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(€66t)60t S6:(t) 6 ?'sr[nN Nuytsir Vol.9. No I (1991)

Taxonomy

Drummondita Harvey

Drummondita wilsonii F.H. Mollemans, sp. nov. (Figures I & 4)

Distinguitur folia aggregatussecus ramulo et flores color; calycis segmentainaequalia et glaber; staturanon robustus. Morphologia simrlit]ldo Drummonditq ericoideset D. miniqto.

Typus: W.A. Wheatbelt (eastern),Parker Range South, East of Skeleton Rocks (precise locality withheld),alt.450 m., 15thJune 1990, F.H. Mollemans27 61 (holo:PERTH; iso: BISH). (Figurel)

Sfrrabs40 cm to I m tall, erect,branching, ericoid. Lear€J green,crowded, linear-clavate, semi- terete,channelled above, 2-6 mm long and 0.6-l mm wide, with scatteredglands, a large,black apical gland,and a to 0.5 mm. Flowers 1-3,tub:ulrrto 1.5cm long, termiral. Calyx.se gments lunequal, oblong,4-5mm long,2-2.5mm wide, with scatteredglands; green with reddishmargins, glabrous outside,minute marginal hairs, insideglabrous. Petah oblong-ovate,12 mm long, 3 mm wide, with scatteredglands; reddish fading to green at base,glabrous inside and outside. Slamezspink, densely hairy in upper one-third. obtuse. Seed3.5 mm long, dark brown-black.

Other specimensexamined. WESTERN AUSTRALIA: W.A. Wheatbelt (eastern),Parker Range South, East of Skeleton Rocks, alt. 450 m., frequency fairly common, 14th July 1990, F.t1. & M.P. Mollemqns3143,3l44,3155,315'l,3158,3159(PERTH); W.A. Wheatbelt,ParkerRange South, alt. 450 m., frequency(locally) fairly common - 5 seen,18th October 1990, F.H. & M.P. Mollemans 3717(PERTH).

Dtstibution. Endemicto the boundaryzone ofthe Avon andCoolgardie Botanical Districts ofWestern Australia, with one known area of occurrenceon a low range located 75 km south-south-eastof SouthernCross, Yilgarn Shire, on vacantcrown land.

Habitat. Found in (Eucalyptusspp.) malleeto 4 m over llakea -Melaleuca-Grevillea scr|ubto 1.5-3 m and heathto 1.2-1.5m on part ofthe east-north-eastfacing slopeof ParkerRange South. Geology is quartzitic (silicified sandstone)as boulder, cobble, pebble and gravel outcrop, bordered up-slope (to west) by outcroppingvolcanics and down-slope (to east)by aeoliansands, with erosionalgullies to the north and south. Soils are pale yellow to yellow olive-brown sandsof a skeletalnature with gravel, pebbles and t cobbles of quartzoseand ferruginous rock fragments and minimal organic matter. D. tlilsonit doesrrct occur on the volcanicsand is replaceddown-slope by Drummondita hqssellii on the aeolian sands.

Flowering period. Observedfor June to July, but probably commencedMay as flowering well underway in mid-June,and extendsto at leastAugust as the specieswas fruiting in October.

Affinities. Drummondttawtlsonll is morphologicallysimilar to both D. erlcoidesand D. miniata. Each hassimilar flowers and leaveswith a largeterminal gland. D. wilsonii diflers from D . ericotdesin the 1-3 terminal llowers and unequal calyx segments,aharacters which it shareswith Drunnondita miniata. D. ericoides differs from both speciesin possessingleaves with apical glands which are sometimespointed. D. miniqta hasthe largestflowem with woolly tomentosecalyces and the largest leaves,which aremoreopenly arrangedandcrowded towards branch apices. D. ericoideshasasimrlat

I - O7LZ suDuallot't J I4 n I/ rt') cd,{totoH sururatlot[ H lJ uuoqlk Dllpuoaunre I etr.Btl

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ottjstsDq d pug DSaDtnaqtorytd nuorthr Dltpuoir.narc''surttle otl{ HI Nuytsiavol. 9,No. I (1993) leafanangementto that of D. miniqta,while D. wilsonii hasleaveswhich aredensely crowded along entire branchlets. D. miniqta, atleastfrom the type locality, is amore robust shrubthan D. ericoides, and D. wilsonii is the least robust ofthe three. Flower colour differs in all three species. Table I summarisesthe similarities and differencesbetween the three species.

Conseryationstatus. Poorly known, CALM Priodty l.

Etymology. The speciesis named after Paul Wilson in recognition of his taxonomic studiesof the Rutacaae.

