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(tg6t) l/Z-t0Z:(Z)9 elstfnN Z0Z NultsiaVol 5,No. 2 (1984)

not presented features or for i:rropercircumscription of his taxa Speciesdescriptions were by Melville. Thi lick of effective keys togethor with the presencein Australia of several hitherto unrecognizedspecies has led to much conlusion in herbarium collections'

The present revision is based on a reassessmentof morphological oharactels ol specles preuiouily consigned to the genera ltwrentia and Salenothqmnus. Despite considerable variation'in habii, trichome type, sexuality and fruit and seedmorphology, it is considered that the division betweel IL\\ren(:ia and Selenothumnuscan no longer be upheld and that the 12 speciesconcerned are best placed in a single genus. Although clearly delimitable groups e;erge, thesedo not correspond to preYiousgeneric concepts,nor does the evidence ieern strong enough to suppofl their recognition at generic level. A possible cxception Is L. berthae, a tptii"t to much at variance morphologically with those remaining in the genus that a str;ng case could be made for its segregation.This has been deferred pending further investigation ol the genera ol lhe Plagianthus aliance.

Taxonomic History The gcnus lirwrencia was erected by Hooker (1840) to accommodate a single species, L. spi(.tta, specimensof which had been sent to him from Tasmania by Ronald Gunn. Hooker (1842) describeda secondspecies, l. glctmerata,sent to him from Western Australia by James Drummond. A third species,L. squamata, was added to the genus by Miquel (184s).

Bentham(1862) united under PlagianthusJ. R. & G. Forst. (1776)non Jaub. & Spach (1845)species variously placed until that date in Asterotrichion Klotzsch, Blepharanthemum Klotzsch, H(tlothamr'r6 F. Muell., Hoheri.l A. Cunn, I'awrencia and Plagianthus.This action was followed by subsequentauthors, including Bentham in Bentham & Hooker (1862), Bentham(1863), Baker (1892),Schumann (1890), Dalla Torre & Harms (1900-7),Leme6 (1934) and Burbidge (1963). Several new speciesof Plagianthus sensu lato were described during this period.

In January of 1909,O. Stapf exhibited specimensand lantern slidesof Plagtunthushelmsii to members of the Linnean Society of London pointing out that it appeared more natural to him to treat that speciestogether with P microphyllus ar'd P. squamotuJ as members of a distinct genus for which F. Mueller's name HalLtthomnus, orginally applied to P microphyllus, would have to stand (Anon. 1909).Amongst those pa icipating in the ensuing discussionwere E. G. Baker and T. A. Sprague.

Sprague and Summerhayes(1926) examined the New Zealand representativesof Plagianthus sensu lato which they assignedto two genera, Plagianthus sensu stdcto and Hoheria, notingit passingthat the Australian speciesof ltwrenc:ia and Hqlothamnus should be excluded from Plagianthus.

In the late 1930s, E. G. Baker prepared the draft for a paper which described three putative new speciesof Lawrencis and provided a partial key to the genus From his dralt it is clear that Baker intended to reinitate the gerruslawrenciz. It is wo hy of note that Plagianthu; berthae ariLdP. helmsiiwereexcluded from consideration,although the significance of this is not explained. Copies of his unpublished manuscript are held at BM and PERTH. Baker anrotated specimenshcld at BM, K and PERTH.

In a more comprehensivestudy of the Plagianthus complex, Melville (1966) divided its speciesamongst six genera. In particular, speciesplaced by Bentham (1862) n Plagianthus sectron lr\urencia were now allocated to two separate genera, namely ltwrencia ser's]u stricto and Selenothamnus, founded, as Stapf had originally suggested, on Mueller's 'uorlelueserd crurouoxel IsruloJ sql s,,r,iolloJr{crq.,tt 3cuecgru8rs ,{J"uorlnlo^epu? cluouoxet JraqlJo uorssncslpplauo8 eql JoJpue oDuah$rJ Jo uoll€cUISSEIc Surnsueeql JoJ srs"q " epnord ereq poreplsuoc slre.t1pctSoloqdroru eq1 snue8 eql 1o sJeqtuetu[e 01 uouruoc su;e11edpcrSoloqdroru lurcgru8rs Jo uorleu€A Jo o?u€r otll uo 'a)uat$D1 poceld sr sseqdrug 3o ,{3o1oqd.touroql Jo tunocce olltsneqyo ue epr,ro:d o1 Surtrdruelleu?ql reql€J ,{8o1oqd;ouj1o stcedsea.Lrlereduoc qll,r\ peurecuocsr uoqros srql ,{Bo1oqdro141a,r.gu.reduro3 '(ssardur) ropu"'I ul puno; eq deur '8uuaql€E sepcqdnp elq€ll"^" Jo uolt"clpul uB qrl\ reqFSol Jo setep pu? selll"u,sroFeIIoJ 'sertrl"col qlr,,r\sueuroeds po,{orlssp aql 1e yo 1sqe,ltsueqerdruoc y lsrrelst ue fq pat"clpul eJB 'txel tueprcJesrql ur pello,rur 1ou selecrldnppetnqulsrp ,{ltuoceJ eql uI ereq,taqe sedf,1u,{sro '.{laleunpog sed,(lo$ fq reqlre peluesuda: er" pe^Io^ur suerurcedsed.{l eql II€ ?rg }q3n"c pue p€oJ eql Uel sel"ls auoq Jreql ol Fueleu ueol SuluJnter {cnJl el{t ueq,tr pedorlsep e:a,r reded srql ur patrc (gg) unueq:eg u"IueruseJ eq] pue ('IAIr'{) "uolcl1 Jo ulnlr"qreH 'suarutceds Fuorl"N eql qtoq ruo4'sedf,1 Sutpnlcut oql lle leq1 Uodal 01 3^eq ()11el33l I 'sasEcsnon8rqu? asr,\\Jeqlour ,(Iuo pelrc er? s-requnu leeqs ro relsr8er urnueqlaq :elqelIEAEeleql\ pellc sI uollcalloc Jo 3lup eql requnu qJolJallot e Jo ecuesqeeql uI suorl"ls l"lols€d ole s?u.lulot peue^ul e1autsur ue,lr3 'oasu3-tpt,nt sertrE)o-l IlnJ ur pelrJ eJe3J3q p3qucsop ,{l,]!,\eu setceds 1o suerurcedselq"llE^" II€ orcua,turry1o uorldoJxoeql ql1\[ sercedsqcu e 1o e8ue: eql lnoq8no:ql s3trsle suorl€lndod Jo uorlecol oql 1ur:ed ot ,{preruud pepualur eru luetutearl ctruouox?l SultrolloJ eq1 ul slsll polr?lep arou eq1 reded srqt Jo uolsnlcuoc eqt le tuJoJ pel"Ilelqqe ut pept,tord st ,{pn1s srql Io esrnoc eql ur po]"louue pu€ peulruexe suerutcedsunu"qreq Jo lsll alslduloc V 'r7rupurspl Jo )uJulteJl rlqerEdLuoJou :r JrJqf 'sop ',{rolur3J I qlnos .,nep.rog (1961) le8ntq a p]el3ld pue urequoN oqt roJ (l/61) eppueddrq3 '?uotcr^ roJ (0861) eloqelSnBeg'"llErlsnv urolseiN roJ (0861) pr€eg'erlertsnv qtnos roJ (6/6I) uouv 'pueysuaenf rcJ (ZL6I) uouy ,(lerueu 'elets qcee ut slsueloq ,{q peldope dlpreua8 suorsr,\rp-qnspcrqderSoe8 eql ol pu" sdeu eqt ot eJualeJet,{q pateclput 's,r\olloJ $ sercodsqr€o Jo uorlnqulsrp eql 1?ql luour}lrsJl clluouoxPJ oql uI el"ls lEql ()1 peuguoc sr sercedse ereq.trerprtsnv uralsoi6 Jo uorpefo.r4 eery pnbg Ieqlruez ueqrue'I " uo Jo erleJlsnv Jo uorFeford ea:y pnbg seuuog " uo pesoduuedns sor€nbs e3r8ep ouo ur ocuerrncco neqt SurpJoceJf,q peldutoc ore,t ,{lalltcedseJ8-E sd"l,1'lut selcedseqt [e Jo uorlnqrJlsrp oql pup I del^l ur paleJlsnllr olJuar,trDTJo o8uEr Jrqd€r8oo8 lelol tql 'red€d srql puo eql te pelrc sueulcedsrunueqJeq eql ur per\.resqouorl"ue,\ plol eql uodn peseq e.r?pelsrl slueueJnseelr'{ (8961) slnorlua3-I ,{q pozrJeururnsosoql elo,r peldope sa:npecord pu" spoqlou eqf €ll"r1snv urrlsal\A ur luesord '€ueqJol{ selJedsJo uorlE^Jesqoplou pu€ uorFelloc uo pu" qsIlIJg tuE^oler pue ueII"JtsnV Joleru eql ,4q peueol ro 1" pouruJ€xosuerurceds unrJ"qJaq elq"ln^" Il" ;o ,{3o1oqd:our ssol8errr Jo .{pnrs e uodn*-iffil"ff:Jllxt epeusuorsrcop clluouox"] eqJ

'uJuatt\t/l pvg snuuoqrcuaps Jo uorlerEdJsaql pJuousanbpue sJopeEqc cqsou8erp I"JoU s.ellr^l3l^iJo SurddeFe^o aqt ot uollualt",,i^eJp (t861) Je{r"g '?xverIIB snLItuDtSDlder{l'secuerTls lEqrJl-qns FuroJur srq Jo euo s3 1rpsldecc€ eq 'xeldu-Ior qSnoqll€ srultuot8Dld3q1Jo tuoruu8ll€orqollr^lsl^l uo tuoururoc ot peullcap (8961) 'eeecE^leJd set"g eql Jo e"a^lel^troqlrl at{l uI sdlrlsuolteleJJI.IeueB Jo ri\el^.3Jslq uI (€861) '(/961) llaqJII l puB (1861)seut"g uosuqctnH fq peldope ss.'t\uor]€.grss€lc s.el|^lol^ sll€^\ 'tlqeq drrcueur eql ;o SuruelJrql eq] Jo ornl€u tql pu€ arntlts3^ uI soJuole11lpJo sIS"q eql 'snuuDqlolDq uo e[^le]{ dq peqsrn8urlsrporo,r ereue8 o,rt esaqJ Z98l ur peqrJcsep$ttJ

,!rur,/l,11r7}o uorsr^ou repuel s N 204 NuytsiaVol. 5, No.2 (1984)

Habit and duration.'fhe branching pattern of Lawrenciq is monopodial. In the seedling stage all speciesproduce a single actively growing main stem. The main axis of the young plant normally soon losesits dominance and severalto many primary branches,o{1en basal, achievethe same stature. Secondary branching is particularly frequent as the inflorescences are produced and during the secondyear.

Six speciesar€ suffrutescent: llwrencis dy'/asa is diminutive and prostrate, L. rePens is decumbent, L. densiJlora(Figure 64.), L. cinereq, L. glomerata (Figure 4) a\d L. spiclta (Figure 3) are conspicuous,erect sub-shrubs.The herbaceousstems of thesespecies become soft-woody with age. Occasionally, such plants die back or are burnt or grazed back to a ligneous branched caudex at ground level from which nsw Srowth is initiated.

The remaining six speciesare small shrubs which may be weakly asaendinglittle-branched plants, such as Lqwrencia buchananensisa\d L. viridi-grisea (F\gure 5), or divaricate shrubs, such as l. berthae (Frgure 2A.) and I' squamata. In L. helmsii and, to a lesser extent, it L chrysoderma (Figure 7A) marked contraaton of densely ramulose secondary stems givesthe plantstheir distinctiveappearance. Shrubby species normally producenew growth liom erectwoody stems.

Although various speciesol Lah,rencia have becn repoded on labels ol some herbarium specimensto be annual, it appearsthat all speciesare perennial, the few instancesof annual duration being attributable to the plants flowering in their first year. A11of the taxa observed in the field appear to reach reproductive maturity in their first year with adequaterainfall.

Thorns. All but one speciesol lttwrenciq are unarmed. Specimensol L. squqmata olien have prominent thorns. These are tapered, depauperatebranches.

Vestiture. yariallori' in trichome types and their distdbution is invaluable in recognizing taxa at all levels it Lawrencia. In this genusr as in many others in the tribe Malveae, the usual multiradiate stellatehat of the Malvaceaehas apparently beelrmodified to produce a wide range of forms which grade imperceptibly into one another.

The commonest hair type encounteredis representedby the haplomorphic (multiangulate), multicellular stellatehairs found in all specieswith the exception of the glabrous ltwrencia spicata a1.d glabrous plants of L. diffusu and L. glomerata. Haplomorphic st€llate hairs (Figure lA) are the predominant hair type found in l. berthae, L. cinereq, L. densiflora, L. dilfusa, L. glomerata and L. repens in which they occur on stems, stipules, leaves and the abaxial surfacesof calyces, invariably in conjunction with bifurcate and simple hairs irto which they merge.

Haplomorphic stellate hairs are found on the abaxial surfaces of petals n Lawrencia cinerea, L. densiflctra and L. glomerata. They a.e present to a lesser extent on all other pubescentspeoies of Lewrencia, although nearly lacking in some.

On the stems, petioles and pedunclesof Lawrencir berthae, haplomorphic stellate hairs are often distinctly raised on wartlike mounds of epidermal and possibly sub-epidermal tissue thus forming tufted stellate hairs (Figure lD), a phenomenon obseNed in no other speciesol Lawrencra.Haplomorphic stellatehairs are also found in l. berthae li the jur'cti.otl of the style branchesand on the surfaceof the pericarp.

