Eucalyptus Rhodantha Blakely & Steedman, the Rose Mallee, Merits Special Attention for Several Reasons

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cuo,ilrfliro{ ACKNOWLEDGEMENTS

We thank the Scott family for allowing access to their ProPerty and for their invaluable contribution to the conservation of E, rhoilanthg.

The Western Australian Bushfires Board and staff of the Moora office of the Departrnent of CALM provided the equipment and exPertise for a controlled burn. We gratefully 'acknowledge their assistance.

We thank Shapelle McNee for her comrnents and advice and for acc€ssto unPublished results.

w 'saDods ar$ Jo Ie r^rns ar{l aouEquapue amsua ol pa}qllrqeqeJ aq pporls slupuruar rallellls lPrR lnq uope JasuoJ roJ AlrolJd rsaqSJq aql s! puEI paJEapun uo luBuul3r 1sa3re1aq1 'pos?q lur{l a}EDtpu-tsuollP8Fsaaul aq ,lfttu ldllsma pa}sullld-pjtq IPnsnrm sl+ ;o luaure8euau pue uopelJasuoJ aql roJ sar8ale.qs 1euort€J rplqn uodn qsuq u $rroJ It.lVJ Jo luauqJeda6 eql rlrlrn uonepo€se u-r l11s.raa1unuIunf, Pue EllPqsnv u.e4sa a Jo ,{11s.ra,r1unaq1 dq pepnpuor 'uolpuBxa 'Po rprEasou r{tF{ PaualPano aroJadar$ s1 sapads arI,L rasqo uroq lou seq sStrgpaas 'sea.re Jo luauqln GraJpue spuqs palEtosl'IJsrrrs aql u1 ,(puerg.ru3.rs pasearJul seq 3u1paa.rqu1 IpJnrln4€e uI lJeI seltds sFIl Jo qusuurar leurs ruay e ,(1uo are a.ritUf, sapeds luqd pa.raSuepua pup a.r€r lsour s,plls.l1snv uJalseM Jo auo q ueupaat5 ry d1ary1g otlflr'poqt snqdlptng

luvvw{os 2.3.1. The breeding system......

3.2.7. Liaisonwith landownersand Shires...... ,,....24 3.2.2. Land acquisition...... 24 3.2.3. Protectionfrom grazing ...... 25 3.2.4. Protectionfrom accidentaldestruction...... 25 3.2.5. Protectionfrom fire...... -- ...... ,.26 ...... 26

vl IIA LIST OF TABLES

Page Table2.1 locationsand estinabd sizesof rermant startdsof E. rhoilantha...... ,....'.....'..6 Table2.2 Estimatesof the outcrossingrate (t) in eleveneucalypt species...... '...... 11 Table 2.3 Total geneticdiversity and the distribution of diversity within and between populations of treesof widespread regional and localised distributions...,. Table 2.4 Estimates of the mean nurriber of migranc (Ntr) o

Table 2.5 Mean seedand seedlingcharacteristics scored in the plogeny of tm E. thodtnlha vs fmm Sbnd 3 and fmm eachof two subpopulations in Stand4...... ,...... 19 Table 3.1 Standslisted in orderof priority for managernentactions...... ,..,.,.,,.....29

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silun:trr{o rsrl 1. INTRODUCTION

Eucalyptusis the most notable and distinctive tree genus of the Australian flora. There are over 500 species and subspecific taxa confined almost entirely to Australia and the genus has great economic,scientific and aestheticimportance (Pryor 1981;Chippendale 1988). Furthermore, because eucalypts are such a dominant element of the flora, the fate of many associatedplant and animal species depends on that of eucalypts. The genus therefore merits the attention of both those aiming to conserve the diversity of natural systems or the viability of agricultural systems, and those whose concem is the preservation of economically useful genetic resources.

Eucalyptus rhodantha Blakely & Steedman, the rose mallee, merits sPecial attention for several reasons. Firstly, there are less than 500 plants in existence so it is one of Western Australia's most rare and endangeredspecies (Lucas and Synge 1978;Pryor 1981;Rye and Hopper 1981). Secondly,the remaining plants occupy soils and situations where their continued survival is uncertain and finally. the survival of this striking species is also threatened by a decline in outbreeding in small isolated standsand by a lack of natural recruitment (Sampson1988).

IdeaIIy, species would be preserved within ecosystemsin vast reserves,but this is often not possible. The agricultural wheatbelt area of south Western Australia was essentially virgin land at the turn of the century but is now virtually completely settled (Main 1987)' During the early settlement phases there was no policy of setting aside areasfor conservation of flora and fauna so the loss of vegetation in this area has been considerable. Speciessuch as E. rhodanthahave been reduced to remnant stands in a matrix of agricultural land use.

E. rhodanthahas been gazettedas rare and endangeredflora and is therefore protected under the Western Australian Wildlife Conservation Act 195G1979. However, the speciesclearly requires management to ensure its long term survival. To be effective, conserYationand management of speciesin small reserves and rehabilitation or founding of new populations should be based on genetic and ecologicalprinciples.

Recently, detailed research into the genetic systems and reproductive biology of E. rhodantha has been conducted by The University of Western Australia and Curtin University in co-operation with the Department of CALM. These investitations, to8ether with ecological principles, form a basis for the development of rational strategies for the conservation and management of E. rhodantha. 'arnlprp8e papualxa ll teql pauodar (196I) rauprpg ro1 par€alJuaaq seq uopelaSa.rleur8uo aqt 'aJurs €aJparg Jo lso\u ur spupls lupuural uoly dluo ul'lou1.sr n4flupotlt.A uoqnqrlsrp ]uasard aq.1

lElIqsH pue uollnqlllqq .z.z

'11961 rod:4y stwtotufrd'Xpue odntonna 7 qtp,u drqsuoqelar aruos Sur,reqlnq slddprna 'saoads rarllo uror; lrurlsrp palrlllsar dlq8rq e se papre8ar rv.'ous a1luupo7l .7 .11961raddoll pup adll) e:og pa:a8uepua pue aler sz pa11aze3Suraq 1r o1 pal pue prrqdq e $u sI DU+ubpor.lr.z 'ra1e1 'sprrqdq leql paqsrlqelsaq)reasar srwto{ufi.d7x tdtntontw.A IEal pue b7qopoqt.X uaa-.{laquolsnJuoJ uo paspq se.u uoqeprdralur sy{J .zJJnI stuotufid sn1dfr1ttn7x tdtatonnut .1 p yo plrqdq aq ol lr paJaptsuoJdaql arurs slddlerna aql Jo uorlpJursselJ(116I) s,uosuqol 'g .sanads pue .rod.r4ur papnlJul lou set oqluopo4l auo o1 pa3uolaq nljuapoyt.g Jo sruJo; IIe l€r.{} 'ra^a,aa.oq/(Z6l) 'sartads paraprsuoJ alppuaddu3 lJunsrp lnq alqeup^ e se D4ryupo11t.3pap:e8a.r '(Z.Z.2\J:g16l pue dlarrea slqt pasluSorar(1961) rauplpD tt p d1a>1e1g;ueurpaatg { dyalelg 'uor.{sp 4n7o17ad're^ sB paqrJ)sap ser,ralorlad lpg pue salpal a}ploaJuel-el€pJoJr{}L\{ urroJ V

'atutdntontptl ,uaprpl sarrasqns pue andtrtotcor{ saltas lpruto;ur aql ul l plood-aut1 sn1dfr.1ttn1pue 'papnl)ur 'eE6I'to rdtutotttw e qllll^ Suotp nou sr ll la d1a1e1g;pg61 ur uEupaats .g dq rprdung 'sdrqsuorleyar rpau paDalloJ lpualer.uuro{ parueu sp,4r11 pue snlels ller.ualsds sll }noqe luaurn8re ol palnqrJluoJser{ srq} pup urljuflpotjt'3 ur palrodar uaaq seq urroJur uoqer&^ alqpJaprsuoJ

'sraMou pallpls pue sa^pal rallprus 'sanads slr dq'1oo11 adtntottau snqdfrlttnX .relrurs aql tuo{ paqsrnSurlsrpaq Aevst41unpo4t 7 '(g pue O I'Z'8rJ) spaas paSuur ua.r.orq1lpp ol u1v{orq8ug1e1uor saap,r .leln8ueul 8urpru1o.rd 'leruaqdsgraq .O .8lJ) qll^{'urJ g lq uD g o1 dn pue dpoo,ra. ssalro a.roursr }lrut aqJ I.Z sluaurelu lurd o1 ,r.ro11adaled qlr.n ruro; p sr a:aq1qSnoqlp sraqlue .no11adql.tl sluaurplr] par a,req dylensn .3uo1 pue ssorJeruJ g'4 o1dn aq deur sra,uolg ur 7 o1dn 11e1se dq paqJqlp pue aseqaql ueql ra8uol 'leuuor 'apr^{ .(A .8lC) dlerrd{ sr der aq1 ruJ 7 pue 3uo1rur g.9 o1 dn aq deru pnq darS qreg I.Z ,g 4e1s dar8 prql e dq paq)e11psl qrll.lt laqun aql uI spnq raaog o1 dn aq deur 1nq ,auo dlensn 'qtpr.\ sr araql ur arues aql lnoqp pup q13ua1ur r.ur g-9 do1 aql le palulod ,alpploJ-JplnJrqro '(V '3lg) .IJIqt are sa^pa.I l'Z alpssnJap pue allsoddo dlleJrddt aJ" qJIr{,^^sa^€al 'an1q-da.r8 .ser{Jup;q snorney8 dlq8rq s1rdq paqsrn8uustp sr lI darS-alrq.aapue lreq u.uorq-da;8 'qloorus 'lqglar{ qlur{ ur r.u g o1 dn s,rlor8 qrrq,u aalpur Surpeards ^ ol e sr otllutpoqt.3

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Figure 2.1 Diagranrs of the distinctive floral and vegetative structures of E. fhoilantha var. tho ilantha A, leaves;B, buds; C, flowers; D, fruiU E, seeds[from Gardner (1951)]' Ir

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'(uoEEJrunuuroJ 'o 's) IaJosJadraddoH ll uoddns tou op suaurJads umueqtar{ 'PIJroN ar{l aJuls alernJJp aq lou Ietu urallEd slqJ ^ aN rPau ol spJP/uqlnos pue ooraqleM '3 uroq pre,^{lsPasalFrr auros'rpl{unD ol (odtuonoa qrli pax$r se^{ ll araq,^) Ja Iu IIIH tuo{

sunlo4ad'nt nqlurpoqt'g to sa^?al a^IlJupqp aq1yo ruer8ulq z'z arn8l{ Perconal communication). Prior to European settlement, E. rhoilantha probably also consisted of a narrow belt of discontinuous populations of varying sizes.