Philotheca Rudge (tnclludesEriostemon sect.NigrostipuLae - Wilson 1970)

1. langei F.H. Mollemans, sp. nov. (Figures2 & 4)

DistinguiturErlostemonfalcatusperparvus,ovatus sepala;parvus petalis; stamina filamenta paene glabra; folia glandis valde exsertis. ryprs: Northern Wheatbelt,NW of ChiddarcoopingHill (preciselocality withheld), alt. c. 465 m., 25th August 1991,F.H. & M.P. MolLemans4127 (holo; PERTH; iso: AD, BISH, CANB, K).

Shrubs or undershrubs,erect, to 1.2 m tall and up to 60 cm across. leave,! exstipulate,shortly petiolate, ascending,clavate, 4-6 mm long, c. I mm thick; adaxial surfaceflattened, canaliculate; margins and rounded abaxial surfacewith distinct glandular venucosities,glands strongly exserted; apex pointed. Flowers 7-3,termital, on a pedicelc. 2.5-3 mm long. SepaLsobovate, c. 0.5 mm long; minutely pubescent,otherwise glabrous; centre thickened, glandular verrucose; marginflaccid. white, elliptic, c. 3-4 mm long,2-2.2 mm wide, pubescentinside, margin pilose, mostly glabrous outside,with a narrow, c. 0.5 mm wide, cenbal glandular-verrucoseband externally which is pink at the apex and fadesto white at base. Stamintl filqments 10, free, alternately5 shortc.3 mmlongand 5 longer c. 3.5 mm, tereteat apexto marginally flattenedat base,glabrous apart from a few scattered apicalhairs;anthers oblong, c. 1.0mm long, orange-pink,white-apiculate, pollen orange-pink. Ovary glabrous,of acute,narrowly pyramidal carpels. SD,leglabrous. Seedreniform, c. 1.8mm long, black, minutely reticulate-foveate.

Other specimensexanined. WESTERN AUSTRALIA: NNE of ChiddarcoopingHill, alt. c. 427 m., frequency (locally) fairty common - 12 seen,Tth November 1990, F.H. & M.P. Mollemans 3716 (PERTH); NW of ChiddarcoopingHill, alt. 465 rn., frequency (locally) fairly common - 6 seen, 8th November 1990,F.H. & M.P. Mollemans 3796 (PERTH).

Distribution. Endemicto the boundaryzone ofthe Avon andCoolgardie Botanical Districts ofWestern Australia,occuning north ofChiddarcoopingHill, which is locatedabout45 km north-westofSouthern Cross,Westonia Shire.

Habitat. Found,nt Allocasuarina campestris! Acqcict ! Thryptomenescrub and open to dense shrublandvegetation to c. 1.8 - 2.8 m tall on pale orangesand over a similar colouredclay crust, near flat granite or amongstgranite boulders. euolspu?s elru€Jc )ped i(Icou 1?llq?H pnbaun pnbaun pnbg slueru8es x,{p3 eprslno esolueuol snorq€lc ,tllooii\ snorqElc ecuecseqndx,{123 9'Z-Z 9'€ E (uu) qlpl,$ x^l"J 9-n 9-9 E (ruur) qfuel x,(1eJ 3uoyq9 elB^oqo el?^oqo e&qs x,{l€C repuels lsnqod lsnqoJ-rluas ernpls luEld (el,{ls xe) sI I E-9 7, 9I qt8uel re,lrolg Jeqluurc pad -a6uEJO ,rollel Jnoloc re^\olc

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Figure2. Phihrher:d ktrgei F.l{. Mollcmans - Holotype (F.l/. 3. i,l.P. MaLlenans 412'7)