Actinomorphic (flattened radiate), multicellular stellate hairs with more or less basally coalescentrays olr a multicellular base or stalk (Figure lE) are the predominant hair type found in llwrencia viridi-grisea and in many populations of L. squamqta. In these species, such hairs are found on st€ms,stipules, leaves and the abaxial surfacesol calyces,occurring

a- -Duldpos,tltp netl eldwts - .DuDpos,furl) _ 'Durapostrnp 1 Jo | / lo oprs .lEujlBd-- I ? Jo ele.s el€ru1ed:uogeluelool pus uonrnpod epcs'eruilode-rqu! 'turapostfutp "I H llsupq .71go D lo e1ors,lellod opuqutg :s{er Jo irurisoleoJ f,.oasu3-tput^ Jo srreqepfla1s cr-qdloruouqci :uoteluouoer pue uote]IJeJal aDqtlaq"I- eseqposrer qlr,r\ - 'Drolljsuap l rl€q olelols 5rqdroEoldeq :s,{€_rJo uoq?cndtFr^I d rruq otdurls Dtolrsuap rreq al€crn;-rrq :uoqcnpo1 g nnl{.tsuap jo nA 1 Jo _ J .Dlzuu4rlZ Jo i )|elJrlsJrqoroLuoloeq przrlerJrdsu0 V ur srreqowllots Jo uoqrnpar pup uorteroq€lE I em8rd

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'lepue.I s0z ,?rra,?,1.u4Jo uotsr^ou S.N Nultsia Vol 5, No 2 (1984) 206 into which they in conjunction with haplomorphic stellate, bifurcate and simplc hair types merge. ia buchdnaneniis Actinomorphic stellato hairs are present to a lesserexterlt in l-4['r'ln' and simple hair types' in *hi"h th"y'o."ur in oonjunction with peltate scalesand bifurcate in all pubescent Bifurcateand simplehairs (Figure 18, J & J) are found to some.dcgree but can be readily sDecies.Thev are oiten the dominant iair types prescnt on stipules' simple hairs are otten ""ur.t""a "J,n" margins of leaves and calyx lobes tntermeshing found at the baseof the petals of male flowers o[ Lawrencia berthae' re thc p redomi nan t Minutely fimbriate or entire multicellular peltatescales (Figure 1F & C) a ar'd' L helmsii' and \n tti"tro-" iip" {oul.d \n Lawrentia but:hananen;is,L thrysodermtt leaves and many popuiations ol L. squamata. Such trichomes may cover stems' stipules' occur in conjunction tt.r" uUu*iul surfaces ol caiyces. In L buchananensis,peitate scales ln the thr€e- other *iifr rt.ttut", bifurcate and simple hair types into which they merge and stellate'bifurcate snecieswith peltatescales, these occur in conjunction with palmate scales' and simple liair types into which thcy merge' L helmsii Bract-like palmate scales(Figure lH & l) are found o\ I{t"tlen(ia chrysoderma' extending ""J i. ,q"ri.i"l"which they-oftendominate the vestitureof stemsand branches, to merBe inlo ont , tfrJ Uutuf portion of stipules and leaveswhere they can clearly be seen peltate scales. of Laurencia herthae Glanrlular hairs (Figure 2l) are found only in the sepalinenectaries (seebelow). all In some forms of I'qwrenciL glomeratq, in most populations of L dilfusa' and in and ol L. spitaru, all vestiturehas beon lost ln otirer taxa the above-groundvegetative plant parts reproductive parts are variously clothed. Thus the vestiture gives the plants or thiir characteristic textures: smoolh and glabrous, scabrous,velvety or shiny and armour- penetratc like. Most of the trichomes are colourless,vitreous and rigid at maturity; Irone can skin.

Iaaves. As in many genera of the Malvaceae, the leavesof l-awrentia are spirally disposed' venation Although spirally ;ipulate, ^petiolate,'sJbsessile or sessile,and have actinodromous arianged, re

Ail speciesbear persistent, paired laterai stipules whioh are shortly adnate to the base of the petiole, or to the base of the leaf blade of sessileleavcs, and appressedto the stem or slightly reilexed. These stipules are translucent and colourless, or opaque and brown o, puii gri"n, membra,rous,succulent and leaf-like, or dry and spongy fhey may be filiform' ttiunguli., elliptic, obovate or sub-orbicularin shape They are minute ot consprcuous' 0.4_1b.0mm long x 0.2_3.6mm wide at the base, and symmetrical or somewhat falcate. are apically acuminate' acute Their margins"Theii aie entire or more or less ciliate and they greatel or obtuse. vestiture resemblesthat of the stcm and leaves,but there is o1len a frequency of simple and bifurcate hairs as is noted above With the exception of L' spiut| anj glab.ous plants of t. glomeruta, both abaxial and adaxial stipule surfacesare usually hairyl Uut the adaxial surface is typically less densely hairy and often glabrous or nearly so. Stipules may be rather variable i[ any one specles.

By virtue of their presenceor absence,length, cross-sectionaloutline and vestiture the petioles of ktwrencii are of considerable diagnostic value The leaves may be distinctly -!

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the only exception being certain populations ol L. squamata in which they are 3-5-dentate towardsthe apex.

Inflorescences.Like other genera in the tribe Malveae, inllorescencepatterns ln l-twrencia are rather plastic and henceeasily modified accordingto the ageand vigour of the individuals. Thus, inflorescencesolten exhibit striking differencesin expressionwhen comparedat different stagesof development,even on a singleplan1.

With the eKception of a single species,the flowers of llwrenciq are subsessileor sessile, solitary, born in the axils of leaves and their associatedstipules. Flowers produced later in the season are often congestedtowards the stem apices. ln many speciesreduction of floral leavesand of the internodes produce terminal and axillary sub-spicateinflorescences. Further reduction of axillary sub-spicate inflorescenceslocated in the upper part of the plant resultsin the congestedterminal spicateinflorescence observed in fully mature individuals ol L. densiflctra, L. glomerqta ?LndL. spicala (Figure 3), as indeed the names ol these speciessuggest. ln all of thesespecies each flower is subtendedby a single leai.

One speciesis exceptional: the llowers ol Lawrencia berthqe are long-pedicellate.The [rs1 flowers are borne near the base of the plant and are usually solitary in the leaf-axils, but later in the season they arc borne in axillary iascicles of 2-5 subtended by a single leaf (Figure 2A).

Flower,s.An epicalyx is lacking in all specieso[ Itwrencia. Peculiarities of calyx, corolla, androecium and style morphology are discussedbelow.

It is worth noting at the outset that three stagesof sexual dillerentiation are found in Lqwrenciu:

1. Hermaphroditism: flowers olall plants bisexual.This condition isfound in L buchqnunensis, L. dilfusa, L. repens, L. spieata (Figure 3) and L. viridi-grisea.

2. Polygamodioecy: flowers of male plants with larger spreading calyx and corolla and a sterile vestigial gynoecium; flowers of female plants with smaller erect calyx and corolla, with or without sterile anthers; flowers of bisexual plants with erect calyx and oorolla, as large as those of males or almost so, with fertile anthem and gynoecium. This condition is found only in Z. glomerata.

3. Complete dioecy: flowers of male plants with larger spreading calyx and corolla, either lacking gynoecium or with a sterilevestigial gynoecium; flowers of female plants with smaller erect calyx and corolla, with or without sterile anthers.This condition is found in L. berthae (Figure 28 & C), L. chr-ysoderrza(Figure 7C, D & F), L. cinereu, L. densifiora (Figure 68, D & E), L. helmsii and L. squamata.

All bisexual flowers of Lawrencia areprotandrous. The style branchesare closelyappressed in early flower and are exserted from the filament tube. Anther dehiscenceoccurs well before the stigmas separate and become receptive. As the flower begins to wilt the style branches reflex into the anther cluster: it is thus theoretically possible for self-pollination to occur at this stage. -fhe Calyx. persistent calyx in hwrencia is composed of five sepals that are plicate and valvate in bud and connate for part of their length. At anthesisthe basal united portron is turbinate or broadly to very broadly obconic; the lobes are flat, erect, narrowly to broadly t angular, and obtuse,acute or acuminate, and they lack accessoryteeth.

-' a :

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.s ,rraarr{r7 Jo uorsr^el{repuel N 210 NulasiaVoL 5, No. 2 (1984)

(0.8-2.0mm), L. difusa (1.2-1.5mm), L. helmsii(0.5-2.0 mm| L. repens(l.l-1.8 mm) and L. viridi-grisea (1.6-2.6mm).

Auricles, basal winglike outgrowths at the baseof the petalsof many speciesof Malvaceae, are entirely lacking rn Lawrencin.

The apices of corolla lobes are acut€, obtuse or truncate and may be entire, emarginate or irregularly dentate. The last feature was observed only rn Itwrencia squamata and L. chrysoderma.

The corolla lobes are usually glabrous on both surfaces.There are severalnotable exceptions to this: corolla lobes of ltwrencia berthae sometimeshave scattered, intermeshing simple hairs in the basal area on both abaxial and adaxial sudaces;those ol L. cinerea are stellate- hairy along the mid-veins on the abaxial suface, weakly so in male fowers, strongly in females; those of L. densiJlora are often sparsely stellate-hairy abaxially in male flowers, denselyin females;those of L. glomerata carl be stellate-hairyalong the abaxial mid-vein.

In dioecious speciesthere are striking differences between male and female flowers in corolla length, the degree of fusion of the petals, corolla lobe width, shape and odentation, and, in specieswith stellate-hairy petals, in the extent of their vestiture. The corollas of male flowers are generally larger than those of female flowers; their petals, which are almost free or only slightly fused, are flat and spreading at anthesis and, if stellate-hairy, then sparsely so. Corollas of female flowers are smaller by comparison; their petals, which are fused to a far greater extent, tend to be carinate and erect at anthesisand, if stellate-hairy, then denselyso.

In colour, the corollas are white, cream-coloured, pale yellow or pale green, and may be tinted red or purple. Corolla colour is often quite variable within a single species.

The petals are membranous and elaborately vascularized with actinodromous reticulate veins which anastamosebefore reaching the margins. They thus resembleto a remarkable degreethe wings of certain insects.

One species,lnwrenciq berthae, rs notable for its sweetly scented flowers, a character detectedin no other speci€sin the genus.

Androecium. The androecium of ltwrencia is composed of monadelphous stamens which arise from five groups of primordia (Heel 1966). This five-ranked symmetry is usually not evident at anthesiswhen it is obscured by the densely packed nature of the anther cluster. Only in L glomerata is the apex of the filamcnt tube ever distinctly five-lobed. In male and bisexual flowers of Lawrencia, the filaments are united into a tube 0.4-7.0 mm long and are free at their apicesfor an additional 0.2-2.0 mm. In female flowers, the filaments are united for 0.1-2.6 mm and, if they have anthers (sterile), are free at their apices for an additional 0.1-0.9 mm. !\4thin thesetwo measurementsin each casethere is a continuum. The filament tube is identical in colour to the corolla and is glabrous in all specieswith the single exception of L. berthae rn which scatteredsimple hairs are often present.

Often the free portions of filaments are paired. At an extreme, this results in two anthers terminating a single hlament, a condition frequently observed in Inwrencia glomerata and L repens.

The number of anthersin eachflower variesconsiderably between taxa. The lowest number of anthersencountered is 5, found regularly in flowers of L diffusa and sometimesencountered in specimensof L. glomerata, L, repens and L. helmsii. The highest number observed is

L -l

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The variable fruit morphology enoounteredrn Lawrenciq rs usefully summarizedas follows:

A. Fruit with indehiscantmericarps; pericarp rugoseor reticulate.

l. Mericarps9-11, all fertile;pericarp with apicaland distal walls stellate-hairy,lateral walls coriaceous and rugose or obscurely retiaulate. This condition is found in I berthae. SeeFigure 2D, G & E.

2. Mericarps 5, all fertile; pericarp glabrous, lateral and distal-basalwalls membranous, becoming distinotly reticulate with fenestrate or alveolate areolac. This condition is found in L. but'hananensis,L. glomerata, L. spit:ataand L. viridi-grisea. See Figures 3 (4,6 &1) and4 (4 & 5).

B. Fruit with dehiscentmericarps, pericarps unornamented.

3. All mericarps more or lesswell-developed.

(a) Mericarps 5, all fertile. This condition is found in L. diffusa and L. repens.

(b) Mericarps 2-5, only I fertile. This condition is found in L. densiflora (mericarps 2-55and L. tinerea (merictLrps2-3). See Figure 6F & G.

4. Stcrile mericarps undevelopedand obliterated by developing fertile mericarps

(c) Mericarps 2-5, only I or 2 fertile. This condition is fouid rl'tL. squamqtq(.mericarps 2'3, 1 or 2 feflile), Z. thrysoderma (mericarps 2-3, only I fertile) and L. helmsii (mericarps 2, only I fertile). See Figure 7F & G.

The mericarp is traditionally the most important structure used to determine taxa in the tribe Malveae and keys to the taxa are generallydepend€nt on its charactet. in Lawrentia the number of mericarps, their lertility, development, dehiscence,shape and size, and the vestiture,colour, texture and ornamentation of the pericary are ol great diagnostic value

In number the medcarps ol lawrencia vary from 2 to ll in each fruit. For a particular speciesthis number may be constant or elseof limited range.

ln a given speciesof l4v,rencia all the mericarps may be fertile, or else only one or two mericarps in each fruit produce mature seed. Mericarps containing sterile, aborted or unfertilized seedcontinue to develop more or less normally in all but three species:in these they regularly fail to develop and are crushed and obliterated by the developing fertile mericarps.In the latter caseeach developing mericarp assumesthe swollen irregularly globular shape of the seedit contains.

In those lruit in which all mericarps are well developed these are shaped roughly like a wedge from an inverted pudding basin. There are four surfacesto each medcarp: two Iateral walls, an apical wall, and a continuous distal-basal wall. There is a single, nearly straight proximal margin and two curved lateral margins.

Individual mericarps are either dehiscent or indehiscent. Indehiscent mericarps separate regularly at maturity as their pericarp becomeslight, dry and stiff. ln one specres,Inwrencia berthae, the pericarp is black, conspicuously hairy on its apical and distal-basal walls and coriaceous and rugose or obscurely reticulate on its lateral walls. ln other species with indehiscentmericarps the p€ricarp is alwaysglabrous, white or tan, thin and delicate,becoming translucent at maturity with a pronounced reticulum. The areolaeenclosed by the thickenings

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Map L Distriburion of ihe gcnus ltr1"rcnrrl Phytogeography lawrencia is endemic to Australia, extending from 18o S latitude southwards (Map 1)' It occursin all Statesbut principallyin WesternAustralia whero 11 of the 12 speciesare to be found. six of which are endemic to that State. Its speciescommonly inhabit the marginsol saltIaker. saline depressions and estuaries.

For convenience,in the lbllowing ta\onomic treatment speciesdistributions are indicated by referenceto the sub-divisions jndicated above which are generally adopted by botanists in each State. However. it has beenfound more meaningful in WesternAustralia to summaize these distdbutions in terms of the drainage systemsthey occupy Mulcahy and Bettenay (1972)recognize six drainagedivisions within WesternAustralia. Their classificationis refined Uy n"tte"uy and Mulcahy (1972) and further elucidated by Beard (1973). These divisions und the luk" systemswithin the South West Division are indicated in Map 2 l-qwrencia occurs in all drainage divisions.