E, rhoilantha var. rhoilantha is now confined to two areas between Three Springs and Watheroo (Fig. 2.3). All stands occur within the Department of CALM Greenough Management Region. Locations of and estimated sizes of rerunnt stands of E. rhoilanthn are given in Table 2.1. The 'extradry climate of the area is mediterraneart' (Beard 1984) with an average of 80 days rairy the maiority of which occurs during May to August. The mean annual rainfall at Watheroo is 428 mm (Western Australian Bureau of Meteorology).

eraldton

O r-'-

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I Perth

Figure 23 Geographicaldistribution of remnant standsof E. rloilantha in soutft-western Australia Occursonly within the Deparhnentof CALII{ GreenoughManagement Region ( - - - - ). uJstu lsPa vS )|?opped c )poPped c qUou e6 lsPa rnJou vZ ls3^L r{uou vl sltlloltad '&^ otllurpoqt,X I auIlaJuaJ v8 9 qlnos vl 6 Peor q (EI urPur lsE€t vs 08r urErn lsallr vl ,T 19yo uopelndodqns) aqplaa,D vu€ LE {Jopped v€ 8 lsEa r{uou ,z ,l lsarr rnJou +r oqrurpoqt'Je^ rttubpott 'x

'? 'oN ,tlllttporl Jo paleuFsg auPu puPls $qumupue$

'g.7 '3rg ur duu aag .dgadord ageau4 .agraa ppoU v + rr$atpolt . to sprn'ls lu"uEar to saz-rgpafrllpsr puE suoll"Jol Iz alqpl

'3gg 11'7 laopped E ur sq SZtnoqe raao paragux are sltrqd 97 Srmrrerua,r 'aup{aarJ aqJ e Suop puq pareap ,(1pg.red uo auros pue aglaA ppor € uo auros ,(Eg pupls) 'ptrq .raq1a3o1padno€ saellEur tI Jo slsFuoJ € puels pmlpru8e 3r4prmo.r.msarp ruo{ JJo paJuaJ .rltwag dlapldruor sr pupls poreallm auo Jo "r{ g,Qaleurocorddy tqapldoxol srydfrInn7 pue .pul.l .ds ,f,a1pt;1 sapuoltd !!q rg lo spuels ryacu[pu aru ararll tltny pue.ds tluoUlute 4tt1so1ta ''rg'\ ^rg .d ,allraatg trtrlts oaltH (.lrs) t1otnfi4 rrltH 'slarq q4aalgy s1tt111dna ^t41uag ^tg'a .lung ..I .g sndappo4 unsourd,s wnlqop4st, sapltlqrnb snuuotqloy3 !fraqqso 's1a';q upurfitg q4saduw tuuonsaxollv 8u-rpnpur uo4v1a3a qtuar{ lrlol auos r{lllr saqrunurrroJ 'puel a,rnd lsor.up sp mrJo asarll p eapun uo aru (g pue t) spupls urarllnos aql Jo o^q IIuO 'uDI arBnbs 9I lnoq€ Jo Eare uE uItIlIA pernqy+slP sptrqs urarDnos g plrE Jarno qJue Jo uDI I uF{+y,^ 'spopped Sw.r.rnrao spusls uralpJou oir l ale ararf,L ur perallecs puu sapsp?oJ uo JrDJo rr ou slrreld ',fi1unoa 8u;lqnpun ,{11q3r1sro IEU uI }os dpups da.r3 uo salo:3 otlltrtpoqt,re^ u&utpotlt.X 'oo o ..._a

Figure 24 Diagrammatic representationof Stand 3 showing the diskibution of E rloilartha arld flre locadon of subpopulation!]a (. ) Stand 3a mallee, ( o ) Other E. thoilantha , ( 77 \ Other species,rnainly Ercalyptus toiltiana F. Muell. ( - ) Fence. (.) Unclearedland along ceekline. The scaleis approximate, 'sl:.t .g olJ Dqlulpoly uro{ ua11od1sa,r.req deur urnssod-dauoqaq1 1eq1 s1sa33ns aJuaprlg 'slrqqeJpue soore8uel '(asnour asnoq) snlnJsna snry se llla.|vrsp t pupls uI pa^rasqo uaaq a^pr{ (ueunp) s6do{]4l1rs pup (urnssod-Aauoyy sryn4sot sadtstal 'salrads qq1 uror; sreaulsua11od u1 punoJ uaaq 'rala.lroq 'seq ua11odr4l uopo1t.7 .8ur3eroypa^Jasqo uaaq lou seq lnq spupts tqluopoqt 'g ur pa^rasqo uaaq oqe seq,suolqp *tfr.uopt1fiq4,:aleadauoq 'salgrallnq ,.dds 'vnlglaw rar{louv pue sr{loru IpJaAaspue saaq a^l}Eu xawtlwoprtl s y ''dds .rplrau swsrlot1 nrpo',snp111td, snqoptd lup ar{t apnlJur sarJads lsalr€r{ oslp slJasul

'(uollpJlunu-ruror .W leuosrad aaNJI I) 9g6I \ qnslnD IZ pue sxnsana.j IZ'apu4slput J 0€ aJa.{ sJa}EadauoqloJ palrasqo staqutnu rumuxpur alls ureur lsa.u aq} }e sluno) 'snutuoz snsual lulod u),ot1 g loJJed alql pue ryqnlunnJ uau4)oqluv pue qn&nr[ tutJouvw 'suaJsafia 'DlJutlstput snwo$ouaqx!1 rfiwU)t1 salJads plJq ralpadauor{aq1 Sulpnpur sarmds .r(ep pralas dq palsa^J€q sr JplJau s1q1 aql Suunp dpreu,relrau ro slunorup snordot acnpord pue lsdep 9g-gg; paa11-3uo1are sra.trolJ 19g6I €NJW) ra.arrogdlpnluaaa r{lqn spnq Jo raqrunu 'snlJlltoz aql sa)npal sntpnuna'1oled aql pup ([r^aa,4,rpnq vtllutpoql .X) snanxtw xfiuo1dtp1 'srlluorJ dq uonepar4 rauurns ar{l SulJnp sJnJf,ospnq yo luaurdola,rapaW pup rl}^{or3 a,rpela8a,l

'aprJr.rroJ lou prp suoseas .qtea Supa,raogJ!ar{} asneJaqalqlssodurl su,u s1ue1daruos u€.{1aq uorleuulod-sso.r3 luar rad 1 ueql ssal palnqlJluor queld lq8ra gqnli uoseasarqua aql Ja^o paproJarsra,ra.og uado aq1;o luar rad .8uuar*og Zt'02 palnquluoJ 1ue1dauo'9961 ur quuld 97;o dalrns e u1 1o uouernp aqt pup psuo Jo auq ar{l ur pue arnpord ,{aq1 sraaou Jo raqumu lelol aql uI dlq€raprsuoJ raJJrp queld pnpr,rrpul

.sra^ ,9g6l .gg6l ou 000€ara^^ araql ur puels auresaqt uI uI pusls ulpur lsa^r ar{l uI Suuamog >1ead1e pap.rora.rara..vr $a.4!ou ggg 'aldruexa rog .sread uaaru.laqdlluerlpu8ts p .uospas d.:err uec uospas u1 parnpo:d sra^rou Jo laqunu lelol aqJ Sugar'rog leuoqdarxa 'ra.ramog .3urra,r,rog uu u1 lueld a13u1se uo papro)ar uaaq a uq sJa^rou g0Z 01 dn >pad 1e uarra 1ueld qrea uo s,rart{oguado 199-1;,r,ray ,fia,r sarnpo.rd dlpnsn 1r1uq1ur qddprna 1s8uor.uelensnun sr saoads '3rg; aq1 19961 uosdurps 19961€I\fq/{ 1g'7 a1s aq1uo Surpuadap 1sn8ny ol aunl to sr{tuotu ralul^{ aql uaa,rqaq a8ue.r satun Suga^ oU lead aql lnq raqrualoN pu" rlJrpr l uaamlaq sralrou br uupoqt J

,{3o1org'g'7

'pauue; ,{laarsualxasr Eare aql ' aJurs punoJ aq IFla. spupls aJorx leql d1a15rn sr \ llutpoqt.Je^ oqlurJpoqt g qlr.tr uorlerJosse 'l!os ul saurlFrxos dpups u1 drlunor 3u11e1npund11q311s ro lpg uo sJnJJo11 .s8ur:dg €u{J rpau 'seale Jar.{loar{l pup ooJar{lp6 Jpau auo o^ l urory dluo r.!!!ou1 sr *n1oqad.E^ o1lurpoUt.A -t- Number of flowers* + % of trees flowering 100 100 90 90 80 80 bo d/u 3 €60 ouo

hJU 50 *t Eto ; z ba 30 30 20 20 10 10 0 0 Jan F€b Mar Apr May In Jrn Aqg Sep oct Nov Dec

Month

Figue 2.5 Estimatednumber of flowers r€leasingpollen and propotion of plants releasingpollen in a sample of 25 plants in Stand 4 during 1986 *, open flowers with intact staminal rings. AIter Sampson(1988).