t pepuedxe,(11eseq (ernturuur ueqm) a1,ts oscqal^ls pepuudxaeqt suruluocqcrq,tr,{l,luc pesserdep 'snorqv13 'Suol 'Suolqo 'elplled u uuog sledrecesnlqo ,&o,rp etrq,$uallod urur S 0 c s.rawrr :asolrd 'alerel 'eqnt 'luur ,{lesuep stueulelyeer; reddn EuJoJ ot JIEqIes€q eqt ur pasnJ g g c Apue 3uo1ruurg c 'esocnr:e,t-;e1npuu13 g {1a1uuratle'91stuatuDl{ loutaws €sEqeqllE elrq^\ ol Surpe;xede eqtte lurd 'qrJpru 'aprslno peue{crql p€orq lutu I c relno uu "{llclluer reSuolsrrEq'urqlr,^\ luecsaqnd snorqElS 'el"loqed '3uol 'e1rq.t 'peuelcrql ulru (9 9)9 c cr1d111e s/rlrd esoJnJJo^-Jplnpuu18 eJtuec:snorqplS 'pr.ceu 'rEln8uuul asr,rreqloluecseqnd ,{lelnurtx ur8rEur:8uol ruur I-9 0 c .tlrdas Suolulurg l-9 0 c '1euruua1',{re111os 'peuesxa alcunpedeuo s.raaroigpepunor xadu ,{13uo.4s1ou spuu13 tnq seqrsocnrre,r 'JIBq '(qserJ re1npuu13lcurlsrp qlr,lr ec€IJnsIcrxeqe le,rol aql ur pe^ool8 ef,eJJnslerxep€ ueq,r) {crql 'Surpuecs€'etE^€lc-repuels 'e1u1ndrlsxe uru g 1-1 c or dn'3uo[ ruurtI-l c uuIuIEI:el€loqed ,{p:oqs 'peqou€rq-qcnlltr sa^Daj ssoJxeurc gg o1 dn puu llel u I ot'sqnrqsfipun ro s4lrqs Surpueids

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Table 2. Comparisonof someimportant taxonomic and ecologicalfeatures of Philotheca langei F.H. Mollemans, E falcatus P.G. \Vilson andPhilotheca basis4,/aF.H. Mollemans.

Character P. langei E. falcatus P. basis\,lq

Plant Erect Sho , much- Spreading, stature/branchi ng branched much-branched Leaf length (mm) 4-6 6 7-14 Leaf width (mm) I I 1-1.5 Leaf shape Clavate Falcate Slender- clavate Leaf apex Pointed Rounded Rounded Stem glands Abundant Abundant Few Gland exertion Strongly Exserted Exserted exserted length (mm) 3-4 7 6(6.5) Petal width (mm) 3 Petal petiole Absent (?) Present Calyx length 0.5 3 I Calyx shape Obovate Narrowly Triangular tdangular Calyx apex Obtuse Acute Obtuse Staminal filament A few Scattered Densely pubescence apical hairs pilosein hairs upperhalf Staminal filaments Free Free Fusedin lowerhalf as a tube Anthers White- White- Non- apiculate apiculate apiculate Habitat Sandloam, (?) sandplain Deepsand yellow Allocasuarina Eucalyptus, mixedheath - (9Zb sunuatlon .l n T H J) ad,{lotoH suxuretpt.i H i'tttfi!-Dq naqtolqd t n8rt

rtrt 16 | . ^,rt JLren.ld rFt dJ 4b..,rji]bl'|J.'P*lv4o..nv ?t :,ai rrt.ttt i s ltz dl eli .r \?.q) )o 1'Ed .- "s, '$'*3-,r ,*. ?rs.-]rr tz r.,.+i.'a:''"2-r l aa.6 P,s p.!-|'rt tr, @ r.€r -6rl'c rrar { .e.n4 Stiiaitt6 ;r;i:!ir;6ffi F,rbtig,/. rr66?:F6J_jg

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(pear-shaped)above point of attachmentwith ovary andtapering upwards, narrowing asstyle matures, denselypilose. Fr it not seen.

Other specimensexamined. WESTERN AUSTRALIA: Trayning District, 180 km eastof Perth, alt. 290 m., frequency(locally) fairly common- c.4 seen,20th August 1990,F. H. &M. P. Mollemans3213 (PERTH); Trayning District, 180 km eastof Perth, alt.290 m., abundance(locally) fairly common, 25th August 1991,F.H. & M.P. Mollemans 4124,4125 (PERTH).

Distribution. Endemic to the boundary region of the Avon and Coolgardie Botanical Distdcts of W€sternAustralia, occuning south-south-eastof Trayning, which is locatedc. 180 km eastof Perth, TrayningShire.

Habitat. Found in dense scrub of Eucalyptus leptopoda, Grevillea eriostachia, Allocosuqrina tlcutivalvis, Baeckeq nuicqta, Baeckeafloribunda, Hqkea coriacea, Melaleuca conothqmnoides, Melqleucqsp.,Dqviesiasp., Choretrumpritzellii, Phebaliumtuberculosum, Pimeleasp. and Hibbertia sp. growing on deep yellow sand.