Two speciesare peripheral to the Great Plateau. Latr|'renciacinerea is confined to the Great Sindy Deseri and Little Sandy Desefl in the Canning Division (Map 6) L viridi- grlsec occuis along the coastline ol the Monger System of the Sluth West Division,. and of the Murchison, Pilbara and canning Divisions; in the canning Division it extends inland to the Great Sandy Desert, th€nceto the No hern Terdtory (Map 4).

I '(Z/6J),{euouog 1tl6l) prsog pu€ (Zt6t) rqecFl I T' ,{euorpg T ,{qtqnl/'I loUV uo$fs NEoJ qlnos t urals,{Spoo^qcelg t ruolsr(Suo^V Z ltroNds re8uor{ I :uorsl^r([lsel\ qlnos aql q suolsf,so3€urBjd uo$I^r( qlnos uors rc uosrq.rnl^I 'X uorsl^Io elcng g lsqtd s ') uorsl^r( erBqld a uolsr rO {elrequry uorsIAI( 3uruueJ C :e{ersnv ure$el\ ur suo$I^r( .SeureJ( Z del I

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Sevenspecies inhabit the Great Plateauand the coastalpJain fringing its easternmargin. Lawrencia t:hrysoderma is found in the Murchison Division and io the Monger System of the South West Division (Map 8). L. berthae rnhabits the Avon and South Coast Systems of the South West Division, although its distribution is disjunct,reappearing in the Eyre Peninsula of South Australia and extending into western Victoria (Map 3). L. densifloru occurs in the north west of the Eucla Division and throughout the Murchison Division, where it €xtends onto the coastal plain; and there is an isolated collection from betwecn Nickol Bay and the De Grey River in the Pilbara Division (Map 6). L. dilfusa is toLtnd in the far west of the Eucla Division as well as in the eastern part ol the Avon Syslem and in the South Coast System of the South West Division (Map 6).L. helmsii is restricted to the south-easterncorner of the Murchison Division and the western half of the Eucla Division (Map 8). L. repens occupiesthe far south-westernarea of the Eucla Division and the easternpart of the Avon System of the South West Division (Map 6). with the exception of an isolated specimen from Loongana on the Nullarbor Plain, L. spicqta is confined to river banks, swamps and estuarieson the Swan Coastal Plain and South Coast Systems of the South West Division with a disjunction in its distdbution until the Eyre Peninsula whence it skirts the coastsof South Australia, Victoria and eastcrnTasmania (Map 4).

Two speciesare particularly widespread within Western Aushalia and in other mainland states, too. Inwrencid glomerata is more or less ubiquitous in saline habitats in Western Australia occurring in all drainage divisions (Map 5). L. squamata occupies the Canning, Eucla and South West Divisions but is notably absent from the Murchison, Pilbara and Kimberley Divisions (Map 7).

Lawrencia buchananeilsfuis known only from Lake Buchanan in nofih-eastern Queensiand (Map 4).

Despite the fact that the species occupy substantially the same habitats, that their geographical ranges overlap and that often several are found growing together or in close proximity, no natural hybrids or intermediateshave been observed:this suggeststhat they are reproductively isolated.

Taxonomic Considerations In this work, taxa have been defined on the basis of correlated morphological charaaters. Such a procedure is invariably arbitrary in that taxa may be chosen to encompasslarger or smaller units of variation. ln particular, I have attempted to define speciesol Lawrencia in the light of character variation documented for other of the Malveae, particularly in recently monographed genera such as Anisodonteu Presl, Kearnemalvostrum Bates, Melacothqmnus E. Greene and related groups (Bates 1967, 1969) atd Malva;trum A. Gray (Hill 1982 a, b, c). This has resulted in a somewhat broad definition of the speciesin which many localized va.iations are not recognizedformally.

Taxonomic problems certainly remain in Lawrencia below the specieslevel, notably in L. glomerata, L. squamqta arld L. viridi-grisea and possibly in L. densifloro.Tbeir resoluiion would require extensivefield work supplsmentedby experimental procedures,both of which are beyond the scope of the present study. Thus the establishment ol infraspecific taxa has not been uldertaken.

Cytogenetic evidence has not been considered. It is likely that in the wider context of the presentlydeveloping classificationof the Malveae 1But., i908, 1969,Bates & Bianclar

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luarulBa{ clurouoxBl 'lu0rsep Jo ,{lrnburdord lse?8nsol peprdrelut eq,{"ur pspunoJ sl tr qcrq,r\uo s3rtrJelru.lrslecrSoloqdrour aq1 s? r€Josurtr1uo peseq .{11ecr1eue3o1,{qdsr oloq peJaJJouorl€cgrss?lc aueue8e{ur oqJ '.rolaq stuJol1eJeue8 a.rour ur possnosrppu€ a,loqe pelueserdecuapr,le lec6o1oqd:oru eq1 r{1r1(luatsrsuocur aq o1 qczordde sq e^erleqI ;(9961)eJp,rle4 f,q snuutoqtouapg snue8 eql .(,,noleq s? pepSorSesuoeq o^eq snuuoqrcuaps seues oql ;o sercadg sarcedg pu€ €xef aueue8e{ul o1 dey aas) snue8 eq} urqlut seraedsSurdno;8;o su"eu InJasne Surpr,to.rdse Ile,r s€ srel*JeqJ lu"uodurr ,{1pc(uouoxel esaql ur ,{lreueSoreteqaq1 ezrsrqdue ()1Bur^les €xEl cueue8E{ur leroi.3sJo uorluSoceJ aq1 ,\\o[" ol .{lluercglns pel"leJ.roc$ uorlErJBAsrqf 'e€o,l.plll eqrJl eql ur ereq,ltasleuorlruSoce.r ruoua8 JoJ srseqeql u3eq eleq serltnuquocstp 'srepe.rzqc qcrqd\Jo ouros ur fueur ur uorJerJe^loullll g eJaqtolJuahra] snue8qnsaq1 u1 .(^\oloq sorcedgpue exel cueua8er3ulo1 f,ey ees)"cua\le snqtuz!&Dldaql;o ,(pn1sreqgn; Surpued xaluDd srue?qns A\eueql Jo uotlruSoceJpaldruo-rd e^€r{ acueq pup petelerroc ,{ltcuqslp ere 'req se4rnunuocsrpasoql nC lrn{ rlr"a ur s1ed.rec.;:oreqrunu aq1pue ,dncued pu" saqJU"Jq 'eqv 'ruecs 'seuetceu e6$ lueuevJ aqi Jo eJn{lsa| eq, IeJolJe 1o ecuasatd aq1 euryedasJo acuesetd aq1 u! serllnuLJuoJsLptoferu ete eJeq peuruexa sarcedsgo exuanbos eqt urqJ4^

,tapue1 LIZ D0uarhrl Jo uorsr^ed .s.N 218 NuytsiaVol. 5, No. 2 (1984)

somewhat depressedabove, basevariously cuncate,acute, obtuse, truncate or cordate, margin entire, coarselydentate or obtuselylobed throughout or towards the apex only, apex variously acuminate, acute, obtuse, rounded or truncate, often lobed. F/ower.rpedicellate to subsessile or sessile,2-5 in axillary fasciclessubtended by a single lcaf, or solitary and axillary, often in densely leafy clusters on reduced lateral branches or at stem apices to form congested spicateinflorescences, unisexual or bisexual,more or lesssexually dimorphic in dioecious species,usually odourless, rarely sweetly scented. Pedicel\ Iacking, very short and stout, or slenderand up to 4.5 cm long. Epicalyx lacking. Cclyx turbinate, 1.0-120 mm long at anthesis,grcen, sepals 5, connate 1/10-9/10their length, often stronglyplicate in bud, adaxially glabrous, rarely densely stellate-hairy or with a lew simple apical hairs; lobes crect at anthesis, triangular, 1.0-4.8 mm wide at the base; accessoryteeth lacking; apices variously obtuse, acuto or acuminate; nectaries usually lacking, rarely with basal tufts of minute, nectariferous,multicellular papillae on adaxial surfaces;venation reticulate, obscure or distinct with 3-5 primary subparallcl veins to each lobe. Cttrolla campanulate, broadly campanulateor more or less rotate at anthesis,exceeding or subequalto the calyx,2- 12 mm long, membranous, white, yellow or tinged green, red or purple, entirely glabrous or else abaxially stellate- or simple-hairy, rarely adaxially hairy, adnate to the base of the filament tobe to form an ovary cup; petals 5, almost free to their bases or connate up to 4i 5 their length, glabrous or else abaxially simple- or stellate-hairy,rarely adaxially hairy; auricles lacking; lobes imbricate in bud, erect or spreading at anthesis,obovate, Qvate or elliptic, 0.6-?.0 mm wide, flat or more or less carinate; apices variously acutq or obtuse, and entire, emarginate or irregularly dertale. Androeclum ol male and bisexual flowers with 5-30 unranked stamens: lilament tube shorter than the petals and matching them in colour,0.4-7.0 mm long, glabrous or rarely subSlabrouswith scatteredsimple hairs; free portion offilaments 0.2-2.0mm long; anthersdorsifixed, terminal, white, extrorse;androecium of female flowen more or less reduced with anthers lacking or smaller in size, fewer in number and sterlle. Gl,noecium of female and bisexual flowers with a single style branchilS from its base or from up to 7/10 its length from the base into 2-1I branches equal in number to the carpels; style 0.4-10.0 mm long, glabrous or stellate-hairy at the junction of the branches; branches divergent at anthesis,filiform, linear or clavate, rarely broadly clavateand lobed, 0.1-1.0 mm wide, variously white, yellow, pink or brown; stigmasintrorsely decurent on the style brancheswhich are papillate over their whole length or on the upper portion onlyi carpels in a single discoid whorl attached by their proximal margins to a central, apically expanded columella, subtendedby a thin, membranous carpocrater; ovules solitary in eachcarpcl, pendulous,attached to the columella abovethe carpeJnotchi gynoecium of male flowers more or less reduced with carpels lacking or fewer in number, smaller in size and slerrle. Fruit an oblate or ovoid schizocarp, 1.44.1 mm high x 0.8-6.2 mm in diameter, exposed or enclosed by the calyx, readily or tardily shed from the shrunken columella and calyx at maturityl mericarps 2-11, completely to partially dehiscent or indehiscent, free from one another, laterally compressedand wedge-shapedor, if sterile, then often crushed and obliterated by the developing mericarp(s); pericarp conaceous, chartaceousor membranous,black, tan, white or translucent,becomlng rugose, more or less reticulate or smooth and featurelessat matudty, glabrous, rarely stellate-hairy, with Iateral margins sometimesprojected slightly to form narrow wings which may connectapically- distally to lorm a distinctive shield, and the apical region often projected to form a short, acute cusp; endoglossum lacking. Seedssolitary in each mericarp, attached apiaally to the columella by afunicle, triquetrous and basicallyreniform in lateral view or irregularly globular, 0.6-3.3mm high x 0.4-2.5mm in radial width, variouslygrey, brown or black, glabrous, sometimes glaucous with a layer of wax, smooth or more or less rugose, sometimes with angular lateral laces,with only a thin layer of endospermsurrounding the embryo. Columella filamentous, slender or stout at maturity, 1.2-4.5 mm high x <.0.1-2.0 mm in diameter. A genusof 12 speciesconfined to Australia.

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,Dra./r{r'ir}o uorsr^adraPue.] sN Nu''tsiaVol. 5, No.2 (1984)

7. Plant erecti vestiture sparseto dense;leaves conduplicate or flat, distinctly petiolate basal leaves merging into subsessileor sessilestem and floral leaves;dioecious; petals glabrous or sparselyto densolyste)late-hairy; fruit with 2-5 mericarps a single one of which is fertile. 8. Plant to 130cm tall; vestitureshortly stellate-hairygiving plant an ashen- grey appearancelleaves strongly conduplicate along prominent midvein; petalsglabrous; fruit with 2-3 mericarps;seed smooth...... 6. L. cinerea 8. Plant to 60 cm tall; vestiture sparselyto densely loosely stellate-hairy, plant green in appearance;leaves flat with severalmore or lessprominerlt veins; petals stellate-hairyon abaxiai surlace, rarely glabrous; fruit with 2-5 mericarps;seed conspicuously rugose on all surfaces.T.L. densiflora 7. Plant decumbent or prostrate; vestiture sparseto denseor glabrousl leaves flat, distinctly petiolate throughout; hermaphroditic: petals glabroust fruit with 5 mericarps all ol which are fcrtile. 9. Plant decumbent,to l5 cm tall x 40 cm wide; vestituredensely stellate- hairy; anthers 5-10; seedsconspicuously rugose on abaxial surface. 8. L. repens 9. Plant prostrate, to 15 cm wide; vestiture sparsely stellate-hairy or glabrous;anthers 5; seedsmooth...... 9.L. dillusa 6. Erect shrubs; dioecious; stems with spongy periderm producing dry hbrous or flaky bark; leaves sessilethroughout, stellate-hairy or with armourlike covering of peltate scales;dioecious; female flowers with or without stedle anthersl sterile mericarp crushed and obliterated by developing l'ertile mericarp(s); seed swollen and irregularly globular, smooth. (Series Selenothqmnus.) 10. Vestiture of young branches sparse to delse with peltate scales or stellate hairs, or glabrous, rarely pannose with palmate hairs, leavesand calyceswith densecovering of peltate scalesor stellate hairs; thorns often present; secondary branches with leaves in fasciclesbut not densely l€afy or contracted; leaf margin entire or 3-5 dentate towards the apex; 1-2 mericarps lertile in each fruit...... 10.L. squamata l0. Vestiture of young branchesdensely pannose with palmate hairs, dense covering of peltate scales always present on leaves and calyces;thornslacking; secondarybranches densely leafy and much contracted; leaf margin entire throughoutl only I mericarp ever fertile in each fruit. I l. Secondarystems openly branched or looselycontractedlleaves 4.0-12.0 mm long; corollas of female and male flowers indistinguishable;anthers l0-20 in male flowers,5-15 in female flowers; pistils 2-3; style of female flowers 2.0-7.0 mm long, branchesoften brieflv lobed towards the aDices...... t t. t . chryso derma I l. Secondarystems much contracted giving plant a characteristic cactus-like appearance;leaves 1.5-7.2 mm long; corollas ol lemale flowers c. /a length of corollas of male flowers; anthers 5-16 in male flowers, 0-8 in lemale flowers; pistils 2; style of female flowers 0.4-3.2 mm long, branchesalways etrtire...... 12.L. helmsii

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Map 3. Distribution of Iawrencia berlhae.