L. oirescens and M. flaoigula are thought to be the main pollinators of E. rhodantha (McNee 1986). Thesespecies have been observed nesting in the west main stand, but the nests were not in E. rhodanthaplants which are very open and provide limited protection. Bird speciesdo not forage onE. rhodanthaexclusively but use a variety of flowering specieswhich occur locally, including B. prionotes,D. ashbyi,G. eriostachyaand C. quadrit'idrs. The speciesbeing exploited alters throughout the year. Some types of bird may forage primarily on plant species which flower concurrently with E. rhadanlha.

23.1. Th€ breeding system

The flowering strategy of E. rhodantha and observations of bird behaviour suggest that it is

pollinated primarili by birds, mainly M. flaoigula and L. obescens.Flowers are large and produce copious quantities of nectar. They are pendulous and the stamen filaments are conspicuously red. These features characterisemany bird-pollinated plant species(Ford et aI. 1979). OI

'arull aurcs aql lp raaog leqt slueld uaa..'tlaq pue spooq.rnoqq3rau dyluuy leus ,(la,rqu1ar 'sJnJJo ulqly* s1ue1dpal€lal uaa^ laq s8ugpur Jo llnsal p se dlqeqord osle uorleuqlod-Jlas 'spooq:noqq8rau uer{l raqlo Surpaa.rqur aurog lueraJJlp lllerrlaua3 leJalas }o lslsuo) spupls aql l€ql pup palFrll aq deru spuels ul{l!1r^ i ou auaS leql palea^ar oslp suoFp3llsa^ul

'd11s,raarp pue Sulssorqno alouro.rdderu slddpana u1 uopeqlyod-pnq lpql srsaqlod,{qaql uoddns dlrs.ra,rrprpauaS ;o uoqnql4slp ayl p'ue o4luopo{r'g Jo spupts l€Jn}pu ul Surssonlno 3o 1az'a1 'aroJalaql 'ur aql 96 dq rnoqqSrau lsareau rraql rro{ palelosl aq dpur queld auros 1eq1os ,{qr1ed sr slpnpurpur Jo uoqnqrrlsrp aql rIJIq^r ulqlpr spupls palplosr lleurs s€ palnqustp lue1d alqrleduror '3 -JIas p ro; Sulssonlno Jo la^al lppuplsqns p surquruur trlluupotlt suollelndod asuap'a3.re1 dlaar1e1a.rur rnJJo qJrq^r saall€ur ro saa4 lsaJoJuaaq a^Eq parpnls saoads raqlo aql 'ralar',r.oH '(Z'Z alqpl) gd{1erna :aq1o:oy pauodar aguer aql }o pua ^{oI aq} le sr 69'0 Jo alur Surssolclnouy

'uoqeurllod-Jlas Jo uoprodord luergru8rs e lnq Surssoltlno lueururope,rdrn1,l{ rualsds Sugeru paxlur z seq t1rytpoqt .J 'suorle8rlsaaur 'pue leql palsaSSns raqlo r{llm raqlaSol 69.0 ser o1luf,potlt.g Jo pupls '(SurssoDlno paqrnlqpun d1a,rr1e1are u1 aler 3u1sso.rr1noJo alprxqsa aqJ alalduror; I pup (SulJIas alaldurorl 0 uaa^^laq a8upr upJ rlJlqi (j) aler Sulssonlno ,a^praJra,aql Jo aleupsa ue appo-rd '(0661 sarnparord asaql l, ta uosdueS 13961uosdureSl sarnparord uoppulpsa pooqlra) r-unrup{pr.u 'g pu€ sralJeur arudzosrSursn Irplap ur paleBgsaauruaaq s"q ufiuopo4t Jo ualsfs Suqeur eq1

'stdIlpJna ueller1sny uralsa6 paleulllod-a1erqa1.rarrpa.nr*og-a8rey 'A rar{lo aqt Jo lsor.upuu otliuopotlt asgap€r€qJ qr1q.ra.suonelndod pa}Fsslp fierus uI paJnpar are Sulpaa.rqur;o saauanbasuocsnoFalalap aql dqaJaq..l4.rusnmJmru e apraord deur uogeu!fiod pJrq lerg '$rprrqdq fnlsaSSns sroqlne asarlJ pue Surssor4no aloruold ol tualMs p Jo uorlnlola aql luasa:da.r deu qddprna ur uoqpulllod p4q Jo uollnlo^a ar{l lpr& palsaSSns(Ig6I) uprol/{ pup raddoH

'saoads qq1 u1 mrro saop uorleuqlod-;1as l"ql pup alqpedurorul-;1as 1ou are daql leql s1sa33nsq)n{.^ paas las u?J suol}pJol palplosr ur 'auJrl s1ue1da13urs €JoruJaqgnd auo ,(ue lE lueld u uo uado aq leur luaudola,rap ;o sa8pls luan;tlp lp sJa^rou lpJa^as aJuls D1tuopoqt g u1 paluaaard lou sI uorleulllod-1yas 1nq uorleurllod-Jlas '(9g6I luarrard ro Dnpar ot ruslupr{Jarue aq o1 lq8noq} d1pIaue3 sr ,{:puelo.r4 aaNJI I) srsar{+up raryp slep ZI lnoqe lqun a.rrqdarar aruoJaq lou saop euSgs aql alrq^,r slsar{}ue Jalrp sdep ua,ras ,ro 'sno:pue1o:d xrs urr{llr sror{lu9aql luorj poqs sr uaJlodaql Jo lsow are f,\luopo+t 3 Jo sra^,\oll Table 2.2 Estimatesof the outcrossingrate (t ) in elev€n eucalj4rt sPecies

Species Source

E, rhoilantha (undisturbed stand) 0.59 Sampson(1988) E. thoilantha (disturbed stand) o.26 Sampson(1988) Eucalyptus obliqua L'}{erit 0.76 Brown ef al. (1975) Eucalyqtus paucifloraSieber ex- Spreng. 0.70 Phillips and Brown (192) Eucalyptus delegatensisR. T. Bak. 0.79 Moran and Brown (1980) Eucalyptustegn4ns F. Muell. o.69 Moran and Bell (1983) Eucalyptusltellulatr Sieber ex DC. 0.77 Moran and Bell (1983) Eucalyptusstoatei C. A. Gardner 0.82 Hopper and Moran (1981) Eucalyptus bitso ^na (Luehm.) Maiden 0.77 Fripp (1982) Eucalyptus citriodora Hook. 0.86 Yeh ef al. (1983) Eucalyptusgrandis Hill ex. Maiden 0.84 Moran and Bell (1983) Eucalyptus saligna Sm. 0.77 Moran and Bell (1983)

An important factor which probably contributes to the maintenance of a mixed mating system in E. rhodantha is selection favouring heterozygous progeny which are more likely to be Produced by outcrossing. The levels of heterozygosity observed in mature E. rhodantha plants were significantly higher than those detected in seeds or newly germinated seedlinF, This suSSeststhat selection is operating during the life rycle. Heterozygotes may be rnore likely to survive at all stages from fertilisation onward; through seed rnaturatio& germinatio& early seedling Srowth and as the plant matures. Similar processes are thought to occur in several other eucalypts that have been investigated (Phillips and Brown 1977;Moran and Brown 1980; Hopper and Moran 1981; Fripp 1982; Sampson el al. 1988) and in other trees (Iedig 1986).

23.2. The distribution of genetic diversity within and between remnant stands

Reviews of the relationships between ecological and life history traits and the Senetic structures of plant populations (Hamrick et al. 1979iLoveless and Hamrick 1984;Karron 1987)indicate that tfrere may be some maior determinants of the distribution of genetic diversity within and between populations. In particular, the importance of the breeding system in determining PoPulation structure is widely recognised (Jain 1975; Brown 1978, 1979;Hamrick et al. 1979; Gottlieb 1981; Hamrick 1983; Loveless and Hamrick 1984). In general, ecological and life history traits which promote outbreeding and gene flow are associated with higher levels of intrapopulational diversity and lower differentiation of populations.