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'lluoslltt 'sucuellonl DlkrNDq .l pw DBUDIDraqpllqd otipu{xa&flJof H J Nultsir Vol. 9. No. | (l9ql) in habit, leaf and floral characters.Compared to the otherDrummonditataxa (D. hassellii,D. hassellii var.longifuliaand D. calida), all taxahave fl owerswhich exhibit variationsin colour andshape offloral parts,butthere are significant dift'erencesbetween D. hassellii,D. hasselliivar.longifuliaand D. calida and what may be termedthe Drummonditaericoides grotp (D. ericoides,D. miniata andD. \eilsonii), in the possessionby the latter group ofleaves which aresimilarly shaped,channelled above and which possessthe large terminal gland. None ofthe other taxa havelarge glandsatleafapices. Severalleaf charactersmay thereforebe addedto petals(used by Gardner1928) to furtherestablish the relationship betweenD. ericoides and D. mini.lta, ^nd to add D. wilsonii.

Pollination mechanismsand flower colour vqriztion in Drummontlita. One of the more striking charactersof D. ericoides, D. miniata andD. leiLrotii is that offlower colour. Thepetalsof D. miniata areorange cinnibar in colour,D. ericoidesyellow-white (Gardner 1928) andD. wilsoniireddishfading to greenat the base. From west to eastthe flower colour of thesespecies, which are groupedhere due to similarities in petalsand severalleafcharacters, grades fiom yellow (D. erlcoldes)through orange- cinnibar (D. miniata) to rcd (D. wilsonii).

Gardner( 1928)noted a similar patternofpetal colour variation in the flowers of D. ha;sellii again from yellow through to red, from west to eastin its distributionalrange. This, he suggestedlimited the value of flower colour as a distinguishing characterbetweer' D. miniata and D. ericoides. Colotr variation in the flowers ofD. ftasselliiis apparentlyclinal in nature,with populationsvarying more or lesscontinuously over its range.Butthis speciesis quitevariable taxonomically and may intimebe subdividedinto more than onetaxon. Parallelsare therefore apparent between what may be termedthe D. hassellii group andthe informal D. erlcoidesgroup (which includesboth D. mhiata andD. v,ilsonii) recognisedhere.

The speciesof lhe Drummondita ericoides group have a broader geographicrange and more northerly distdbution than that of the D. Dassellilgroup, but arerestricted by habitatrequirements to certaingeological features (Table l), ofapparentlygreat age. Although exhibiting an identical pattern of flower colour to the D. hassellii group,the speciesof the D. ericoides grottp,rather than fbrming a cline, apparentlyoccupy ditTerentlocational foci within a hypotheticalcline. What we seenowadays in these three speciesmay be three divergent members of a formerly more widespread,common ancestor.

Flower colour has definite advantagesfor pollination, and in the D. ericoides group the flower colour of a common ancestorappears to have beenretained by eachspecies in the group. In the flora of south-westWestern Australia the coloursseen in this group areamongstthe four mostcommon and Drumtnondita is bird pollinated accordingto Keighery (1982).

Similar colour variation to that presentirboththe D. ericoidesgroup and the D. ftassellllgroup is also found in the flowers of Diplolaena ()with flowers againvarying in colour from west to eastfrom yellow throughto red (G.J.Keighery pers. comm., P.G. Wilson pers.comm.). In this instance, however, the colour variation is in the (P.G. Wilson op. clt.), which in Diplolaena are quite long and prominent. Armstrong (1979) lists birds as pollen vectorsit Diplolqena.

Drummondita miniata is thought to be bird pollinated as Keighery (pers. comm.) has observed brown honeyeatersvisiting the large flowers of this speciesat the Cue Rifle Range (Cue is the type locaity of D. miniala). The relatively smallerflowers of D. ericoidesand D.wllsonll (Table 1) would seemto be a factor preventingbird pollination. However, brown honeyeaters,for example,possess a o1rq,tperelleun ,{lpquasse paureueJ spq stBJlsqns eql'(,raproJlta p|tpuowwwO lo tplrqBq)uotppleC .r€eu)ped elrqld pu€ (tluosllM xtJrpuoww,?lclo tElrqBq)qlnos e8uuu ra{ryd ss qcns sllrq uO

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'Iereruseqr,{q pesrlln eq,{po uuo puu slcasur.{qpelrsr.t ^,rr", "r" r:iifii,lX""lffiiffitt:X: '(6961) ,{req8re; olSurproaJv sercedso,trl eseqlgo sJs/ri\oU :elnqnl ,roJruueql ele.ueued,{lrsue plnoc feqt qcrq,l.r.{q'(ruruoc sreducqurg q) xedeeql te e8upuaddue{rl-IeqteeJu qlr,r en8uolrepuels Buol

n|^lstsnq .l puc laduol n)2q1t41qJ llaotlt r ot1pu.rlrllrrro.suDuallol H d NuytsicVol. q. No. | (1991) the climate has changed through time. However, on adjacent more recently developed aeolian sandplainsand associateddeposits both the substrateand the climate haschanged.