Subgenus LAWRENCIA

Iawrenr ia Hook. subgenusIawrencia

Plagianthus sect\ot Lawrencin (Hook.) Benth., J. Linn. Soc-.,.Bot.,6: !23 (-1862);E- G. Bali-er, J. Bot. 30: 72-3 (1892). Sida section Inwrencia (Hook.) F. -Muell., Pl. Victoria l: 162-3i1862). Itctotype (iere designated): Pl. spicata (Ilook.) Benth. [l,. spicata Hook]

Prostrate, decumbent or erect sub-shrubs,or erect shrubs;with or without thorns; vestiture stellate-hairyor squamulosewith peltate scales;flowers subsessileor sessile,axillary, solitary, often crowded in leafy fascicles, odourless; sepals adaxially glabrous; petals never with intermeshing basal hairs; filament tubes glabrous; style branches glabrous at their junction; mericarps indehiscent or dehiscent,2-5 in each fruit, all fertile or only 1-2 fertile; pedcarps glabrous, laterally reticulate with fenestrate or alveolate areolae or unornamented; seed triqu€trous or swollen and irregulary globular, smooth or rugose. 'uaor8 'oleurqrnt rleqt €/Z-t/l el"uuoc qcdos:8uol urur 9 61-99 rQz3 selcrcse;djeel ur 'fr"lrlos 'elrssesqns ',{relllxe 'lEnxesrq 'so,^llol pep,rorc uauo ro eJrsses slr,!ofur laloU ur elnJ" 'elnf,B 'se^Eel ,4l,tror.reuSuIuoceq ot esnlqo {lpeo:q se,lrel luseqJo xede I"JoU ur tnoq8norql ojrluo oslo Jo xod€ spr€.,'\ol ,{Iuo aleluap Suuroceq tnoq8norql elrluep ,{lesrzoc se,teal 'eteeunc-elnce 'sure^ 'tsll pseq yo u€reru ol -esnlqo es€q f,EurJd t-l ql^t ]rupslp uorl€ue^ 'apr,r 'ote^oqo sop€lq uru g 0 x Suol uur 0't se ll?urs s€ dl,{orreu .ro cqdqye ,{y,,nor;eu 'ete^o 'epI^\ dl,roJr"u sa^DallD,to /b apon Iuc 0 t x Suol ruc 0 / u€qt ssol {I1ejaua8 1nq 's]"JIeJ 'et"Ao epr,\\ ruc 0 9 x 3uo1 ruc A l l sr e8rel s" uouo ,{1,r,io::eu:o crldqla ,{1,,ro.l:eu 'Jrtd\p '.8uo! 'snorlanbr-r1 'tnols sa^DalIDSDq Io apolq ruc 0t o1 :o 1euo3r:1 a/oltad:elEurunc" 'eJrtue 'reln8uer:1 fy,,no:reuxede urBJEru'apr.t\ruur gg x 3uo1ruru ll-Z ,{l,rorr?u'e{rl -J€ol pu€ ueJr8 salndr$'.se,'eallerou alrssosro elrssesqnsolur 3ur8.reruseaeel pseq eleyoqed 'eleuJetl" '8uqce1 Iltcurlsrp sa,rarT lnoq8norql snoJqelS 1ue1d a.fr7r7sa7crlrporqdeur:eq 'srlrds ,{;eol osuop turoJ ol polcerluo3 surals ,{:epuoces Jepuels pue el}esor 1es"q € 'Surpuocse'lFt rrlo{ SurSre^rpsluels tnots leJe,\asot auo qlr,r urJ 081 ol qruqs-qns

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Etymology.'fhe specificepithet draws attention to the densespicate inflorescence chalactelistic of this species.

Nol"r. In his protologue,W J. Hooker (1840 loc. cit) cites specimenssent to him by "Port Ronald Gunn collected at Arthur, Van Diemans Land; and at Port Fairy, South Australia, growing on the side of a salt-water inlet, where the ground was marshy". Neither these specimensnor any duplicates of them have been found in the course of the present study,nor are theycited by Bentham(1863 loc. cit.).

A specimen sent to J. D. Hooker, Gunn 746 (K), has a note in Gunn's hand appended: "Numirous specimensof this plant collected at various times are packed up separately. They are from Port Fairy, South Coast of New Holland, but the Lawrencia spicata occurs at Flinders Island and I doubt not upon our Nofth Coast although I have not yet seen it. Remove Port Arthur from the habitats as I now hardly feel satisfied about having seen "Mr it from that place".Further, Hooker (1855loc. cit.) notesthat Gunn informs me that the Port Arthur Station formerly assignedto this plant by him is doubtful". Clearly, Gunn 746 was dispatched well after publication of Itwrencia spicata lt is just conceivable that it is a duplicate of Gunn's original collection from Po Fairy probably made between 1836 and 1838 (see Willis 1949). On the other hand it could as easily have been gathered at some later date on Flinders Island. ln view ol this uncertainty it seemsbest to designate the above neotype. Fortunately, L. spic(tlq is sulficiently characteristic for there to be no mistaking it lrom the detailed illustration provided by W J. Hooker which is reproduced here (Figure 3). Mueller ( 1862loc. cit.) publishedthe n ame Sida ldwrencia indicating that it was composed "glabrous ol Itwrencia spicata ar,d L. glomerata distinguishing a erect variety with simple "procumbent spikes"and a or ascendentvelvet-downy branched variety with glomoratespikes". Collections cited by Mueller under this name are referred variously lo L. glomerata, L. spicata and L. viridi-gisea in the presenttr€atment.

A specimen of Lawrencie spicata from W J. Hooker's herbarium and now held at K "sydney is annotated Bynoe." Benjamin Bynoe was a surgeonwith H.M.S.'Beagle'on Kingt 1837-1843expedition to suNey the north-westem coast ol Australia. Bynoe made botanical collections on Depuch Island, the Abrolhos, the Victoria River, off Bass Strait and in New South Wales (Maiden 1906). His specimen of L. spicata was cited tentatively by J. D. Hooker (1855loc. cit.) Subsequentauthors recorded the occurrencaof this species in New South Wales under Plagianthus rplcaras, including C. Moore (1884 loc. cit., 1893 loc. cit.) and Maiden & Betche (1916 loc. cit.), although no further collections from that state are known. Jacobs & Pickard (1981 loc. cit.) consider its presence in New South Walesunsubstantiated, It is possiblethat Bynoe €ncounteredL. spicata at the Victoria River or off Bass Strait where it is still to be found today.

With the notable exception of a single immature specimen collected at Loongana on the Nullarbor Plain, the distribution of Lawrencia Jpicalc shows a pronounced disjunction between south-westernand south-easternAustralia. It seemsmost likely that the Nullarbor specimen is of merely sporadic occurrence from seed dropped by trains, although future collecting from this region may demonstrat€ otherwise.

Despiteits disjunct distribution, there is no morphological-geographicalvariation discernible tn Itwrencia spicata. The geographicalrange of Lawrencia splccta overlapswith those of I berthae, L. dilJisa, L. glomerata and L. repens. L. spicala has been found growing in close proximity to these other sDecies.

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Fig$re 4. Ilrnencia glomenta. L Flower.2. Corolla. 3 Staminal column and style branches.4 Immarure fruit. 5 Ripe mericarp. 6 - Medcarp opened to reveal seed.7 Ripe seed.8 - Lower leaf. Drawn from Drummond 55 (holotype). Reproducedfrom Hooker, Hook. lc. Pl.5i 1.417(i842).

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Map 4. Distribution ol lawren(ia glometta

Distribution. Found in the Avon, Eyre, Irwin and Roe Districts ol the South-West Botanical Province,in the Ashburton, Canning, Carnarvon, Coolgardie,Eucla, Giles, Helms and Keartland Districts of the EremaeanBotanical Province, and in the Dampier and Fitzgerald Districts of the Northern Botanical Province of Westem Australia: in the western half of the Central Australia Pastoral District of the Northern Territory; in each and every Region of South Australia; in the Mallee, Melbourne, South West and Wimmera Study Areas of Victo a; in the Far Western Plains (North Subdistdct) and Western Plains (North and South Subdistdcts) Botanical Divisions of New South Wales; and in the Gregory South PastoralDistrict ol Queensland.See Map 4.

Etymology. The specificepithet draws attention to the clusteredflowers usual in this species.

Nole,r. Bentham (1863 loc. cit.) clearly based his Plagianthus spicat1.6var. pubescens on the second of Mueller's unnamed varieties ol Sida lawrencia ( Laffencia spicatc) discussed above. All of the specimensdesignated here as lectotype and lectoparatypeswere cited by Mueller (loc. cit.) under Sida lawrent:ia and have been examined and initialed by Bentham.

In the dralt of a paper made in the late 1930sconcerning lrwrenciq, the British botanist E. G. Baker described and named two putative new species,which were segregatesof Z. glomerata. These putative new specieswere founded on diflerences observed amongst the 'ale^oqo ,{lpeorq ol el"Aoqo Jo 'etecl"J seturtetuospue cqdrlle o1 c4dr11edl,rorr?u 'Jeeurl 'snorlenbut '.repuols 'eleqrc [Itt'oJJeu apolq :3uol ruu] 0S 01 a/ollad:oleurunJ" ,{l^\olJeu xede 'epr,,ir 'reln8uer4 'o{rT-JEal ur3:eru luur 6 0-t 0 x Suol luru g 6-9'1 dl,trorreu :o urrojrlu pue 'seqcuelq ueer8 safdrls:se^€ol leroU elrssosqnsotur Sur8reruse^eal psEq eleyorled,{ylcurlsrp pacnper uo peJelsnlc ro eleuJelle sa^Dal acue.reedde,{e$-uee€ elqcurlsrp e 1ueld eq1 'sfer 'sJrBr{ 8ur^r8 luocsol€oc,{1pseq ssel Jo eJoru qtlr re el oq1 e1e1e1scrqd:ourourlce pue 'eldurs 'selndrts 4pJJnJrq tJoqs qtr,r\ esoluourol,{lesuep secf,pc;o seJe}ns I"Ix"pe pu" sa^Eel 'sutels 'Surpuacse 'peqcuE.rq-elllrl'lp1ruf, troarnmsa/l crlrpoJqderureq ,{11eo,r,r 00I 01 qruqs '(fU '.1861 ,tfSN'tI'dS'htg :osr:HfUEd :oyoq)gy11 Dpuo7 S'N reqotco 'ErTerlsnv 'pn"t{ urelsed\ turod ol p€oJ uo peelseuoH ErqEprsJ ol lJoulnl lV .s/rd4z 'rnltn8urlsrp elrJEJsnquouru enctdsuol enbsnquog snqrlssescau srlelorledsnqrTas"q srrloJ 'otelled cau olellels otueunpur enb e 'squuts tsuauouoq)nq )uat$t)-I DasttS-lplll uJuabrLrl o]€lncrloJ ourTe,{qrlus"q-rp}srp le rleloltll 'snlrsodruoc arceyord:ecr:od snqrluecsrqapuranburnb srrdJBcr.rerua snlcnlC snlDser\esuap 'snlrporqd"uJeq sD"llals snqr^aJq srTrd snsouEJ ec.redxeln:; tse oasuS-lpt l a)ualt rr-I 'Aou ds'repue-l uasr.rS-rpr.rr,rEIJua.urE.I t

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uJuathtT Jo uorsrr.a)d'Jepuel s N NuytsiaVol. 5, No. 2 (19E4)

Frg'xe 5. Lawrencia viidi-gfisea. Dravn fiofi Iander 1149 '1ue1suol SurureorersrslJe:eqc Jeqlo IF'seruellxe IeorSoloqdroru eseq] uae,rleq '8ur{"ads unnurluoc lse,\\-lseeuE sr eraql dlp€org sartadss({} Jo uorsr^rp-qnscruouox?} eql 'peapur ro.1luerun8re Surledruoc ou eq o1 reedde p1no,r ereql lueJaJJrp,{.re,l reeddz uec uorleuedJo eguel srqlJo seruellxoo,\\l aql ur04 stu"ldq8noqlly ecuereeddeur srsuauouDqcnq 'T Jo sepcs elelyed eqt Surqceordde sruJe tueJseleoc ,{l3uorls qlr^\ sJreq elellels ur parllola orp pu? so^?el ol"doqo flpuorq 01 al"Aoqo uoqs .reeq ot puol arl"Jlsnv pllueJ uo.rJ esoql :,([esEq ]uecseFoc dluoqs ,{luo er" uroql eqlolc leql sJreqelellels al€rp"r eq} Jo surJe aqt puE 'se,ree1cr1dr1le :o crldr11edl,r\orr€u 'Jeourl 3uo1 eaeq erl"Jtsnv ujatso/A ur 'p:eue8 suorplndod lelseoc uror.1sluu 1d uy ernpelrqcre .pel slr pue rrnlrlsel s1tur ,{ltelncrged 'uorteu€A 'sarcN lEcrqder8oeS-l€crSoloqdjouralq€roprsuoc sdeydsrp Dasl.t8-tpt ^ oDuarAoT

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'seunp-puES lelseoJ Jo eprspJB^\eel eql uo 'slelut {alr/rds ls8uoruElueun-uord osl€ sr ll suorsse:depouqes pue sa)I"le{e1d puelut l"tsBoo 'ssuenlse punorE pue ur eddels luepccns ur qros dEIcpue pues eurl€sqnsuo stnxxo tpllqoq ':ee,{ eqt }noq8no:qL pouad 8mn,\\olJ '9 o:n8rg .releruetput urru g g-g g x q8rq 'J?pLreF (rtru gz-gl D auryo) qlpr,{\ IErp"J ur rutu gl-80 x q8rq urur 9Z-60'qtoolus 'u,rojq 'snorlanbul 's8ur.n pa?S luersrqepur prelelplsrp ,ro]JEu ruJoJ()1 pe8uolord u€J"ru 'sece]]ns 'snouEJqrualu'uel l"sEqlElsrp pue IErel"l uo elElncrlerpue luf,cnlsu€rt SunroJeq lo olrq,r di?Juod :g sdr?Juoru:rateu"rp ur rutll 0t-0€ x q8rq urru I t-02'olrlqo InU llt -[/Z reddn uo esoteu8rls,{lesJo.rtur '{urd ro elrq,r 'ruroJrtu'os"q o} ae4'S seqcue]q:8uol ruur 9'S-Zl el,{ls qlr,\\ wnDaou[9 0Z-01 sJeq]uE:8uol ulur 0 l-S 0 sluoruBlg]o uorgod ae.ry '8uol :snorq€I8 ruur 0 f-g l oqnl luau"lu qlr/|l^uru)aotpuv alEurSleuleseJrde :opr,r ulru 9 Z 'ele^o 'et€uuec 'qfua1 -9 1 ,{llq8rlssoqol :snorqElS :reql 9/7-91/1 eleuuocsplad :8uol ulru ',tro1a,{ 'J"ln8ueul 7 9-9 tr apd ro elrq,tr o oto) etrLae,{lp€oJq secrdE:apr^{ ru(ll g Z-0 2 'eteurqrnl seqol lq13ue1rlaql t/Z-Z/l elvuuoc spdes :8uol ruru IS-8[ r.4rJ s3lrrcse] ,{;eel ur pep,rorc ue{o 'fJelrlos 'flel|xE 'alrssos'lenxosrq ,rJ.r,alo1J alEcunll lo esnlqo 'olnc" xede'lnoq8norqte:r1ua {1ue:'peqolg flereru seturtetuos'rodEoq1 sp:earo1 e1e1uep,{Ies;eor Suuroceq,{lpnsn 'a:nuo urSreur'eteeunc-esnlqo ol -elnce eseq'suro^ dJ?ujud snoncrdsuorg-l 'eJnJSqo 'lpu 'f,pr,r qllr lcurlsrp ,4lFnsn serurlarrrosuorleuo,\ urul 97-1 x 3uo1 urur gt-g 'ropuel ,rrrrrr{r-1 Jo uorsl^ed s.N 236 Nul'tsiaVol5, No. 2 (1984)

Herbarium specimensoi I'owrencia viridi-grisea have been relegated to L glomerata in the past.

