus dr e p a no p hy I L a) f orcst.Queensland Mazanec,Z.(1986). Oviposition bchaviour and dispersal Departmentof Foresky,Technicill Paper No. 7. of Perthida glyphopa Common (Lepidoptera: Incurvariidre)."/ozr nal ofthe Australian EntotnDlo gical Havel,J.J. (1975). Site-vegetation m:rpping in thenofihern Society25, I49-159. janah forest (Drrling Rangc).1. Definition of site- vegetationtypcs. Fdre.rtJDepartme t oJ Weslern Mazanec,Z. (1987).Natural cncmics of Perthidaglyphopa Austalia BulletinNo. 86. Common(Lepidoptera: Incurvariidae). Journal of the AustralianEntomological Society 26, 3Q3-308. Jacobs,M.R. (1955). Growth habits of the Eucalypts. Forestryand Timber Bureau, Canberra. Mazanec,Z. (1988a).Immaturc stages and lifc historyof Chrlsonotomyiasp. C (Hymenoptera:Eulophidae), a Mazanec,Z.(1974). Influence of jarrahleafminer on the parasitoidofjffrah leafminu.J our nal oft heAustr qlian growthofjrrah. Australian Forestry 37 ,32-42. EntomologitalSociety 27 , 219-292.

Mazanec,Z. (1978).A samplingscheme for estimating Mazanec,Z.(1988b). Jarrah leafminer, a pestofjarrah. In: populationdensity of the jarah leafminetPerthida The j arra h fo res t - a comp le x M edit ena ne qn e c o sy ste m. gllp hopa(I-epidoptera:lncurvariidae). Jour nal of the Eds.B. Dell, J.J.Havel andN. Malajczuk. Kluwer, Australian Entomolo g i cal So c i ety 17,27 5 -285. Dordrecht.pp. 123-131. L6

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Assessmentofjarrah crowns (basedon methodof Grines 1978).The valueof theindex ranges fro m 0 to 24.

LEAF'DENSITY (excludeepicormics on bole) 9 very dense 7 dense 5 avemge(refers to ideal, not the stand) 3 sparse I very sparse 0.5 all leavesdead 0 leavesabsent

If muchfruit is present,increase the scoreby l.

INCIDENCE OF DEAD BRANCHES 9 none present 8 < 5070of branchletsdead 7 > 50Voof branchletsdead 6 < 507oofsmall branchesdead 5 > 50Vaofsmall branchesdead 4 < 50Voof large branchesdead 3 > 50Voof largebranches dead 2 < 50Voof pimary branchesdead (epicormics present) I > 507oof primarybranches dead (no epicormics) 0 all branchesdead

CONTRIBUTION OF EPICORMIC BRANCHESTO FOLIAGE 6 no epicormicspresent (foliage concentrated at extremitiesof branches) 5 on crownor bole < 25Vo 4 on crownor bole > 25-507a 3 on crown or bole > 50Vo 2 on crown and bole < 50Vo 1 on crownand bole > 50Vo 0.5 on crown andbole lffiVo 0 tree is dead

98