11 ZI

'spuBls '1vtol uaartlaq dllsra,rrp aql Jo Iaâl aql JIPrIlnoqe se^r lr'salJads raqlo uI sv 'luaraJJrp se.rr uopepuara;Jrp ro laâl aqt q3noq11e dpuelryu8rs a;a.n daq} ler{l paleJlpul (l Z '3rg;1.rede ru g91 dlaluulxordde suopelndodqls olllutpoqt g d.rurlrqre o^{l uaaataq pu€ ulqly{ 'suorsrltpqns dlrsra,rrp Jpauag Jo uoqnqulsrp aql yo uosrreduroJ uollulndod uaa.rlaq ler{1 aJl.l{l dlaleurlxordde se.n suoylelndod uaa^ ]aq uo-ruodord aql ler{l puno, (7861){)lrureH pue ssalaô'I "4ralâl p u1 suonelndod lue1d urqly"r ap)s leJol E uo Jn)Jo osle deur uoqequaral;lp JqauaD

'u.ralled anrl aql palJolslp a^eq deut dlrsrarrrp;o slalal pu€ arnparord Sugduresarp

aJu\st\lutpoTt 3 ul aJuetsrpJrlauaS uo paseqsdlqsuonelar 3u11ardra1u1uaq.\r paslJJaxaaq plnor{s 'spuBls uollnpJ ruaqlnos aql u.rojj Z pue I spuEls ,o uoll€.rPdasaql uroJJuaas aq UPJse aJuElslp '(LL6I lzcrqde.rSoa8qll,lr pasearJuraruelslp rpauaS upau prauag q qallDog) sanads lue1d raqlo 17 yo suopepdod.rol palrodar lpql ol r€llurls se^r (I90'0=O) aJuelsrpJllauag Jo aleurqsauParu 3qI .(g.z .8lc) sarnsearuaruelsrp rrlauaS dq pale.qsnlI osle se.{ uo!}epuaraBlp ^rol lnq }uPJIJIuSls

'slueuurar dep luasa.rdaq1 ueql palelosl ssol ala^.r '1eql €ugeap o1 roud suollelndod uaa.tlaq îog auaS,o slaâl qgrq dla,rqelar ol palnql4+Paq treur 'e '(1861 lr'fl4ryrpo1t uI raddoH pue uPtor{ 198613rpa1; slddlerna Surpnpur'sapads aarl lsour ur pa^rasqo u:a11edaql sl snlJ r*ol d1a,rqela:lnq lupJlJlugls se*\ f,qiuvpoqt g Jo spuels luzuruar '€'z aqt uaaataq uopelluaraJJlp aqJ a1qel u1 uaarS are s1d.,{prnaraqlo lPlaâs 1o suogelndod puu oqluopo4t g Jo spupls uaa^ltaq puB urqll^ d11sra.r-rpJo uognqlrlslp aql 8ul^roqs salPu4lsg

'sluplrual Jalleurs aql ul /r{ou aua8 parnpa.r pue ryrrp €urpaetqur Jo lJPdurl ar$ auo)Jaô '1nq o1 q8noua aq 1ou deur asaql suraas11 filsraarp alouord qJnI1'âôqe polsrT sJllsPalJeJBqJ aAPq '(vt'z saop tqluapo\r'? uollJas aas) s€llsru palPlosl Jo puels lFrus P ul punoJ se.^{Sulpaarqur ur aspalJul luear;yu3rsp pup azls uollelndod puP fllsra^lp rnauaS;o saJnseauraruos uaa.,',rlaq 'luaredde suo4elaJJoJlea.u ara^r araql aJa^ azls uouelndod IIPus pue uo11e1uar'u3e:;to s]JaJJa 'sarcads arg'ra1a^roH asaql ul dlrsrallp Jo sJueualur€r.rrar{l ol SupnqrJluoJ srolJeJ se palsa?flns 'suollplnu ara.^4. Jo uollplnrrnJJe aql pu€ dllauaSoralaq lulodural pue IePPds qll^{ palelJosss 'q8rq 'd1ua3uol pup salo8dzoJaloq SuunoAPJ uoqJalas 'stualsds Surleur paxrq a.ra.u uraql

Jo IIe ul punoJ dlrs.rarrrpJpauaS Jo slalal aBBJaaeaql lnq azls ur a3wet v1\utpo1t'e Jo sluPu\uall

',u.og 'lJlrp aua8 ;o qurulsuoJ aql puP uoqJalas ol sasuodsar 'GL6I'lt ,(repuoras are 1ro1rlqdroru,{lod Jo raqunu aql uI uol}Jnpar pue saIaIF ro ssol arIJ t, raN) lJauaploq p raue dlplder sasearJulazls uopelndod JI [Prus aq deul dtrso8,{zoralaqJo ssol aql qSnoqlp (916I ulp{) ,{lrsoSdzo.rapqur uoprnpar P pue suonelndod ulqlu Sulpaarqul pasearJul 'dllPJllsJoaql 'suo11u1ndod o1 peal deur suollulndod llprus olur sapads 1o uorleluaurSer; aql pue Jo azrs pup uollnqFlslp ar{l dq PaIJaJJPs1 dllsraarp Jllaueg }o uollnqglslp la^al aql Table 23 Total genetic diversity anil the diskibution of diversit5r rvithin and between populations of trees of wid€sprea4 regional and localised geographic distributions Hn total genetic diversity. Ilg , mean gmetic diversity within populatioru;. DS? mean genetic diversity between populations. Gg7, mean proportion of diversity between populations.

No. of Species populations H7 Hs Gsr (%l Sourcea

WIDESPREAD Pinuspoulrosa Engelm, 11 0.289 0.2& 1.5 I Pinus ruilialz D. Don 5 o.lt7 0.098 16.2 2 E. granilb t2 0.190 0.767 12.o

E. saligna 7 0.260 0239 8.0 J Eucalyptuscloeziana F. Muell. 17 0.270 0.240 11.0 4 Casuolina cunninghamiana m 0.292 0208 28.7

REGIONAL

Eucalyptus caesiaBenth. 13 o.t76 0.068 6r.4 J E, oucis 10 0.320 0222 24.4 6 E, Iane-poolei 7 0.324 o272 13.7 Eucalwtus dioercicolor F. Muell. 13 0.299 0270 9' 8 E, rhoilantha 5 o25s o2/r 10.1

LOCAIlSED Eucalyptts lateitica Brooket & Hopper 0.318 0.278 12.6 Eucalyptuspanilens Brooket 0.r70 0.155 8.2 Eucalyptus johnsonianaBrooker & Hopper 0.139 0.084 39.6 Eucalyptus sabaecBrooker & Hopper 0.t97 0.170 13.7 aSowcesof data: (1) Flamrick (1983),(2) Moran and Hopper (1982),(3) Burgessand Bell (1983), (4) Turnbull (1980),(5) Moran and Hopper (1987),(6) Sampsona al. (19SS),(D Sampson(1988), (8) Coates(personal communication).

l5 VI

.^rou sanads p'q;o:noppqaq pue Jaqunu aqJ aua8 puorlelndod.ralur.roy saqrungoddo Burprlord 'sporlad pa8uolord roy suolleunsapu^louTun ol spuplsJo lno paôur auos pue spupls uaa,+rlaqqsnq paleapun .(6961 Jaô ^{aU oslp sp{g aaNJW) s1ue1dzua; e uaa^ laq paleJluaJuoJaJa^{ sluauraôru pue sarrolrJ.ralpaqsllqelsa ,lsEJluoJ .ramou sanads auros uI lxau aql ol ur 99 o1 dn Bur,(g uaql pue ua11odSuqxlor pue srar'log uo Surpaaypaa.rasqo a.ra,u ua11od Dqruopoqt.g parr.rer 1eq1 sapadg 'sp;1q dq saJuplsrp a3.re1:aao palrodsuerl aq,(eur ua11odb1luzpotlt.g .srolBurrlod d1a111gsoru .y,1 aql paraprsuoJ anrl' nln8wll pue sunsa a-7 q8noqlp ragod uq1urpo4t.X Attet,sryw1sot.1 'urnssod dauoq aql pue sarlads prlq praâs tpr.0palp4snll (9961)aaNrl l ;o suorle8qsarrurar.la

'19961uosduregy slood uaylod uaamlaqdlrs_raarp Jo lunoup aql Burler,uqsa,(q pue (Sg6t) upllpls yo poqlaur aq1Bursn apa.u uaaq .19961 aaeq .,uogauaS yo salpuqsa lJarrpul uosdrue5), pue g salrsur salalp a^gJutlsrp suralled psradsrp aql Sululurpxa dq pue ;o 19961aaNJJ I) sluarua^our rolBurtlod ro suollE^Jasqo urorJ paulelqo uaaq .urelqo aleq oqluopoUt.Aur ^{oU aua8;o guaurarnspaur lJaJIq ol llnJgrrp a.resuoqelndod lprnlpu uqlr^r 1es:ads1paua8 pue uaa^rlaq ^ oU aua8 ,o salpu4sa a^qelgupn$

^r.oIJauac .9.€.2

.(gg6l) .suoqeJol uosdups laljv puets roj I.Z alqpJ aas aqluppoqt.x spupls to lupuruar arp uaamlaq sd1qsuo11e1ararg fq,uoqs (SZOIIaN aO) sarnsparu aJuEls.rpcllaua8 ;o srs.r(pueralsnp 17t IO.I1 uo pasuq ruer8orpuag 9.7 am8lg

esuslstpi[eue5

z0'0 ts0'0 90'0 I0'0 0 1'0

a E I v

l "_L: I

of Figure 2.7 Diagrammatic representation of E. thoilantha at stand 4 showing the position subpopulations 1 (wes0 and 2 (east) and Are area bumt in APdl 1986 14th' 1985' ( . ) Sampled plant, ( ,,! ), Other E ' thoitantha- (8 ), E' rhoilantha burnt APril ( - ), Fence. The scale is aPProimate.

found in standsvaried betweenyears, aPParently because of the number of flolvers and availability of nectar.

Seedmorphology and observationsof seeddispersat also suggesta Potential for gene flow over relatively large distances. seedsare winged and are blown readily through the Patchy vegetation by strong prevailing winds.