Lawrencia viridi-giseq is a conspicuouselement of the hind-dune flora along the coastline of Western Australia from Leeman to Broome. Its geographical range overlaps with those of L. cinerea, L, densiJlora,L. glomeruta arLdL. squamata L. viridi-grisea has been lbund growing in closeproximity to theseother species.

5. Lawrencia buchananensisLander, sp nov.

Lav,rencia buchananercisest frutex hermaphroditus trichomatibus peltatis denseobsitus. Fructus e mericarpiis quinque indehiscentibuscompositus, pericarpio facie laterali et distali- basali hyalino et reticulato. Haec speciesl-qwrenciae |iridi-grisea simillima, a qua facile indumento peltato nec stellato, foliis basalibussessilibus nec petiolatis floribusque conspicue majoribus distinguende.

Typus: Lake Buchanan, Queensland,June 1976,J. Gusteen85 (holo: BRI)

Shrub to lO0 cm tall, ascending,hermaphroditic. Vestiture ol stems, leaves and adaxial surface of calyces densely squamulose, predominantly with peltate scales, but also with scatteredsimple, bifurcate or actinomorphic stellatehairs, the latter with more oI lessbasally coalescentriys. Leovesmore or lesssubsessile throughout, alternate or clusteredon reducad branches; strp&/esmembranous, glabrous, linear, c. 2.5 mm long x 0.5 mm wide, margin ciliate, apex acuminatel blade narrowly ovate, obovate or cuneate, 10-30 mm long x 2- 60 mm wide, flat, venation obscureexcept for a prominent mid-vein, basenarrowly cuneate, canaliculate,margin entire, apex broadly acuteto obtuse.F/owers bisexual,subsessile, axillary, solitary, often crowded in leafy fascicles.Callt twbinate, green, 7-0-100 mm long; sepals connaie 2/5-3/4 their length;lobes narrowly triangular,2 0-2.5 mmwide; apicesac\Lle Corolla white, 6.0-8.0 mm long; petals connate 1/4-l/3 their length, glabrous; lobes erect, ovate, 2.0-2.5mm wide;apicesimarginate.,4ndroecium withfilament tube 2 4-4.0mm long' glabrous; free portion of filiments 0.6-1.0 mm long, anthers 20. Gynoecium with style 45-65 mm longi branches 5, free to base or connate for up to l/3 their length, hliform, dark brown' introrsely stigmatose. Fruit oblale, 1.5-3.5 mm high x2.542 rnm in diameter; mericarps 5; pericirp tan, membranous, becoming translucent and reticulate on lateral and distal- basal surfices, indehiscent. ,.teedtriquetrous, smooth, brown, 2.1-2.4 mm high x 1 2-1 4 mm in radial width. Columella slender, I 5-3.5 mm high x 0.2-0.3 mm in diameter'

Fktwering period. Jrne. ". Habitqt. . . forming clumps in lower part of salt meadowsadjacent to lake. Heavily browsed by cattle and roots sought by pigs. Withstands long immersion in salt water'"

Conservation status. I'Twrencia buchananensisis known only from a single population restricted to a highly speciltc habitat where both its above-ground and below-ground parts are heavily UrowieO Uy cattle and pigs respectively.Thus it must be considered a rare and vulnerable speciesaccording to the criteria establishedby Leigh et al. (1981). (holo: Specimensexqmined. QUEENSLAND: Lake Buchanan, June 1976, J. Gasteen85 ShI); I-ate Buchanan, c. 130 miles (208 km) SW of Charters Towers, 1L^S l'aveny 52 (BRt).

Distribution. Found only at Lake Buchanan in the South Kennedy Pastoral District of Queensland.See Map 5. srrossnoasd,{B puesedecspuel errl-uoou ,ur o, ,"0nu"1ffi:lt,.|j#tX"ff{}##I}# 'qtoorus 're1nqo13 ro oso8nJ .(1:eln8errrpu" uallo,lrlsJo ,lrnrJ ,tuecsrqop snoJtenbul paas:peluaureu:oun drecuad loglre;7-1 fluo ro elueJ ll" qc"e ul g-Z sd:ecrreru:etpl€p ro ruro;r1g sa1f,1slluesud s.roqlueelrJels qll/tr sercedssnorceorp;o sre,nog 'sqnrqs-qns elElueJ:suJoqt lnoqtlr ro qlr& :sqnJr{spe.re Jo lcela ro tuequncop ,eteJtsoJd

'[ bryq xo saoNolpLuonbs .(1961) .ueo DrruatuaT I e1yrqa1t1(.brl,{xr s€N) sruDutonbs5 radri; 6-g€€ 'Id '10cr(I .llns prlsnv'seuleg :(/96I) ts :z ]d ]c usc,uosurqrlnH :(9961)g-?I9 :02 'ellr^lehl 'lets 'topu€-I .t\sX snuutoqrcualas ^ou (efir1a1t1) snwuoqlouaps orlles oDual aT s n N I{ YHJ ON E 7 g S uolrra S 'snue8 eql ut sercadsroqlo ou q1r,,ilSulroJB punoJ ueeq seq pue pelelosr dlpcrqderSoet pue ps{col-puel sr slsuauDuDlpnqDrJuatMt/I .sapN 'pu€lsu$nO u:olsee-quou ul ueueqcng e1e1 1e,41uo punoJ ue3q s€q uox"l s({t l"ql lceJ eql 01 uorlualts s,'rre:pleqlrde Jurcedsoql .t8olou1,!g

-DlDids pu9 ,(O) .E l.) 7 (+) D2su3-tpt.t1^7 srsuduDubqrnqDpuar t2-tIo suorrnqursrc del^l

Ltz m)uat.atl Io uorsr€\'Jepue.I SN Nuytsiavol. 5, No. 2 (1984)

Series HALOPHYTON Lawrenciq serlesHalophyton Lander, ser. nov.

Suffrutices prostrati decumbentes vel erecti, glabri vel stellato-pilosi, dioici vel hermaphroditi; flores feminei sine staminodiis; mericarpia omnia fruclus beneevoluta; semrna triquetra, laevia vel angularia vel rugosa.

Typus: L. diffusa (Benth.) Melville

Prostrate,decumbent or erect sub-shrubs;dioecious or hermaphroditic;without thorns; glabrous or stellate-hairyl stems without spongy periderm; leaves distinctly petiolate t-hroughout or petiolate basal leavesmerging into subsessileor sessilefloral leaves;female floweis withoui sterile anthers, style branches filiform; mericarps in each fruit all well developed;seed triquetrous and smooth,angular or rugose

Etlmology. The name refers to the saline soils frequented by speciesin this series

The four speciesin this seriesare extremely closely interelated and display a considerable number of independent reductional series in characters such as habit, vestiture, leaf size and marginal incision,mericarp number, anther number, seedsterility and external sculptunng. They have been much confused in the past. species in this seriesate confined to western Australia and exhibit a striking north-south distribution pattern (Map 6).

6. Lawrencia cinerea Lander, sp. nov.

Suffrutex erectusrlioicus indumento densoe pilis paryis stellatiscomposito cinereus.Fructus e mericarpiis 2-3 dehiscentibus,pericarpio membranaceopellucido non ornamentato compositu;. Speciesnova versimllitir lawren

11116.Near Lake Auld. WesternAustralia, 22" l5'5, 123" 45'E' July 1967'A S' George gl:Z (iroto:PERTH). This sheetincludes male and femaleplants.

Sub-shrub to 130 cm tall, ascending,dioecious. Vestilureol yovrLgstems, stipules, leaves and adaxial surface ol oalycesdensely cinereous-tomentose with small simple, bi{urcate or haplomorphic stellate hairs. I,e.tye.ralternate or clustered on reduced branches, distinctly peiiolate basal leavesmerging into subsessileor sessilefloral leaves;Jl,P,,/e.r green and leaf- iik", soon becomingbrown and scarious,triangular to broadly ovate, 2 5-5.0 mm long x L8-3.0 mm wide, adaxial surfacesometimes glabrous apart from a densebasal tuft of hairs. margin entire, apex acute to obtuse;Peliole stout, semiterete,to 30 mm long; blade tatrowly ovati to ovate,elliptic or obovate,7-35 mm long x 2-23 mm wide, stronglyconduplicate, v€nation indistinct except for a prominent thickened mid-vein, basecordate, truncate, obtuse or acute,margin coarselydentate throughout, apex broadly obtuseto narrowly acute.F/"w?rs unisexual, subsessileto sessile,axillary, solitary, often crowded in leafy fascicles.Cal1r o/ triangular' male flower turbinate, 5.0-7.5 mm long; sepals connate Vz-3,4their lengthi lobes 2.0-3.b mm wide; apices acute. Corttlla of male flower white becoming purple, 5.0-7.5 mm long; petals free almost to their basesor connate for up to t/3 their length, weakly stellate- haiiy on abaxial surface along mid-vein, otherwise glabrous; lobes spreading, ovate, 2.0- 2.5 mm wi<1e;apices narrowly acute to acutel margin entire. Androecium oJ male flower with filament tube 3.1-4.0mm long, glabrous;free portion of filaments0.4-1 0 mm long; anthers 10-20. G1,noe<:iumof male flower lacking. C'al1x of female Jlower turbinate, 3 0- 5.5 mm long; sepalsconnate 1/ 10-1/4their length;lobes triangular, 1.3-2.0 mm wide; apices 'snorlenbul 'Japuols ,ol?rlrJ urur 0I 01 alorlrd:elnJ" xed€ urBrcur ,opr,r urut g.6-E.6x Buol 'el"rl"J 'clldlll3 .o{ll-Jeel ruur 0 h9 Z ueuo ,{l^\ott"u pue ueeJ?sapdls:se^eel elrsses ,ewuJetle I€Jog Jo elrssesqnsotur Sur8rorr se^€al l€s"q e1e1ot1od,{lpurlsrp sa^Dq slr"q et"[ets 'eldrurs ctqd:ouroydeq pu-ltsl"ctnJrq qtr,r ,{ir?q ,{lasuep01 ,{lesJEdssec,{pc;o ocegns prxeqe puE 'seFdrls 'surets ,SurpuacsE.ll"t se^eel lo aln $al snorcoorp urc 0g ot qntqs-qns

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,!rr|rJr{r7 Jo uorsr^oulapuE.I s N --

240 Nu]'tsiaVol 5, No.2 (1984)

long; blade obovate or broadly-cuneatein basal leaves,obovate, ovatc or narrowly cuneate in floral leaves, 4-25 mm long x 2-23 mm wide, flat, venation obscure or more or less distinct with l-5 inconspicuousprimaryveins, basenarrowly acute-to broadly obtuse-cuneate, margin coarsely dentate, apex acute obtuse or truncate. F/owers unisexual, sessile,solitary' often crowded in leafy fasciclesarranged in dense spikes. Calyx oJ mak flower turbinate, green, 3.0-6.5 mm long; sepalsconnate 215415 ther length, adaxially glabrous; lobes triangular, 1.8-3.5mm wide; apices obtuse, acute or acuminate. Corolla of male .flower rnembranous, white, 4.0-7.5 mm long; petals almost free to their basesor connate for up to l/3 their length, glabrous or weakly stellate-hairy on abaxial surface; Iobes 1.5-2.8mm wide, spreading, ovate to obovate; apices obtuse, entire; margin entire or conspicuously crlrate.Androecium oJ'maleflower with filament tube 2.1-3.4mm long, glabrous; Iiee portion of filaments 0.9-1.5 mm long; anthers 20. Gynoecium oJ male .flower lacking, or rarely presentbut stedle and much reduced;style filiform, 2.5-4.0mm long; branches3-5, 0l- 0.2 mm wide, free to bases or connate for up to 1/3 their length, dark brown. Calyx oJ female flower turbinate, green, 2.5-6.0 mm long; sepalsconnate /2-3Atbetr length, adaxially glabrous; lobes triangular, 1.54.5 mm wide; apices obtuse, acute or actmrnate. Corolla of female flower white, 2.0-5.4 mm long; petals connate for 1/5-l/2 their length, glabrous to densely stellate-hairy on abaxial surfaces;lobes erect, carinate, ovate, 0.8-2 0 mm widel apicesobtuse to acute, €ntire; margin entire or conspicuouslyc\liate. Androecium of .female flctwer rnuch reduced with fllament tube 0.5-2.6 mm longi anthers lacking. Glnoecium of female Jlower with style 2.0-7.0 mm long; branches 2-5, free to base of style or connate for up to I / 3 its length,filiform, 0. t-0.2 mm wide, brown, introrsely stigmatose..Fi'ail somewhat oblateto ovoid,2.0-3.0mm high x 2.0-3.4mm in diameter:mericarps 2-5; pericarpwhite, membranous, becoming translucent, dehiscing to releasenaked seed a single one of which is fertile in each fruit. Seed triquetrous, angular on lateral surfaces,rugose on all surfaces, btack, 1.3-2.2mm high x 1.2-2.0mm in radial width. Columellsslender, 1.0-2.5 mm high x 0.1-0.3mm in diameter.Figure 6.

Flowering period. July to October.

Habitat. Occurs or7subsaline sand or clay soils in succulentsteppe and low shrubland around playa lakes, saline depressions,dry watercoursesand on limestone and gypsum ridges,often colonizing disturbed roadsides.