D. hasselliihasdeveloped on the aeoliandeposits, and in theParker Range south area there is aclear segregationbetween D. hassellii and D. )eilsonii. D. reilsonlloccurs only on skeletalsoils developed on the geologically old rocks,while down-slopeit is replacedby D. hasselliionsandplains lapping onto and having extensiveoccurrence adjacent to the range. Even thoughboth D . wiLsoniiand D. hassellii occur essentially side by side at the boundary between outcrop with skeletal soils and adjacent sandplains,no evidsnt intermediateforms (hybrids) were observed.There is a clear subsbaterelated line of demarcationbetween the two species,and this may also reflect a clear geneticsegregation.

It is possibl€that if D. ericoldes,D. wilsonii and D. miniata arebrought togetherexperimentally they could hybridise,as a relationship(albeittenuous) has been established between them using several leaf characterscoupled with the petal characterused by Gardner(1928). But this would not occur in nature due to the vast distancesseparating the thre€ species. D. ericoides occuls near Geraldton, D. mtniata in the Cue district and D. wiLsonii occurssouth-south-east ofSouthern Cross,D. ericoides is separatedfrom D. miniata by a distanceof 360 km, while thesetwo speciesare separatedfrom D. wilsonii by distancesof 600 and 550 km, respectively.

Heathvegetation is thoughtto haveflourished in Australiafor almost50 million years(Specht et a/. 1979),and had its beginningsin the pan-Australianrain forest which existedwhen Austualiawas paft ofGondwanaland.The break-upofCondwanaland allowed for altcredclimatic influences,particularly around southernAustralia, which was associatedwith the gradual separationfrom west to east of Australia and Antarctica. As Drummonditaexhibits evidentdivergence, associated with climatic and physiographicchanges stemming from continentalseparation, there are clear indicationsthat it is a genusof great antiquity.

Acknowledgements

I would like to thankJim Armstrong,Paul Wilson andthe staffofthe WestemAustralian Herbarium (PERTH) for assistancein work associatedwith the preparationofthis paper. The initial collection of the three speciesdescribed here were madewhile I was employedon contractwith the Departmentof Conseryationand Land Management(CALM) on a surveyof rareflora in CALM's Merredin District, a project fundedjointly by the Austalian National Parksand Wildlife Serviceand CALM. I wish to acknowledge the funding bodies. Thanks also to Mr P. Brown, Dist ct Manager, Department of Conservationand Land Management,Merredin, for permissionto collect in ChiddarcoopingNature Reserve.Jim Armstrongand Paul Wilson pointed out aspectsof this paperrequiring improvement. Special thanks go to my wife Nellie for her untiring assistanceand support,at home and in the field.

References

Armstrong, J.A. ( 1979). Biotic pollination mechanismsin the Australian flora; a review. New aland J. Bot. l'l| 467-508. Gardner. C.A. (1928). Contributionesflorae Australiae occidentalisVII. J. Roy. Soc. West. Austral. l4: 79 80. Hawkeswood,T.J. & Mollemars, F.H. (ined.). CalothatnnussuperbusT.J.Hawkeswood & F.H. Mollemans(kptospermoideae: Myrtaceae), a new speciesfrom south west Western Australia. NuylsirS:311-318. '8tz-gtz:8 '(eBecElnu) Brs nN !?lleJlsnYuretso?\A uJo.IJ se!.ods ^\au e DalrltxDraqloltlld,, (2661)c d'uoslnA '/0Z-16l:I 'eEecstnu '(l/6I) er$,(nN ,,e{eJlsnvuolse/A Jo ,([edlcuud ,(Fx€Jeql uo sEouclrlouo).eJ. g d'uoslr^\ 'ssI-.I :IBrsfnN,peaceln\'umlpqaqdptreuoualsolra'oa or)ereueBeqlJo uolsl^oj cllllouoxel V,, 10/6I) D'd'uoslLlA ( Illeplatsulv :,{usduroJ 'sopnrs '( '1 Suqs{qnd cgquarcs rer^eslg) el-g dd pcq.{puv g spuelqruqsparelau pue spuqqtBeH pg) lqceds u 'sur-ldoH 'srogou "I .4 spuellEeq uBrleJ|snv:suqt(qr SuFe.Alogpus q1^\oi3Jeuossas (6/61) L I >l v 4, /lt|U d lqceds

'l1uosllt 'susulellold Dtt$lsDq d pw la3uol otautol1lld Dllpuourrrro H d