Estimatesof gene dispersal from the distribution of rare non-Il|atemal alleles ranged htween approximately8 and 185rn (mean57 m, s.e.26.62',).Comparison of adult Plant and Pollen Pool Sene

15 9I '199611uosdure5 raryy l 96'0 f,LSADJ'7 t9'0 qJnfi'7 8t'z npod-aun1'X 0I< nqluaPo4t'x

tsa(uN) sar)ads

azls aldures roJ paparror lN = lsa(uN) qddlurna pa1nq14qpd1euol8ar Jo sapads r:lqp'oprrr- uE ropory'r? Jo spuqs lusuuar uaa^tlaq paSuEr{JXe (urN) querSlru Jo rJqrunu u"aur arp Jo salerrrps:l t'z aIqEI

'Upp rpauag Jo tlnsal p se luaraJJlp dlplpua8 au.roraqpuP urroJ spootpnoqqSlau IpJol tpr{l os palJlrlsar Sulaq .nog auaB uorlelndod-ulq}la }o dlrpqrssod aql a}eulrulla 's1ue1d lou saop suollelndod uea^,rlaq^4oU auag q8rq suraes11 ul lensnun lou seaau:a11ed srql 1eq1 'pallurll punoJ (t861) {JurxpH pu? ssalalo'l'^ allar € uI ,(lluaredde s1suorlelndod u!ql!^t ..',rou aua8 /rala^roH 'zuouepdod;o uoppquaraJJlpaql Sulplqar aJro; allsar{or apln-salJads lueuodurl ue pup SuFpalJ o1 roFd q8rq se^r suollplndod uaa^rtaq ,r,'og aua8 lPql lsa8Sns rryuapoy'7 u1 suogelndod uaa^{laq ^ oU auag Jo saleurgsapue dlrsralrp tqaua8;o sdaarns tuor; aJuap! a aql 'slpsar Suprlguor a,r13o1 readde suollelndod uaa.+rpqpue unn1lr ,ra.ogaua8 3o suo11e311saau1

'@86t'lt t, uosJrl) srpueudp uollelndod lua.unr aq1ol upql a8ueq)xa aua8 ;o su.ragedIPJIJolsl{ ol luP^alaJ aJour uoFpruJoJu! upluoJ poqlaur sltll Iq palPlnJIPJ sanlP^ asneDq pautqqo aq d€tu saleuqsa q8gl 'lJullxa ^{ou a:e 1eq1suollelndod auos ;o azls aql ueql ssal dlqeqord lnq slueuruar luasard ;o azls aql ueql ra3re1 puP sldfleJna ,o sarJads pslnqFlslp dlpuo13a; raqlo aarql roJ apPru salBulgsa .|rou '(t'z aua8 leuollqndodralu! upr{l ralear8 apnl$8eur Jo srapro o/vr}ro auo sE^{lI alqPJ) qg1{ ifta^ 'exel se^r poqlaru s1q1uodn paseqnlluupoqt 'g uI rr{og auaS;o aleurpsa aqJ uaa^{laq drualslsuoo auos sI arar{l pue apn1ru8eu Jo srapJo arou Jo o^ l ,{q Ja;;rp deru suorlelndod prnleu u1 ^rog auag '^{oU aJuls role)Ipul ue sapr.ro:d1r 1nq aspard ruo:y ret sl poqlau aqJ auag Jo sla^al qBJrtr1sa33ns '^rou 'uoqurauaS 0'I uer{l JaSJPIsalPugsa seaJsr{.^{ auaS ,rao1aleJlpur 0'I uer{l ssal sslErul}sg rad stu"$ltu Jo raqrunu uparu aql sp ^rog aua8 Suneurnsa ;o poqFur u padola,rap (9861) uDIlelS

'suopelndodqns 'pa11u41ser"r'gede ru gg1Ipo oaalaql uaaMlaquauod ,o luaura^ourlBr0 palsaSSns(tZ'31g'!V 'oN) puels parpalJun uB uI suorlelndodqnsdre.qrqre o^ l uaar\laq sapuanbarl 23.4. Outcrossing rates in natural and disturbed stands

Outcrossing rates in plant populations are not fixed but are affected by several genetic and ecological factors including the size and density of populations and the availability and behaviour of pollinators. Theoretically a reduction in population size will be associated with an increase in inbreeding and reduction in fitness.

There was a substantial increase in inbreeding in E. rhodanthain a small remnant of 14 plants isolated in land cleared for agriculture (Stand 34 Fig.2.4; Sampson1988). The level of outcrossing recorded in this stand (?=0.25) was significantly lower than that determined for an uncleared stand and substantially lower than rates reported for any other eucalypt (see Table 2.2). It was attributed to a substantial increase in self-pollination, probably as a result of reduced plant and Pollinators numks.

The increase in inbreeding was not associated with a decrease in reProductive outPut as determined by the number of capsules produced per plant in one season,but these investigations were very limited. Observations suggest that the smatl remnants of E. rhodanthaare less fecund than the larger populations and the plants appear less vigorous. This may be simply the result of environmental degradation. Ily'hether increased inbreeding is associated with decreased fitness in E. rhoilantha is a subiect that requires further investigation.

2.3,5. Fecundity

Plants in distubed and relatively undisturbed stands maintained moderate levels of seed set

(Table 2.5). The number of fruit set Per year was low when compared with the malority of eucalypts which produce numerous, usually thousands, of small flowers and fruits. This reflects the different flowering strategy of E. rhodantha which produces a few large flowers with large nectaries in each flowering season. The number of flowers and hence the number of capsules may vary substantially between years.

There were significant differences between plants for the number of seeds per fruit and the mean height of seedlings at eight weeks (Table 2.5). When within-plant variation was taken into account, no significant differences were found between plants or between stands for any of the other characters assessed(Tables 2.5 and 2.6).

77 8t

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199611uosdure5 .ragy

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oz sz /.€ N 9[-0 €I{ t}{) a3ue.r (6r) r'9 (8'r) 6'€ (Er)€z uearu 'oN g986I roJ fuqd/ras SIIruJ

LT, N t8{) a8uer su (6'|i)r'9r IIE III 'oN Vga5I roJluqdnas qTn{I

7 uoneJndodqns 1 uoguJndodqns t alls t aIs € alls

'poroJslou'su 'sasaqlraredur sro.rlapJepuuls .196l .qal uI '986l 'JaO .suopeool .3r.g palunoD g ur palmof, o uonqndodqns pu" prsrs roJ l.Z pue 1.7a1qel aag t pusls uI -3 suopqndodqns on1 uI pue € puEts tE qwlil ttllwporp yo uoganpord paaspra apsdu3 SZ ans; Table 2.5 Mean seed and seedling draracteristics scored in the progeny of ten E. rhoilantha plartls ftom Stand 3 and from each of two subpopulations in Stand 4 Values shown are the means of two replicates. See Table 2.1 and Fig. 2.7 for stand and subpopulation locatioru.

Germination Final Seedling alive at Height at at 7 days (7o) germination (%) 8 weeks (7,) 8 weeks (mm)

Stand 3

Mean IJ.J 47 42.75 54.97 Range 0-27.5 30-77.7 30-77.7 36.7-68.7

Subpopulation1, stand4 Mean 10.5 54.5 46.25 57.42

Range 2.5-35 17.5-80 tJ-oz.J 55.0-75.0

Subpopulation2, stand4

Mean zo 72.8 63 JZ.3

Range o47.5 57.5-82.5 50-72..5 JO.l-OO.Z

After Sampson(1988).

plants indicated that more than half of the fruit on a plant could open during a single year, with the highest opening rates occurring between October and December. Despite this supply of seed, natural seedling recruitment has not been reported tn E. rhoilantha. A limited investigation of the effects of fire on seedling recruitment and vegetative regeneration was conducted by Sampson (1988). Following a moderate April burn, 29 of 30 plants began coppicing at the onset of the first growing season following the fire (November 1985) and some flowered during the 1988 season, two years after the fire (McNee personal communication). No significant damage to or loss of the new vegetatfue growth was observed. Ttris was attributed to the exclusion of sheep since rabbits and kangaroos still had access to the area. New shoots on plants in paddocks where stock have been grazing are usually stripped of leaves.

The fire was not followed by seedling recruitment. A substantial quantity of seed held in the canopy (=27 000 seeds) was released following the fire but only six seedlings were observed. None of

10 UC

'9961 '? aun{ Suunp uUtuupo4r Jo pate lunq e ur palueld ara.r,rs8u11paas aq1 slo4uoJ paJuaJlm qly paredruor sarnsolouaparuay u1 Supprrns sSulpaas flUlutpoqt.g to a8eluacraduearu aql g'7 am8lg

(r0uourpue read)aurrJ

L86L 986I v I I r/{ v II J I a N o s vl--T t[

('I

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"'S 08

.slerpenb palnquluor ,{1qeqo.rdsquar.unu pue arnlsrotu pos ;o flddns aq1;o uonelrurrl paJual ulgll^ .ralear3,{11epue1sqns lnq'luepunqe sen ' srtbllrdn.4 pue wnsoutds.g,.dds 'sanads ,1 'l f,floqlutcl dlureru a^peu Iera as pue saplouatfiuomuqtn prm DFpuaIoJ oxa\ptr.tv ^1nq{ 'sarceds waay ,llureur paam go qlaaorS aql pup papaa^.r.lou aJa r slerppn$ .s8u;1paasaaa; 'paarasqo ,suals ,salerqalraa € uo palrasqo sp^ a8eurep pazu1 rourl I lou sp^r ualorq se qans a3.reyo1 alqqnquue uaaq a^pq lqgnu lEql a8purpc '(g'Z .3rg :99.6<4,1sa1-1;sSurlpaas palueld yo d111e1rou aq] uo patJa lueayuSrs ou ppq sal"rqal.E ^ a8rel dq SurzerS og anp a8etuup Jo uoqpuJurlla aqtr

'DUlurpotlt g Jo pupls lumq aql ul uoneunula8 SuqurI rope; rofeur aq1 dlquqo:d s?i!t pup'putJ]r pue s:olepard dq a4J ar{t raup paas aql ;o uorlaldap 'lurnq prder aql ol pal sI{I sp,raEaJe l1erus e dluo asneraq arIJ aql 8uv{o o, palelles lou ara^a slue se qJns sJolEpa:d paas 1eql palsaggns (886I) uosdureg 7961 1sn8ny ur aarTesP,u s8uqpaas asaql to the mortality of E.rhoilanthaseedlings planted in quadrats. It was undear why there was no increasein the mortality over thesummer period @ecemberto Febnuary)whm melrn temperatures are high and rainfall negligible.