Selected specimensfrom 56 collections exqmined. WESTERN AUSTRALIA: c. 16 km SW of EaraheedyHomestead, R.J. Chinnctck94l,(AD)i betweenMt Morgans& Laverton, C.A. Gqrdner 2441 (PERTH): 19 km NW of Cue on road to Weld Range, NS lander I 125b(BH, BRI, CANB, MEL*, NSW, NT, PERTH); On Giraliaroad, 28 km E of Exmouth- Minilya road, N.S. Laruler I139, (NSW, PERTH); 8 km S ol Wooramel Roadhouse, on North west CoastalHighway, N.S. LarulerI156, (BH, BM, BRt, CANB, K, NSW, PERTH); Nickol Bay (as "Nicol Bay') & De Grey River, April-Sept. 1863,-/8. Ridley s.n. (holo: K).

Distribution. Found in the Ashburton, Austin, Carnarvon, Fortescue and eastern margin of the Helms Districts ol the Eremaean Botanical Province of Western Australia. See Map o.

Etymology. The specific epithet draws attention to the dense inflorescencesolten observed in this species.

Notes. Several specimensheld at BM, K and PERTH have been annotated as Plagianthus gardnefi by E. G. Baker. However, the paper by Baker in which he set out to descdbe and name this putative new species(see notes und.et ltwrencia glomerqtus) was never 9SIl )aPuLfl uro{ u^\erc - JErl rurtnpJ f-sruoqJof I peosodtu I{ qp^\ dIecu.(u Ierelel snouerqurotupu€ ellelunloc ^toqs ol pr^our0r_c,.rEJuJr! - .}F_rJ , .dna Jldurs E r{tr^\ trruJ Jlnlerllull C otnlelutul J fu€Ao Jo ocuorslqop Je{p seqru€rq olf,ls pue €lloror alernoC g Jml Surpuarqns pue io^\ou elErneJ C[ .ouorsrqap row reqtuv 'oqnl .otolftsuap .g J leurulets pue ra^\ou eleu Jo elloroJ - g llqeq lueld apruod y blcu Mt/I .tr\Brl

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'lopue.I ,?rra.r,1.1r,lrJo uors el{ s.N 242 Nu],tsiaVol. 5, No.2 (1984) published, hence the combination is invalid under the International Code of Botanical Nomenclature. Curiously, Blackall & Grieve (loc. cit.) adopted this invalid name for specimens ol P. densrflorus(= L. densiflora) occurring in the region covered by their handbook

Ittwrencia densiflora is remarkably constant in its morphology over quite a wide geographical range. Although it varies somewhat in the extent of its branching, the density of its inflorescence and the density of its vestiture, nowhere does this variation appear geographicallydiscrete or extensive€nough to warrant sub-division of the species

The geographical rar'ge ol lrwrencia densiflora overlaps with those of Z chrlsoderma, L. glomerata, L. helmsii, L. squamata and, L. viridi-grisea. L. densiflora has been found growing in close proximity to theseother species.

8. Lawrencia repens (S. Moore) Melville, Kew Bull. 20: 514 (1966); Beard, Descr. Cat. Western Austral. Pl. 84 (1970); Marchant & Keighery, Poorly Col. & Pres. Rare Vasc Pl. WesternAustralia 6l (1979);J. Green,Census Vasc. Pl. western Australia 67 (1981). Plagianthusrepens S. Moore, J. Linn. Soc. 34: 179 (1899);C. A. Gardner, Enum. Pl. Austral. Occid. 78 (1931): Blackall & Grieve, How to Know Westem Austral Wildfl 2: 345 (1956). Type: Gibraltar, near Coolgardie, Western Australia, September 1895,S Moore s.n.(holo: BM; iso: K).

Plagianthus repens var. pentandra E. Pritzel, Bot. Jahrb Syst' 35: 361 (190\' :fype: Bullabulling, Western Australia, 29 October 1901, L Diels 5204 (iso: PERTH)'

Sub-shrub decumbent, to 15 cm tall x 40 cm wide, hermaphroditic. /errllure oi stems, stipules, leavesand abaxial surfaceof calycesmoderately denseto denselyhairy with simple, bifurcate and haplomorphic stellate hairs. lzaves alternate,petiolate; slrprilescolourless and membranousor green and leafJike, narrowly tdangular, 1.54.5 mm long x 0.3-0.6mm wide, margin entire, apex acuminate; petiole slender, canaliculate, 245 mm long blade cuneateor deltoid,8-10 mm long x 3-12 mm wide, flat, velation obscureor more or less distinct with l-3 inconspicuousprimary veins,base obtuse-cuneate or truncate,margin coarsely dentate, apex obtuse. F/owers bisexual, subsessile,axillary, solitary, often crowded in leafy fascicles.Cal1:r turbinate, green, 4.0-6.0 mm long; sepalsconnate 3/5-4/5 their length; lobes broadly triangular, 2.0-3.5 mm wide; apices acuminate. Corollo white, often tinged yellow, 3.5-5.0mm long; petalsconnate l/5-l/2 their length, glabrous;lobes €rect to spreading, ovate, 1.1-1.8mm wide; apicesobtuse. Androecium with filament tube 0.8-1.0mm long, glabrous; free portion of filaments 0.5-1.2 mm long; anthers 5-10. Gynoecium with style 2.0'2.6 mm long; branches 5, free to their bases or connate for up to 1/5 their length, filiform, light to dark brown, introrsely stigmatose. Fruit oblate,2.24.3 mm high x 3.0- 4.0 mm in diameter; mericarps 5; pericarp white or tan, membranous, becoming transluaent, dehiscing to releasenaked seed. ,.Seedtriquetrous, smooth, angular on lateral faces, rugose on distal face only, black, 1.5-2.2mm high x 1.2-1.7mm in lateral width. Columella sletder, 1.8-2.5mm high x 0.3-0.5mm in diameter.

Flowering period. August to November.

Habitat. Occurs on subsaline sand, clay and loam soils in succulent steppe on the margins of small playa lakesand salinedepressions, less commonly found on lateritic soil in Eucalyptus woodland.

Conservqtionstalus. On the basisofthe singlenamed specimenavailable to them at PERTH, Marchant & Keighery ( 1979) rroled,lhat lltwrencio repens\ttas rcslricted to the Ravensthorpe area. Subsequent collecting and the determination of previously unidentified matedal at PERTH and from other herbaria in the course of the present study has shown this species 'J3leu"rp ur (uru e 0 -20x qSlqtrtru 0Z-gl'repuelsD aLuryo) qtpr,rInrp"r ur ruru tl-90x q8rqruru /l-tl ')3elq 'secEJ 'snorlenbul 'lu3cnlsu"Jt lerelel uo :z1n3ue paa5 peespeleu 3sEelal01 Surcsrqep 'snoueJquou Sururoceq dJecued :S sdrecuaur :relauerp ur ruur 0 t-g Z x q8rq ruru g 7-g 1 'ercIqo 'uJoJrlg 'u,rolq 'epr,\\ 'ses"q tlntl esoleur8rts,{Ies:o:1ur 1q31 :o e1rq,r,r ulur I 0 c 'g 'S rreql 01 ee4 seqcuerq :8uol urur g 7-9 1 e1,{1sq:l,!l. wruJaoul) sJeqtuE:8uol ultu 0 l -8 '8uol 0 stuerrlPlg.lo uorpod ee{ :snoJqel8 urur 0 l-l 0 eqnt tueru€lu qtr,\\ uru)aotpuy 'esnlqo 'et"^o 'Surpeerds 'ql8uol secrdz:epr,r uru g l-Z l ol tcolo soqol :snolq"F rrortl € /Z ol 'a1[],\\ 'eleur\Iln]E eJEuuoJro s3seq rrorlt ol lsoru|? oo.rgslu1ed :8uol lutu S t-0 ollolo) 'reln8uerrl € secrd€:apr,r\ uru 0 t-0 2 ,{lpEojq soqol :seqol eqt uorlcunl eq1]e sJrerlalEllals 'q}8uel Jo ,re] e qtr,r Jo snorqelS,{11urxeqe JreWZlfril ?t"uuoJ slEdasi3uo1 urur Eg-gg 'ueorg 'ot€urqrnt dlpeo;q rf7rz3 selcrJsp+IJeel ur pep^\olc ual.Io ',{rBtrlos',{JellrxE 'elrssesqns 'l"nx3srq 'elnce s./?rlo/J papunor ro 3snlqo xede 'eleluep ,{lesl"oc ur8Jeu 'alerunc-esn}qoJo -a1nc€ 'orncsqo 'lEU 'luelnJcns 'epr,\\ es"q uorleue,\ lBqAleuos utur g 1-9 x Suoyruu g 'protlap 'tr1cJq1e 7 9-6 7 Jo ete,\oqo'elE^o apolq:8uol ulul Sl-Z'et"lncrlEu"c'Japuolsalorlrd:esnlqo 'elerlrc 'epr,r 'JElncrqro ,{lpeorq ro esnlqo xede urSrrru utru y 1-t g x Suoy rulu c Jo 'snouEJqueur t oledo'el€Aof l,uoJJ"u pu€ ssoltnoloJsalnd4s :lnoq8nolql e1elor1ed,{11cur1srp 'eleljjetlc- 'sJrerl saADaTsno:qel3 ,{I1unsnlnq alEIletsrrqd:oruoldeq pue alecrnJrq'eldrurs qllr 'solndrls'sulolsJo snorqulSqnsro ,{JrEq,{lesuepf1a1e:epou sec,{pc;o ecelns 1erxeq"pu" saleel '3pr,\\ atnn\al JtltpoJtldEuueq tuc gJ c ol 'lJ"duoJ 'el"JlsoJd 'qruqs-qns edrlnururo '(aoquaq 7 -IAI['],) 9LZpuowuruO Z're^ru us^\S ')) 'ra^ru '.(opeuolS .I-t) :(ounutoq Z-I lg glz puotuuruO Z u",rs LtI puotuwruO 'f 're^rd ' ,ErTe.rlsnv up,us :sadllotodoltaq () :olJolosr T olcal) tol puoluwnqf T t"p s 'Je,\rd urelse N ue,trg :(poleu8rsepanq) ad{tona7 (ttOt) Sf prcJo I"rtsnv Id runug 'reuprEC 'rolEg V J:(2681) Z/ :0t log f C e:(€981) 061 :l lerrsnv IJ' t]ouegsnsnllrp 'ueerD snqtuDl:lDld(l36l) /9 erl€rlsnv urelse/d Id cs"A snsueJ f :(6/61) 19 erFrlsnv ',{:eq8re; uralseiN Id csp^ er"U serd ? IoJ ,{uood s, lueqcr"l :(0/61) tg Id IErlsnV 'prl?Jg 'oll^lel{ u.retse/d tEJ rrso(l :(9961)flS :02 llng ,\\a) ( qtueg) EsngrpBrJuar^{s1 6

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,7r.rr./.11r7Jo uorsl^al repu"1 sN NuyBiavol. 5,No.2 (1984)

Flowering period. Septemberto November.

Habitat. Occvrs on subsaline clay soils in succulent steppe in and around playa lakes and salinedepressions.

Conservationstatus. On the basisof the singlenamed specimenavailable to them at PER'I'H, Marchant & Keighery (1979) noted that Ltwrenciq dilfusa was restricted to the Fitzgerald River National Park. Leigh et al. (1981)categorized it as rare but not currentlyconsidered endangered or vulnerable. Subsequent collecting and the determination of previously unidentified material at PERTH and from other herbaria in the course of the present study has shown this speciesto be of much wider distdbution. It can no longer be considered poorly collected or rare and would seemnot to be endangeredat present.

Selected specimensJrom 16 collections examined. WESTERN AUSTRALIA: c. 4.9 km N of Lake Cronin Crossroads,R.-/. Chinnock 4136(AD); near Woody Lake, NE ofEsperance, June 1970,A.S. Georges.n. (PERTH); 14 km S of Clear Streak Well, c. 70 km SE ol Norseman,K. Newbey 7575a(PERTH); Hunts Well, Nov. 1891,R. Helms s.n. (AD, K, MEL); YuinmeryHomestead, near Youanmi Lake, Oct. 1981,A.U Mile*-skis.n. (PERTH); FitzgeraldRiver National Park, c.20 km N of Bremer Bay on E side of No.2 Vermin Proof Fence,(PG. Wlson 10163PERTH); 6 km N of Borden, Sept. 1966,P.G. Mlson s.n.(PERTH).

Distribution. Found in the Eyre and Roe Districts of the South-West District and in the Austin and Coolgardie Districts of the Eremaean Botanical Province of Western Australia. SeeMap 6.

Etymology. The specific epithet refers to the prostrate, spreading habit characteristic of this species.

Noler. ln his protologue of Pl.tgidnthus dilfusus, Bentham (loc. cit.), list€d four syntypes, namely Swan River, Drummond 104(K), Drummond 137 (K), Drummond 246 (K, BM) and Drummond 275 (K, BM). These specimenscomprise three distinct taxa which can easily be separated.Of them, Drummond 104 corresponds most nearly to the protologue and has been designated as the lectotype above. Thus the remaining collections become lectoparatypes.However, Drummond 137&246 are clearlyspecimens of lzwrencia glomerata Hook.; likewise,Drummond 275is clearlya specimenot L. berthae(F. Muell.) Melville.

The geographicalrange ol ltwrencia dillusa overlapswith those ol L. berthae,L. glomeruta, L. helmsii, L. repens and L. spicata. L. dilfusa has been found growing in close proximity to eachof thesesoecies.

Seri,esSELENOTHAMNUS

Lewrencia seriesSelenothamnas (Melville) Lander, stat. r'oy. SelenothqmnusMelville, Kew Bull. 20: 514-5 (1966);Hutchinson, Gen. Fl. Pl. Dicot. 2: 53 (1967):Baines, Austral. Pl. Gen 338-9 (1981). Type: S. squamatus (Nees ex Miq.) Melvrlle f=l,svrlsncia squamata Nees ex Miq.l.

Erect shrubsi dioecious; with or without thorns; vestiture stellate-hairy or squamulose with peltate scales,never glabrous; stems with spongy periderm producing dry fibrous or flaky bark; leaves sessilethroughout; female flowers with or without sterile anthers; style branches filiform or clavate; sterile mericarps crushed and obliterated by developing fertile mericarps; seedswollen, irregularly globular, smooth. ,(l) (+) osnJlrp1 pue(O) tuadar'7 6) orollisuap7 DarauDrrrurr&ry Jo suorlnqutsrc 9 det l

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The threc speciesin this section are extremely closely related and carelul attentlon to all the detailsnoted in the diagDostickey is often neededto differentiatebetween them. Two of these speciesare confined to Western Australia; one is widespread, occurring in all mainlandstates (Maps 7 & u).