Seedlingrecruitrnent follows only a small proportion of fires in mallee eucalypt populations and occursas a result of the coincidenceof both fire and envircnmentalconditions suitable for seedling establishmentWe[ington and Noble 1985a). Observationsin the small burnt site of E. rhoilantha demonstrad that fire may have an important role in the reproductive biology of E. rftod.antha by stimulating the releaseof large quantities of seedfrom the canopy. Similar suggestionshave been made for many oth€f,sderophyllous plants (Gill 1981). It seemsreasonable to suggestthat, prior to the dearing of vegetation, lrore extensive periodic fires in E. rhoiltntha populations may have stimulated the release of seed and satiad predators. lvhen other conditions were suitable, recruitrent would then ocrur.

2l a(. I suoqdo luaura8eueu pup suoqpJaprsuor:gaua8 aq1;o auos paquJsapa^€q (986I) u^{org pue 33ap 'suollepdod;o 'alqrssod uolt€llllqPqar aql roJ Ial)nu sPpasn aq plno) spuels palPlosr prus aq deu-r puel 1e.rn11nlr8eSurpunorrns aql ol salnqule alqerlsap u-{€ual ro uorlerolsal aql lo uollplllqpqaJ 'uopepergap 1nq alqrssod aq 1ou deru rualsdsora 1eur8:ro aqt 1o dra,rora5 r1aq1 lua,ra.rd o1 luaura8eueuraruos alnbar 11r.radaql salpJrpul DqlurJpo1tA Jo slupuuJal aql Jo ssaufleursaqJ

'puel prntlnrrrSe paftap ul spupts IIErusraqlo aql ur q8rq ,{lqeqord oslp aJB slalal Surpaarqul azls uollelndod alqeh rumururur aql ^^olaq sr puels s1{l sa}pJlpul

1s1ueld711 pg puets Ilpurs aqt u! punot Sulpaa.rqur1o 1a,ra1qSrq aqJ uleuaJun sl salJads sHl 'alqrssod ;o lepualod ur.la13uo1 aq1 ra8uol ou s1suollelndod luelxa ruo{ ^rou auaSSurnuquo: a:urs 'JaAa.+roH'ruJal uorls aql u! lspal 1e'sapads sn{l JoJ azrs uouelndod alqen lunrurunu aql a^oqp 't '? dlqeqo,rd a.re1s1ue1d 09t'9 puels puu slue1d081 puuls) tqtuopotlt Jo slupuruar paqrnlsrpun 'lplllJns dlaapelar oml ar{I ro; uralqord p aruoJaq plno^r uoqerre^ Jo ssol pue Surpaarqur rpnl.4r ,+rolaqsuoqeyndod;o aJnlJnJlspue azls aql luasardar ol lulod^ al^ llaua8 e u:org palpurrlsa 'Uodal 'xalduor ale sdAI I slql u1 sI (dAI I) sazrsuollelndod alqPr^ rumururu Jo uoltprurtsE

'pauluralap aq o] spaau palrasuoJ aq o1 suogelndod;o azrsa^llJa;Ja alqerl urnurrulrr pup raqurnu aql qloq'{1sraalp a^lasuoJ ol Japro uI 'snr.{J 'suopelndod Suoure ,uog aua8 alqeua pue 9urpaa,rqurur saspaJJut 'suoqelndod Ielluelsqns luaaard paq.rnlslpun ur punoJ uralsds Suqeur aql ulpluleu lp^r lErll Ia^al p lp sJaqunu uopelndod Sururelurerudq dlrsra,rrp alJasuoJ ol aq uaql pfrcqs otljuvpol,lt J 'qoeordde ro; d8alerls uorlplJasuoJ alerrdordde aq1 yeraua8 slql lroddns uorlaas snol,ra:d 'suortplndod aql ul paqFJsap t\tu,po\r'3 ,o suopPSpsa^ul ar{J Suoure r"rog auaS alqpua pue 's.raqrunuuorlelndod Sururelureurdq dllsoSr(zoralaqaloruord o1pue SurpaaJqulaJnpal o1seu. s1ue1d Surpaarqlnoyo uopelrasuoJ aW ro, d8ale-4sluaula8uueur snor^qo ar{t l€q} papnpuor (9861)Slpa-l

'(9861 'Jolraqurrs pue 9lnos) arueuodur! augd;o sI salrasar uorlplJasuoJ ur slelrqer{ alqellns Jo uopealasald pue uolspo.rd 'suo11e1ndod aq1 1ue1dyo 8up1as pr18o1ora aql uorJ paJJo lp aq louueJ suorleraprsuoJ asaql 'Ja 'dlrsraarp a^ioH prrSoloqdroru pue arru)nrls rqauaS uogepdod ,sapads aq1;o uollepa.rdde ue sarpbar sapads 1ue1d;o luar.ua8eueu pue uoJle Jasuocaq1 roy sa13a1er1sptugdo 1o u8rsap aq1

f8aleqs p.rauag '1'6

'o4luuporlt 7 ;o suorlelndod Jo plr.r aql ur lp^llJns panupuoJ ar{l aJupqua puu aJnsua o1 s1luarua8eueur 1o a,rrlralqo aq1

I|c EDVI{VY\I 'S in the founding of plant populations. The information on the genetic diversity within and between stands and the mating system presented in this report provides a background on which decisions about founding and rehabilitation of populations may be based.

lf E. rhoitantha were still distributed as it was before European settlement, the appropriate conservation strategy would be to have several reserves containint entire PoPulations scattered throughout the range of the species. This would conserve the range of morphological and genetic diversity, maintain levels of gene flow and cohesion in the species gene pool, and maintain habitats to conserve the flora and faunal diversity that contribute to community stability. This strategy can be modified and, with appropriate management, the remaining stands of E. rhodantha can provide a viable system of populations to ensure and enhance the survival of this rare flora.

Researchhas indicated that stands of E. rhodanthamay be subdivided into genetically different neighbourhoods and that gene flow within populations is limited. When populations are subdivided in this way, it is important to preserve as much of the entire population as possible because the loss of a portion of it may be as critical in a genetic senseas loss of the entire Population (Hamrick 1983). Therefore,entire populatio^s of E. rhodanthashould be conserved where possible and the larger, fecund populations should take priority (i. e., Stands 4 and 5, 1 and 2).

Recommendations for conservation should not be based solely on genetic data. Ideally they should include information from morphological and physiological studies (Hamrick 1983). Strategiesto conserve E. rhodanthashould therefore include both of its morphologically different varieties. The investigations of the responseoI E. rhodanthato fire and field observations provide limited but useful physiological information.

One approach which attempts to use genetic information to conserve variation is to maimise the conservation of locatly-cornrnon alleles which are presumably maintained by selection. The Pattern of distribution of this type of allele in E. rhodanthais artificial becauseits stands are remnants but all the efsting locally-common alleles could be conserved by preserving Stands 1,4 and 5. Another approach is to select populations on the basis of genetic distance. This is not advised for E. rhodanthasince estimatesof genetic distance for it may be biased and relationshiPsunreliable.

3e Managem€nt actions

Kelly et al. (unpublished report) have described in detail the allocation of responsibility for rare flora within the Department of CALM. In general, the Provision of advice on management prescriptions is the responsibility of Flora Conservation Research Program staff and the tz

laleayd; €g puers (r) ls8uudg aerql;o a4r{S)Z puE I spuErs (q) (alP Ird) 9 pue t spuPrs (E) :uoprsmbre.ro; AFolrd Jo JaprouI spusfsJo Suqs{ e sI 3w^ .olloJarll 'uoqruExa uappnsJo Isu aqr Su.Dnpar (p) puP 'lueuurar Sunsrxaaql plmorE peqsllqqsaSqaq s,uopqndodir,rau yo pooq11a1[eqr (J) 'saIaIIe

uouuror-,ql.pJolpue &Isa Ip IEDISoIoqdrourpuE rFaua8 Jo aSuerer{l SuI rasuot (q) 'lEllqeq qI pue lusuurar arluo 6{Pnb Pu? a^s ar{l (E) lq paupnrapp aq pFor{s {uolrd uaql elqlssodlou $ srq 'slupuural 'essqrmd JI IIe aNasuoJol aq plnoit^ d3a1e4spapl arp'queuuar ^aalos arE araql aJuls ro agusrfJxe'uoDEuop ,{q ,raq11a'srr|'JJo tUlrbpaqt 'e rIJIrIr uo puel arJnbru pFor{s luatuuPdad aql lEql elqensap dHSIq q lI 'puq parE)Ptm uo ulsutar spuqs o,^,rldluo pue eunPJpue Erou '3 ,o uoqe^rasuoJarD roj pa^rasaruaaq sBqrpFllr^ puel vo tllutpoqt ;o suauroadsou aje arar{I

uoElslnbr€ puBl 'z'e6

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.JIqs ,s.Eu]r CpU I pue sa4nls opupl Jo uonurado.ot aq1 qlpr uopUapun 'anupuol aq plnoqs suopre luarua9eueur 1JV plnoqs sIrD leql ppuassa pup alqp sap qloq 'sraun sI u (quw) tuaucrdac spEou ulelt aqt puB sar.rqs opuq aleaud ;o acueprsse puE Ilr/r^poo3 'lq8r$roJ 'g aql q8norql ,fta8,re1 pa,ragpe uaaq ssq tqfitpollt Jo uorlE rasuot ar{l r ou Ipun

Itr,qs puE sJitul opuq rI|I/t uosr-El1 'I'z't

'JJEls uoperlEulrupv pue.1 pue aJIIpIr.M pue rlJuprg uopJalord aJIIpIIM leuo€aU Jo rqlllq-rsuodsar 'sreur\ eql sI suoFElndd EJoH eJsf, parBIJt Jo sJauinoPu€I Jo uopu)lJpou para IaPPuEf opuel Jo irouuJlJllou roJ lqualEru arEdard pue snlqs s,puel ar0 arlllnJaFp ol lauuosJitd uopErlsTurupv 'Paterol 'pa,rpba,r puE-l puE eJIIpIIM Jo AlKlsuodsor ar[| q lI a,re suogelndod arau ;1 'sa4l'o sE palEpdn aq pFoqs ralsrSar aul l44slc puP lguol8au ur Drold arsu F$uapIJuoJ 'g Jo slsl uo par4slu.reur aq ppoqs ,tllutpolt 1o suollupdod urttou:l Jo slplap 4IIEJoI aslJard

'aapruruof, uoplslnbry spurl uope,r.rasuo3 aq11o,$grqrsuodsar ar$ sI puel Jo uoplsJnbogl 'srar.rrJo aql puog8ar arll Jo $Jllqlsuodsar aqt s! prog arEr ;o uogralord puu luaua8eunu *lr' (d) Stand 3 (entire stand private) (e) Stand 5 (Shire of Moora) (f) Stand 7 (private)

Investigations sugtest that about 25 plants spread over about 4 ha of natural habitat is above the effective population size for E. thoilantha (Sarnpson 1988). However, although most pollen dispersal may occur within neighbourhoods, neighbourhoods are not isolated. Gene flow within and between stands is an important component of the genetic systern. Therefore, a minimum viable population should consist of more than one neighbourhood (subpopulation) of about 25 plants.

It is very important to maintain populations of pollinators. Birds forage on a variety of species and need species other than E. rhodantha for nesting. To ensure and enhance the survival of E. rhoilantha, it is essential to provide and maintain areas containing a suitable variety of species and habitats.

Therefore, 35 ha is the minimum recommended reserve area for viable populations of

E. rhoilantha. There are two remnant stands in uncleared areas treater than this size (Stands 4 and 5) and they are the highest priority for conservation. Smaller remnants in suitable habitat may still be useful as long as they are above the estimated effective populadon size of 25 plants. These stands may fulfil the requirement of msuring and enhancing the survival of E. rhodantha over the short term, but their value for long-term conservation is reduced.

3.2.3. Pntection ftom grazing

Plants that are to be conserved requirc protection from grazing. This may be achieved either by fencing or by agreement with landowners to exclude stock. Evidence suggests that.-it is not necessary to exclude rabbits to protect E. rhodantha from grazing, however, the rabbit population should be controlled to prevent damage to the habitat. Part of Stand 4 has been adequately fenced by the landowner. Complete fencing of Stands 4 and 5 and the associated habitat should be the highest priority. This may conflict with the landowners requirements for grazing and shelter belts for stock. It is therefore essmtial that managemmt actions include discussion with the landowners about the position of fences and provision of alternative shelter belts.

3.2.4. Protection from accidental destruction

Stands of E. rhodaztha should be protected from accidental destruction or damage by bulldozing, off-road vehicles, rubbish dumping, spraying of potentially damaging herbicides and recreational land use.

E 9Z

'a3ero1spaas uual-9uo1 pue uope^Flnr qSnorql paaaqae aq deu'11 'aseaslp ro sarr' sp qrns saqdo.rlseler q8notql suogJupxauappns Jo Arlqssod atp aJnpar oI m1uopoy.X ro1alqesl^pe q uopelrasuof, nlrs rg

uogE^.ersuoJ4rs 1I 'g'z'e

.groda.r .12 paqqlqndun p d1a;1 uralsds sr.qtldope ol 'sa pa8anorua uaeqaaeq saqqs leJo.l rasar peor uo sea.reluerguSrs dlvluauruorraue SupE rpruap :o; rualsds Suqruru plag e pedolaaap ssrl q1dl I aqg .3u11ruurpeau os1e$luarqarlJ Suop s1ue14 'que1d asaql ro; uogaalo.rdtunrupr.ur e apraord ppon 3upgeur Jeaun .suonpradoacueualupur ,Z ,I ,.{qa8uruep o1 pa{.qns a're pue sp€oruo palerol a.resuau.oads palelosr a,uos pue 9 pue e spusls

8u11reurreau-r1 .4.7.6

'sua.reuoperaua8ar .soN papoaspa.Ilp pup lprqeu r4 spoarnJo lo4uor arll uo uonzurroyurapprord gg-z pup lg-l q€r{s 'sapads uoqeruroJul I I-lV) Jo luar4redaq a,rpeu raqlo ro nluepotp'g a8eurep,(eur sapnlq.raq a^IlJalas Jo asn araqru aJueqrqslp IIos runurunu qlli puBq dq pa,rouar aq plnoqs spasM

's.nu,raopuq , aluaud pue CUW p.reog uouaagor4 a,rqpr1.r8y 'pare aqt aq1 u1 asg.radxa Wl\{ JJels nnvJ Jo luau.4.redaq rD!\{ asl€Il ppoqs luaugedap aq1 'spaa,r,r Jo sroJIrJO dq paptmorrns aroprar{l are pue ptrq p.rnlprlr8u ur aJE spus}s }so14[ .alqBrlsap 'X s\ ryflryoU lo uoEE rasu@ JoJ po :Esard aq o1 are ;uq1 spuep reau ro ul spaa{. Jo loquoJ aqtr

Io4uor paaM .g.z.g

'seere3u,rprmo.r.ms ur uopJnpar lanJ Jo $Iearqarg Jo uopJn4suor,{q sarg pallorluoJrm ruo.g paDaloJd pue sutnq paq;nsard uro.r;papnprc eqp1rcqs rqrurw 'g 'paFnuuroJor lott s\ oq&rpopt.X yo uope.nuaSar aartela8aaJo lrr:)rrrllrutar Sulpeas ;o uopotuo.rdaql JoJIool luaura3eueu e sE anJ Jo asn aqr

e4t l!o.g uoptaprd 'g'z'e

.(.62.9uorpas eas) uoFEIIr.qEqarpue arnsop .roy,Q1.rq.rd lsaq8g aW q Z pusls to $rou 1;d p,rur8 aq1 .palellllqeqer puBI aql pup paleu.ural ro palue^ard aq pFoqs spue5 uFRIir ,ro o1 gaoelpe sue,re14 3up4141 Seed from cultivated plants could be supplied to nurseries since the specieshas ornamental value. This may have the added benefit of reducing pressure on natural populations from illegal seed harvesters.

Seed from artificial plantations should not be used to re-establish or found new PoPulations unless seed from wild populations is not available'

Seed from wild populations should be collected and placed in long term storage both as a precaution against sudden extinction and to conserve potentially important economic genetic materials. This seed should be representativeof the range of genetic and morphological diversity and sampling should include populations with locally-common alleles (Stands 1, 4 and 5). In practice, all stands of E. rhodantha can be sampled with little effort. Within stands, sampling should be random. Information about the status of seed collections should be made available to field officers to ensure samples are rePresentative and to Prevent unnecessary re-sampling.

3.2.9. Rehabilitation of existing stands

Rehabilitation of the smaller existing stands is required to fulfil the oblective of management. It should be done in co-operation with landowners, Shires and the MRD. Officers are referred to the Department of CALM Information SheetsNos. 5-87 and 2€8, and Edmiston (1987) for advice on methods of rehabilitation.

The number of E. rhodanthaplants in smaller stands that are to be conservedshould be increased to at least 25. Reestablishment of other species,particularly those exploited by pollinators, is also essential. Ideally, artificial populations formed around the nuclei of remnant stands should form subpopulations of related individuals and levels of genetic diversity should be maintained. Seeds for re-planting should therefore come from existing plants. This may produce a founder effect so that the new population is not representativeof the original population but complete elimination of genetic variation is rare even when the number of founders is small. If sufficient seed is not available from the stand being rehabilitated, then it should be obtained from the nearest population.

E. rhodanthaseed germinates easily with the addition of water' Approximately 30 per cent of 6- week-old seedlings planted into the field in June were surviving 15 months later. Improved survivorship may be obtained by planting older seedlings with deeper root systems or by watering seedlings throughout the first summer. Investigations indicate herbivory by rabbits, kangaroos or insects is not a significant causeof seedling mortality and therefore it is unlikely seedlings would need protection from thesefactors.

27 8Z

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IrEqaIq €I'z'€

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tu,uo4uo141 '71'7'9

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aa,o1yaua8 pp1;prv .II.Z.S

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suonelndod -,r,rauSurqqqplsg .0I.2.€

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. Infor.rnation on the locations of Phytophthora infections should be regularly updated and the management actions required to prevmt the infection of E. rhoilantha consewation areas should remain under revision. It is essential lhat P, cinnamomi and other pathogenic Phytophthora species be elduded not only from areas in which E. rftodantfta occurs but also from the surrounding communities which contribute to the support of pollinators.