10. Lawrencia squamata Neesex Miq., in Lehmann, Pl. l-. Preiss.1: 242 (1845). Plagionthus squamqtus(Nees ex Miq.) Benth.,J. Linn. Soc., Bot. 6: 103(1862); C. A. Gardner,Enum Pl. Austral. Occid.78 (1931).Selenothamnus squamolrii (Nees ex Miq.) Melville,Kew Bull 20; 515 (1966);Cochrane et al, Fl. & Pl. Victoriat. 1617(1968); Beard, Descr. Cat. Western Australia Pl. 84 (1970);J. H. Mllis, Handb. Pl. Victoria 2: 381(1973); J. Green,Census Vasc.Pl. WesternAustralia 67 (1981);W R. Barker,in Jessop,Fl. Central Australia 212 & t. 246 (1981);G. M. Cunninghamet al., Pl. W New South Wales492 (l9ll2); Jessop, List Vasc.Pi. South Australia 62 (1983).?l.,pe: Southern River, Perth, Westerr Australia. Preiss1231, Sept. l84l (holo: LD non vidi; iso: MEL). Southern River is a branch of the CanningRiver, near present day 1-hornlie.

Halothamnus mitrophyllus (F. Muell.) F Muell., Pl. Victoria l: 159 (1862);F Mueller, Second Gen. Rep. Gov. Parliamentary Pape^ Votes & Proc. of the Legislative Council 1854-5(A. No. l8): 10,sine descriplione. Plagianthus mk:rophyllus F. Muell., Fragm. Phytogr. Austral. l: 29 (1858)Benth., J. Linn. Soc., Bot. 6: 103(1862); Bentham, Fl. Austral. l: 190(1863); E. G. Baker,J. Bot. 30:72(1892); Bailey, Queensland Fl. l: 110(1899); Maiden & Bstche,Census New South WalesPl. 136(1916); J. M. Black, FI. South Australia,edn 2, 3: 557 & t. 718 (1952);Blackall & Grieve, How to Know WesternAustral. Wldfl. 2: 345 (1956).Lectotype (here designated):Murray River, s.dat., E Mueller, s.n. (lecto: K; isolecto:BM, Mel 98788& 98?89).Lectopat.rtlpes:Spencers Gulf, October 1857,E Mueller s.n.(MEL 98790,98812 & 584132);Spenccrs Gulf nearPort Pirie, October 1851. E Mueller s.n.(MEL 584133);Murray, s.dat.,F Mueller s.n.(MEL 98791,K); Seaflatson the Murray, s.dat. F. Mueller s.n. (MEL 98787);Lake Albert Salt Springs, 1854,E Muallet s.n. (.K); Between Guichen Bay and Lake Albert, s.dat., E Muelkr s.n. (MEL 584131 - material alfixed to sheet only); Guichen Bay, South Australia, s.dat., E Mueller s n. (MEL 584131 pro pte).

Plagianthus incqnus J. M. Black, Trans. Roy. Soc. South Australia 49: 214 (1925); J. M. Black, Fl. South Australia,edn 2, 3: 557 (1952);Blackall & Grieve,How to Know Western Austral. Wildil. 2: 345 (1956).Lawrentiq incana(J. M. Black) Melville, Kew Bull. 20; 514 (1966);Beard, Descr. Cat. WesternAustral. Pl.; J. Green,Census Vasc. Pl. WesternAustralia 67 (1981);W R. Barker, in Jessop,Fl. Central Australia 2ll (1981);Jessop, List Vasc. Pl. South Australia62 (1983).Type: Gawler Range,South Australia,September 1912, J. Whites.t. (holo: AD; iso: K).

Shrub to 100 cm tall, spreading,dioecious, with secondarybranches often terminating in a rigid thorn. Vestitureof branch,esglabrous to denselytomentose with simple, bifurcate and actinomorphic stellatehairs, or squamulosewith minutely fimbriate peltate scales,rarely pannose with palmate hairs, clothing a thin spongy periderm which produces grey fibrous bark on older stems; leavesand abaxial surfacesof calycesdensely tomentose with simple, bilurcate and actinomorphic stellate hairs, or squamulose with minutely fimbriate peltate scales.lzaves denselylasciculate on short lateral branches,sessile throughouti J/Du1e.tbrown or pale green, membranous,narrowly to broadly triangular,falcate, 0.4-2.1 mm long x 0.2-2.3 mm wide, vestitureof abaxial and adaxial surfacessparse to densewith peltate scales or palmate hairs, margin ciliate with stellate hai$, apex acuminate or actte, blade linear, narrowly elliptic, ovate, obovate,spathulate or suborbicular,{lat or conduplicate, 1.6-19.0mm long x 0.5-11.0mm wide, abaxialand adaxialsurfaces dense with eitherstellate hairs or with peltatc scales,or with palmate scalesat the base merging into peltate scales, 'eprclepv ' 'JJnlg fJ Jo gN rur{ 0l c ed I"foU :((V) 6t qIwS 7J tBunl.IEoN ld '"eplo 'dpad suou!\d :(CV) BOgguolqlo1 N U,J Jo ,^Aurl 0Z c :(QV) gVtt u7tlltol W yJ 'epl"l3pv 'lelC raqooJ Jo ,,!\ rut 9 I r :(CV) 816 lqanquaqaotx 11O Jo AN Ix{ E6 c '€ue,\\tlq/K srellB/d :(CV) 9001 1M1 Ur pu" uoJnsog uee,ttaq'e{"'I puoruEl( :(CV) l00tl nllplg 'lH 'pBor "uelleg-fldo3 uo '.{a1do3 UDL c :VI'IYU1S1V Jo ,465 '\r)lg) HJ1OS (7 6sLalund 'E 's 'peelsrtuoHurLc qlnos uDt :cNV-rsNEEnb ,a'u ,tt .94 ,l I o8z '('Mht 'x) lo 3N 'o'YI 'g 's 808stowlN ,09.Ie I ,09"zz 'e8pe,^A "IA},,iI'Hg'(V) rI,\{ I€rtueJ Jo,^[ (tu{ 9't) etllu J r:(HIdEd IN L6LVz1o7 'Xd'g,,gZ 'S,l.Z 's8uudg 'CV) "SEI .96 elsnoqpq :(J-N 9169zloT Xd'A,tZ "(,tl'S 'E 'S '.,ue^€q,,lreN,, ',,,{qreddeN,,:kIv) 4rI 'Xd :[tSN ,t? 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'rapue.I ,lJ!./rl&r7 Jo uolsr^ou SN 24s NuytsiaVol. 5, No.2 (t984)

Smith 1127(AD); 9 miles (14.5 km) SW of Malinong Hall, MC.R. Sharrard 1224(AD); DalhousieSprings, c. 80 miles (128 km) N of Oodnadattain the vicinity ol Spring Hut, D.E. Symon 3251(CANB, K); Shell Beach,lnnes National Park. J.Z. l4/eber4229 (AD); Head of Bight, PG. Mlson 162l (AD); St FrancisIsland, c. 60 km SW ol Ceduna,N.M. Wece 136 ( AD\. VICTORIA: Raak Salt Plains,9 miles (14.5 km) WNW of Hattah, l.C. Beauglehole 40583 (MEL); near Kiatta, R. Melville 988 (K, MEL); Wimmera, s.dat.,J. Dallechy s.n. (K); nearAnnuello, Sept. 1971,R. Iluades.n. (MEL). WESTERN AUSTRALIA: 10 miles(16 km) S of Leonora on road to Menzies,TltFl. Aplin230( (PERTH); l7 miles(27 km) N of MeiroseHomestead, "/. S. Beard6534 (PERTH): "Brooke') lsraeliteBay, 1885,,t TC Brctctks(as s.n. (MEL); 6.5 miles (10.5 km) W of "Yelma", '772 Winchester, s.dat., C. Chapman s.n. (PERTH); Bern Pool, R."/. Chinnock (AD); CanningtonSwamp, WattleGrove, R. Cctveny8258 (NSW, PERTH); c. 30 km NNE of Stokes' lnlet, Hj. Eichler 20322 (AD, PERTH); Lake Auld, A.S. George 9131 (BRI, MEL, NSW NT, I'ERTH); 13 miles (20.5 km) from ChesterPass down South Stirlings Road, Dec. 1964,A.S. Georges.n. (PER1'H); 12 km NE ol Norseman,K. Newbey 8095 (AD, PERTH); Rawlinna, PG. Mlson 5798 (PERTH); I km W of Meckeringin flood plain of Moftlock River, P G. llilson 6403(AD, BH, BRl, MEL, NSW' NT, PERTH). N end of Lake Moore, PG. Mlson8635 (K. PEITTH).

Distribution. Found in the Avon, Darling (Drummond Subdistrict),Eyre, Irwin and Roe Districts of the South-WestBotanical Province, and in the Austin, Canning, Coolgardie, Eucla and Helms Districts of the EremaeanBotanical Province ol WesternAustralia; in the western seotor ol the Central Austraiia Pastoral District of the Northern Territory; in the Eastern,Eyre Peninsula,Flinders Range, Gaiidner-TorrensBasin, Lake Eyre Basin, Murray, Northern Lofty, Nullarbor, south-easternSouthern Loliy and Yorke Peninsula Regionsof SouthAustralia; in the Ballarat,Mallee, Murray Valley,South West and Wimmera Study Areas of Victo a' in the Far WesternPlains (North and South Subdivisions)and WesternPlains (North and South Subdivisions)Botanical Divisions ol New South Wales; and in the Maranoa PastoralDistrict of Queensland.Sce Map 7.

Etymology. The specific epithet relels to the densely squamulosc vestiture of peltate scales found on the stems, leaves and calyces of the type form ol this species,giving the plant an ashenappearance.

Nolen. Like lltwrentia glomeruta (see above), L. squamatu is a widespread and highly polymorphic speciescomprising many aliopatric populations. Nonetheless,it is rather surprising that specimensformerly referred to L. incana and L. squctmatashould have been located in differentgenera by Melville (1966).Barker (1981)has drawn attention to this bewilderinganomaly. To be sure, branchingpattern, leaf shapeand size,thorniness, and hair type are extremelyvariabie, but examinationof all availableherbarium specimens in the courseof this study revealsa continuum in each of theseaharacters. Specimens with predominantlyactinomorphic stellate hairs usually have longer,broader leavesthan those with predominantlypeltate scales, and they have a generallymore northerly distribution. Plants at either extreme can look strikingly different, and in the past L. incanu and, L. squqmqta sensu stricto have been distinguished on this basis. However, in their essential characters of flowers, lruit and seed these and other forms are remarkably constant: in keepingwith the principlesexpressed earlier in this paper,I am unwilling to maintaintheir separation,even at an infraspecificlevel.

Lawrentia squqmqtqis the only speciesin the genusto possessthorns. Theseare not invariablypresent and their presenceor absenceseems not to be geographicallycorrelated. R. F Parsons(pers. comm.) has suggestedthat small-leaved,non-thorny plants and large- 'stueld olBureJpue .V '6961 ,E elertt r{toq sepnlcurteeqs ed.{1o1oqeq1 (H-tXSd :otoq) 918 Ia4{/bS A reqotco 'S '€rTerlsnv 'e8ue.11 ,sued,{e13 ,tf .l.ll 788 o9Z uro1se16 ple/d Jo puo 16 r11rgqpg :sndr(.; ':nlrn8urlsrp srqd:oulp :olqenxes ceu snqrurloJrunanbsqyo.roc snqrc,{1ec 'snquoferu 'egede sr11o; sr8eul ououaagtuel €nb e 'euttllt-urs llsLqal! au)uat&D7 seraeds 'snlrsodruoc Je€H etuecnlsu"rl o3c"u€lqueu otdlEcuad snqrluecsrqepE-Z strdl"Jueu o snpnrC etuotsrsuocs4e11ed s1l1d a aued ord eu8Eul oluerunpur ,snqrlu€ruJoJsolerloJ esuop selneJsrlcerluoJ urnluenbtle srrrcpunJ3sstueJ sncrorpxoln{ lse olunposttq) DDuaJhzal 'Aou 'repue'I ds Buuopos,{rqJBrJuaJ^{BT J I 'sercads reqlo eseql o1,41rurxo;d esolc ur '7 'stsuauDuDq)nq Sur,tror8punog ueaq suryolowonbs Z Jo uortdecxa eq1q1r,r snueSeq1 ut sorceds:eqlo IIe Jo esoql qtl,,lrsdelJ3^o oloLuonbsDDu intr[ Jo a8u€r lucrqde]8oe8 aql urerede qrns pe^,3sqo loue^eq r '],]#,1"'J#ffiii'ii:,"JtJ";:it""iff"':;,'ilx};,,]: sr srql tslql(\ s|os snoosd,{3-uou',{ru"ol SurlrqEqurJ3l1"l eqt pus slros snoesd,{8,,{pu"s 'pet"8er3os .7 ,pe^€ol Surddncco:aur:o1 eqt rllrl ,{lecrqdepeaq feur apwonbs 1o s1te1ddulorlt

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,rrlrl&r7 Jo uorshel{ repue'I s N 250 NultsiaVol. 5, No 2 (1984)

Shrub to 100 cm tall, spreading, ramulose and much contracted to form dense lealy stems,dioecious. Vestiture ofbranches, yellowish,densely pannose with palmatc scalesclothing an exceedinglyspongy periderm which producesconspicuous dry flaky bark on older stems; leavesand abaxiil surfaceol calycesdensely squamulose with fimbriate peltate scales.Z€ave,! sessilethroughout, denselyfasciculate;sllpales pale green,dry and spongy,narrowly triangular, 1.5-7.0mm long x 0.5-2.3mm wide,laloate, abaxial surlace densely squamulose with palmate scalesat the baie merging into peltate scales,adaxial surface almost glabrous or with dense vestiture of palmate scal-s on ihe mid-rib merging into peltate scalestowards the margin and apex, margin ciliate with simple, bifurcate and haplomorphic stellatq hairs, apex acuminate; blaii ovate, narrowly obovate or spathulate,4-12 mm long x 1.7-30 mm wide, conduplicate, reflexed, abaxial and adaxial surlacesdense with palmate scalesat the base merging into peltate scales, base narrowly cuneate, margin entrre, apex acuminate' acute ot o-btri". F/owe'rsunisexual sessile,axillary, soltary. Call'x ofm?/e.flotreru turbinate' green' 3.4-4.5 mm long; sepals connate 3/5-4l5 their length, abaxial surface densely squamulose with palmate scalesmerging into peltatescales; lobes triangular, 1.0-1.5mm wide; apices acute to broadly obtuse. Corol/a of male flower white, 3.5-5 0 mm long; petals lree almost to base or connatefor up to 3/5 their length, glabrous;lobes spreading,ovate,0.8-20 mm wide; apices acute, obtuse or truncate, irregularly dentate. Andrttecium o;[ male Jlower with filanent tube 1.7-2.5mm long, glabrous;free portion of hlaments03-1 5 mm long; anthen l0-20. Gynoet:ium of male.flower very much reduced and sterile, sometimeslacking. th€ir length, Calyx of female Jtottel tu rbinate, green,3.3-5 5 mm long; sepalsconnate I /2-4/5 abaxial surface denselysquamulose with palmate scalesat base merging into peltate scales, adaxial surface glabrous; lobes tdangular, 0.8-2 0 mm wide apicesacute to broadly obtuse to base or connate Corolla of female ./owers white, 3.0-5.0 mm long; petals almost free for up to ji5 theii length,glabrous; lobes erect,ovate, carinate' 0.8-2.0 mm wide; apices obtuse to truncate, entire or irregularly dentate.Androecium ol Jemale.flower with lilament tube 0.7-1.6mm long, glabrous,free portion of filaments0.1-0 9 mm long; anthcrs5-15, stetlle. Glnoecium of female flower with style 2.0-7.0 mm long; branches 2-3, free almost to base of style or connatefor up to 7/10 of its length' filiform or clavate,0l-1 0 mm wide, apices often briefly-lobed, introrsely stigmatose. Fruit ovoid' 2 3-4.0 mm high x l 8- 2.5mm in diameter;mericarps 2-3; pericarp white or tan, mernbranous,becoming translucent' dehiscing to releasenaked seeda single one of which is fertile in each fruit, sterile mericarps crushedand almost obliterated.Seed swollen, irregularly globular, smooth' brown' l 5-2 6 mm high x 1.1-1.5mm in radial width. Columellafilamentous, c. 2.0 mm high. Figure '7

Flowering period. August to .Ianuary.