Guidelines for dieback hygiene procedures can be obtained from the Regional Office of CALM and the Moora District Debad< Protection Plan.

Table 3.1Stands listed in ord€r of priority for managementactions Priority determined on the basis of existing threat and b€nefit to the overall shategy. Refer to the text for details. r, Land acquisition is the responsibility of the Conservation Lands Acquisition Committee.

Managemmt action Standsin order of priority (first to last)

1. Land acquisition* 4andt |a d2, 3a, 3 6, 7 2. Prokction from grazing 4andt rehabilitation stands 3, Protectionfmm acridental destruction l andZ 3 6, 7, 4and5 4. Protectionfrom fire 4 and5, 1and2, 3 6, 7 5. Weed control 4and5, rehabilitationstands, remainder 6. Linear marking l and 2, 6, 4 and 5, isolated roadside/firebreak plants 7. E: sifu conservation 1,4, and 5, remainder 8. Rehabilitation of existing stands 1ard2, 3a, 3 6, 7, 4and5 9. Establishingnew populations not applicable 10. Artificial gene flow not applicable 11. Monitoring 4andi 1and2, remainder 12. Dieback 4 and 5, areaswith Bankia ard other susceptiblespecies

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rue.6ordlraua8errera arnIo rural .e.t REFERENCES

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31 CL

'g0t-t61'9o1. srilrur, uopeluaurSery1o slaaayqSrq Suraeqsuorlelndod Suouru.nog aua8 3urrnsea141 '(tg6l) 'd'X /âue1pue.g.O'aIpM.V ,uosrs-I 'oluo3 .yodar :luaura8euen pr.rc.Ipup uoqp^rasuoJ Jo luarugedaq 'uor8ar paqsllqndun slsaro; uraqlrou arp u! uogJaloJdpoads 1o paau ur s1ue1draqlo pue erog arer parDIJaC"I'uosqo1 pup 'W'am{8ouoq'O''q 'g'raddo11'1 'pro}raHÎ'C'sateo3'.g .y'd11a; 'gg-11\ fi8o1otX funuo4ryoa7 sraua8uorgueld peardsapr.'"rpup patJglsar d11erqder3oa8 '(lg6I) 'C ur dllgqeledurorur-Jlaspu€ rusJqdrorudyodrpauaS;o s1aaa1;o uosueduoJ V I'uore) 'p.uaqueJ :aJuar)SJo .,{urape:y uerler;snV 'IL-€g'dd "I 'Cl'M '>J 'tllb4snv Capry pup saôlg'H spg) u qrnrasay qstg p sapadgu1 'arrlcurd uolle,rrasuot o1 rar4ad outolosluI suralsdslpueS;o a)uelalal aq1 '(Zg6l) 'H'S-'saruel '96V-69n' L srttvwalshs 's1ue1d '(9161) ')'S'u1eI puv fr3o7oc7to mana6 pnuuy u1 Sulpaarqur Jo uounlola aql 'uopuol :ssar4.Qlsranlun a8pr:qrue3 '9€-9I 'dd ('sal^reH 'g '{ 'la{uerC 'H 'O spS) mo luol put fiopo1 nl satnosay 'ualsds .(916l) ').S,u1el t1aua1 dotl u1 SulpaarqJo slraJ;aar{l pup arnpnrls uo1lu1ndo4 'gt9-gz9'62 .D1ro$ frutqogto punol uaqo4snv '(186t) 'd snldlpnT ;o r.ualsdsSurleur aq1 pue uopeulllod prtg D'uero;4l pue'.q.g,raddog '1,4'g'1r1 '699-299.dd .1,1 uolrolq Surddq3 :suo5 pue ,{peag daung 1.suqdo11 '{'y 'V'plourv'M'D 'srapmes'V'O 'uo4nqa8al pue aSprqrng'V spg) aagt1,1lo slunuluay 'elleJlsnv lo alog a41 :uorloabsuo) aJrytN uI uratsa6 uI spaJp lprnl"u ;o luauraSeueu 'U86D 'I'I ^n'I'u^rorg'I .y'su11do11 ar{l ul asn :o; rualsds Supolgour V taspoo5 pue t f '6Iz-zIZ 'o9 f3o1or3 'el1er1sny .(6961) .D.d,puplloH lsear{lnos ;o uo11e1a3a,raallpur eql w1 sn\dtlntnX 1o srlueulp lqglaM '002-gll'Ol stlltuayslg ptn fi3o1ot7lo maraay 's1ue1d Trnuuy u! uopelrpl rpauaS alqelralap dlprrlaroqdortrala pup sJltslralJerer{J '(6L61 '5.'I'uo}lrnl drolsrq a;r1 uae.{laq sdrqsuorlelag pup'g'A lJBqu}.I,.I .I ,IrgureH 'uopuol :s8rmlurnSTurue{uag '8t€-9€€ 'dd ('seuroqJ '.1 'M pup 'ap,ftgre11 'g 'sraqrlleql 'I 'S'xolplPn 'J 'suorlepdod I auoqJs'I/{ spg.)'uo4ua0flo) pur eqauaS u1 1ueld p;n1eu Suoue pue ulqui uollBlrp^ r9aua8 yo uopnqlrlslp aqJ '(€86[) '.1 'I'{JuurpH '9v1 '7 frt1s1wa4toqfiq4 'suo11e1ndod uy ssatSot4 1ue1d puu aruaplla ;11a.roqdor1ra1g'(196l) .o'1 'qallllo5 '08I-L9L 'r9 uzptog lortuolog 'sr11eura1sds '(LL6D 'q 'l 'qalllloD ynosslq ary lo sltuuv 1ue1dpue aJuaplla rrlaroqdo:1rayg 'exaqueJ :atuang ;o durape)v uBIIeIsnV .ILZ-EVZ.dd .d .I 'saôrD .H .d .I 'V -ttotg ('alqoN pue lllg [ spg) unp4snv aql 'sarlJ '(1861) 'W 'V 'llID puv ?t!J uI ol sapads 1ue1drepJsul uerle4snv Jo sasuodsararl4depy '989-8r9 'Z '?ntlnztBy 'pIIBIlsnV .(1961) .y .J .raup&D otlt4snv un$eM to prnol uralsaM Jo s€rl Ledig, F. T. (1986). Heterozygosity,heterosit and fitnessin outbreeding plants. In Conseroation Biology: The Scienceof Scarcity anil Dioercity. (Ed M. E. Soul6.) pp. 77-104. Sinauer Associates:Massachusetts. Loveless, M. D., and Hamrick, J. L. (1984). Ecological determinants of genetic structure in plant populations. Annual Reoiew of Ecologyand Systanatics 15,65-95. Lucas, G., and Synge, H. (Eds). (1978). The IUCN Plant Reil Data Book. International Union for Conservation of Nature and Natural Resources:Morges, Switzerland. Mairy A. R. (1987). Maragement of remnants of native vegetation-a review of the problems and the development of an approach with reference to the wheatbelt of Western Australia. In Nature Conseroation:The Role of Remnantsof Natioe Vegetation. (Eds D. A. Saunders, G. W. Amold, A. A. Burbidge and A. J. M. Hopkins.) pp. 1-14. Surrey Beatty and Sons: Chipping Norton N.S. W. McCredie, T. A., Dixon, IC W., and Sivasithamparam,K. (1985). Variability in the resistance of BanksiaL. f. speciesto Phytophthoracinnamomi Rands. Australian ]ournal of Botany 33,

ozt-o5/. McNee,S. (1985). 'Pollinationbiology of Eucalyptusrhodantha'. Grad. Dip. Nat. Res.Thesis. Curtin University: Perth. McNee,S. (1989). 'The Pollination Biology of Eucalyptus rhoilantha'. M. Appl. Sc.Thesis. Curtin University: Perth. MoraryG. F.,and Bell,j.C. (1983). Eucalyptus.InIsozymes in Plant Geneticsanil Breeding.(Eds S. D. Tanksley,and T. j. fton) pp. 423441. Elsevier; Amsterdam. Morary G. F., and Brown, A. H. D. (1980). Temporal heterogeneityof outcrossing rates in alpine ash (Eucalwtus delegatensisR. T. Bak.).Theorctical anil Appliet Genetics57,701-105. Moran,G. F.,and Hopper,S. D. (1987). Conservation of the genetic resources of rare and widespreadeucalypts in remnant vegetation. In Nature Conseroation:The Role of Retnnants of Natioe Vegetation.(Eds D. A. Saunders,G. W.Amold, A. A. Burbidge, and A. J. M. Hopkins.) pp. 151-152. Surrey Beattyand Sons:Chipping Norton N. S. W. Nei, M. (1975). MolecularPopulation Genetics and Eoolution. North-Holland: Amsterdam. Nei, M. (1978). Estirnationof averageheterozygosity and geneticdistance from a smallnumber of individuals. Genetics 89,583-590. Nei, M., Maruyama, T., and Chakraborty, R. (1975). The bottleneck effect and genetic variability Eoolution29,1-70' T PoPulations. Phillips,M. A., and Brown,A.H.D. (7977). Matin8 system and hybridity in Eucalyptus pauciflora, Australianlournal of BiologicalSciences so,337-344. Pryor,L. D. (7981). Ausbalian EndangereilSpecies: Eucalypts. Australian Government Publishing Service:Canberra. Pryor,L. D.,and Johnson, L. A. S. (1971). A Classification of the Eucalypts. Australian National University Press:Canberra.

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