Habitat. Occlrs on subsalinesand or clay soilsin lightly woodedsucculent steppe on margins of playa lakes, particularly on gypsum ridges.

Specimensexamined. WESTERN AUSTRALIA: 44 miles (70.4 km) NW of Cue on road "Modoonga", t; Mileura, TE.H. Aplin 2531(PERTH); 26' 56' S, 117" 36' E, J.S. Beard 6612 (PERTH): EneabbaFlora Reserve,Oct. 1961,C. Chapman s n (BM, BRI, CANB, K, NSW NT, PERTH); Salt lake SW of Mnchester, J. Coleblt-Mlliams 117 (PERTH); SaltlakesWof Mnchester,J. Colehy-llilliamsll8 (PERfH): 9 km lrom Winchestcralong road to Eneabba,0.5 km S of road, S end of Yarra Yarra Lakes, M.D. Cnsp 5469 (CANB); "Barnong", A.LI. Humphries P48 (PERTH); N of Weld Range, opposite Madoonga Homestead beside road from Cue, N.S. I'qnder ll22 (K, MEL*, PERTH); N of Weld Range oppositeMadoonga Homesteadbeside road from Cue, N.S Itnder 1123(CANB, "Puttingup'), "Roderick", PERTH)I locality unknown (as s.dal, G Maxwell s.n. (MEL); c. 150 km N of Yalgoo,A.A. Mitchell 968 (MEL*, PERTH); Billi Billi Clavpans,W end "Barnong", of Weld Range, 26" 58' S, ll7" 33'E, R. A. Saffrey 816 (holo: PERTH); April (adrloloq) 9l8,lr4&s tuo{ u^\?rc 'sotuoqiul raln^o paesposEaler pur aluals pue dftruau paqsnrcSurul?tar trn{ tu.ds 'ellorot H pe.1.qlr/\ $u3csrqrp ssarord uI tlnlJ ernl€ru pue elloro. por.qluA ol^ls g lo^\ou C Jo 'to?ld C oteuoC (l r!^\ou .teur Jo €lloroJ J JeaT g tlq€q at€l,t V Du)aporllqc DDuaraq L :.lnSt:I

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Nultsia Vol. 5. No. 2 (1984) 252

1954.D.G. Wilcox s.rL.(PERTH); N end ol Mongers Lake, 28' 46' S. 117" 22' E, PG' Mlion 8604(PERl-H).

Distribution. Found in the lrwin Dist ct of the South-WestBotanical Province and in the Austin District of the EremaeanBotanical Province of WesternAustralia. See Map it.

Etymology. The specific epithet draws attention to thc dense yellow vestiture rrf stellate hairson the young branchescharacteristic of this species.

Noles. A mixerl collection of Lawrencia t:hrysotlerma and L. squamata communicated to "Puttingup." Ferdinand von Mueller by G. Maxwell is annotat€d 1-his locality is cited by ( | 1375)und er Plagidnthusmicrophy llus L. squamata). A thorough searchh as failed M ueller I "Puttingup" to reveal that any place name exists in Western Australia with the spelling "Put..." and no aboriginalplace name anywhere begins (NomenclatureAdvisory Committee, Officeof the SurveyorGeneral, Perth, pers. comm.) From the handwrittcnlabei it is possible "Pullingup" that "Pullingup" was intended.lhere is not a in WesternAustralia, but there is a Pulingup Spring near Pallinup river at 34" 18'S, 118' 23'E Although L. squamata is to be found in that vicinity today it is far removed from the presentrange of L. chrysoderma: it would seem that the attribution of Maxwell's specimen of the latter speciesto this area is in error.

The geographical range of llwren(:ia chrysoclermaoverlaps with those of Z densiflora' L. glomeruta, L. helmsii, L. squanata and L. viridi-grisea. L. chrysodernta has been found growingin closeproximitl lo all the.eipecies.

12. Lawrencia helmsii (F Muel]. & Tate) Lander, comb. nov. Plagianthus helnrsii F Muell. & Tate, Bot. Centralb.55; 316 (1893);Trans. Roy. Soc. Soulh Australia 16: 339 (1896); Anon., Proc. Linn. Soc. London 1908-9:9 (1909);Morrison, J Nat. Hist. & Sci. Soc. WesternAustralia 3: t.2 (1907);C. A. Gardner,Enum. Pl. Austral.Occid. 711 (1931)l Blackall & Grievo. How to Know Western Austral. Wildfl. 2: 346 (1956). Selenctthamnushelmsii (F. Muell. & Tate) Melville,Kew Bull. 20: 515 (1960);Beard, Descr.Cat WesternAustral. Pl.84 (1970);Erickson et ai., Fl. & Pl. WesternAustralia 154 & 1.487 (1973)iJ. Green, Censusvasc. Pl. westernAustralia 67 (1981);A. Mitchell,in B. D. Morley & H R. Toelken, Fl. Pl. Australiat.Tgc&d(1983).I2(tott)Pe (herc designated): Lake Lefroy,Western Australia, 7 November 1891,R. Helmss.n. (lecto: AD; isolecto:K, NSW),981553261).LectoparqtfPe: l,ake Annean,October 1893,S. Drron s.n.(AD).

Sftrub to 100 cm tall, ascending, ramulose and much contracted to form dense leafy cactuslike stems,dioecious. Vestiture of branchesyellowish, densely pannose with simple, bifurcate and haplomorphic stellate hairs and palmate scalesclothing a spongy periderm which produces dry flaky bark on mature exposed branches but normally obscured by denseleafy lascicles;leaves and abaxial surfacesof calycesdensely squamulose with minutely limbriate to entire peltate scales.Z"oveJ sessilethroughout, denselyfascicuJate; stipules pale green,dry and spongy,narrowly t angular, 1.2-3.5mm long x 0.9-3.0mm wide, falcate, chartaceous,abaxially glabrous or spa$ely to denselysquamulose with palmate scales,often becomingglabrous towards the margin, margin ciliate with simple,bifurcate and haplomorphlc stellate hairs, apex narrowly acute to acuminate, blade tarrowly obovate, elliptic, ovate or orbicular,L5-7.2 mm long x 0.7-2.2mm wide,conduplicate, reflexed, vestiture on abaxial and adaxial surfacessparse to denseat base with palmate scalesmerging into peltatesscales on lamina, base narrowly cuneate, margin entire, apex acute to broadly obtuse F/owers unisexual, sessile,axillary, solilary. Calyx o;f maleJlowers turbttate, 1.5-45 mm long, green; sepals connate at least 4i 5 their length, abaxial surface densely squamulose with stellate 'g de1r1eeg "II"JtsnV urelse1\ Jo acurlord 'uounqu$I(J IEcIuEtog ueeeruaJg aql Jo stculsr6 erpreSloo3 pue urlsnv eql ur punoC

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,rr,/aJdlrTJo uols$el{ repue'I s N 256 NuyisiaVol 5, No.2 (1984)

The primirive hair type in La\rrencia is takar. to be the haplomorphic, multicellular stellate hair common to all but the glabrousspecies in the genus.Modification of this basic hair type hasapparently proceeded in severaldirections (Figure l). From the basichaplomorphic stellatehair (Figure lA or D) can be derived the bifurcate hair (Figure 1B), sirnply by reductionin the numberof rays;further reductio n leads to a simplehair (FigureI C). Planation and fusion of the rays of the basichair type result in actinomorphicstellate hairs (Figure lE) and, with completefusion, peltatescales (1F & G). Subsequentreduotion of the lused rays of peltatescales, coupled with a changein orientationof the hair, producesthe unique bract-likepalmate scales (Figure lH & I) observedin the seriesSe/enolr/rr?rut Continuation of this plocessproducts a sin.rplehair (Figure 1J). Neitherpeltate nor palmatescales ocour elsewherein the Plasianthus alliance.

In Latrrenci{t, the protective role usually attributed to plant hairs is probably three-ibld. Firstly, hairs effectwatcr economy by restrictingmovement ol air near slomata and thus Ieducingtranspiration. Secondly, hairs may preventlissue damage from high incidentlight by vi ue of their great rellcctivityand by providing an insulatinglayer' Thirdly, they tnay help to avoid damagefrom salt-sprayby preventingdroplets ol sea-wateror salty lake- water from reaching iive tissues. HaiIS Qn the flower buds and solt vegetative parts of Lav,renciamay aisodiscourage insect predation.

l-he intermeshing basal hairs on the petals of male flowels of Lawrencia berth.reassociated with the sepalineneotaries of that speciesare discussed beiow.

The bristlelike bairs on the persistentmericarps of Ll\t rencia Dellh.rcprobably assist their dispe$al(see below).

Z.,aveJ.Variation observed in the leal-characten of Lawrent'ia lalls well wilhin the limits recoriledfor other membersof the ifibe Malvcae(Hutchinson 1967, Bates 1968a).

Although stipules are usually present in the Malvaceae (Cronquist 1981) and in the tribe Malveae, throughout the Plagianthus alliance there is a trend towards stipule reduction and loss(Bates 1968): Glnalrrx Alel.,like Ltwrenciq'has conspicuous and penistentstipules; stricto and Hoheria have stipules which arc small and caducous; Plqcianthus seDsu 'l'he Asterctfi(,hionis exstipulate(Melville 1968). considerablevariation observed ln texture, vestiture and size of the stipulesof Lawrencia is striking

Melville (1966)and Bates(1968) have noted the trend towards palmati-pinnatevenation in the genera ol the Plqgisnthus alliance. ln the present study venation rn Lqvyrenciawas observedto be actinodromousthroughout, but with a marked tendencytowards reduction ol the lateral primary veins with only the mid-vein really distinct or r'r'iththc renation quite obscure.Leaf bladesin the intermediate category ce ainly har,'ethe superficialappearance df being pinnate:on closerinspection, laint lateralprimary veins radiatingfrom a central point towards the blade base are invariably revealed. Leaf venation is variable even within ipecies and no overall pattern it I'av)rencia was discerned.ln other generaof the Plagianthus alliance reduction of ihe lateral primary veins has proceeded further than in lQwrenciq with the deveiopment of stronger secondary veins from the mid-rib giving and is coupled 'lhis a distinctly palmati-pinnatevenation. syndromeis most advancedin certain species ol Hohe a \ hichappear tu ha\efrinnate \enrlion

Heteroblasticleaf developmentsuch as that observedin Lawrcnciais widespreadin the tribe Malveae. Elsewhere h rhe Plagianthus alliance it has been reported in Plagianthus and Hoheriq(Allar' 1961,Melville 1966,Salmon 1980). 'eauB\I! snqluDldold eqt ur pea:dsapr,t oq ol sruoesuorlrpuoc srql crldqlo ro ol€Ao d[oraru uouo or€ asoql (selcedssnorceorp Jo sJe^\ou epru ,{lpnsn) aseq aql ol ee4 tsorule sletad qtl,r esoql ur 'le,\e,roH ue,re'serceds,{u€ur ur ele^oqo flFcrsaq ocueq pus elppru eq] 3^oq? lsop€orq sI ',{1nueJ leql eql Jo I"crd,{t l"ql ot urroJ uI puodsaJroc o1 ruees qeled aql olJuahvl uI 'suorttuuep crreua8ur pe,(oldureueeq e,reqBIIoJoc orll Jo 'o€aJBAI"I suorlerr?d SuDIuls eJotu ,reJ E l eql ur ,4cu?lsuoce,rllelar slr al\dseq qlorc) 'snqluol8Dld puP.uoqoE ar? polotl uaoq seq aDquaq a)uat&try Ja l"q} 01 r?Iurls runJJed " qcq,\\ .roJecue\le snquo8tild eq1ur ereue8reqlg '(.to1eq aes)serceds srqt Jo .{ft}cou eql qll^\ peprcoss" eq o1 rueddr aoquaq 7 Jo sre^\ou e1eruJo soqol elloJoc eql uo srleq IBs"q Surqseur:e1u1oroq eurrl tsrr1eql ro; pelroder sr rrllzatD pue aDlluaq Z qloq ur ocuesardrlaql ',{lrrueJ eql lnoq8norql peepur pus o€o^l€n aqul eql Jo Breue8 ,{uetu uI uolllsod slql uI punoJ ore sreq Surgerces-:epouqanoqtlv (996I elFIeI,{) uolq)upntsv ur pue 'sro,rou e1ertreJpull rptrr qloq ut pa,\resqoeq uec deql eJeqtd'nrtDutg ur ,(po ecuelle srulluotSold eql ur eJeq,reslepunoJ eJ? aoquaq 7 Jo spdes eql Jo eseq aqt 1" ec"J.InsIErx"p€ eql uo 'reFllecqlnru 'Jeqund punoJ esoqt ol oou€Jsodd?ur rplnurs azpded snore3uepeu Jo sun] '(8/61 '.9I6I '196t ur,tul d' e.Ioo],{ slunJ'9961 slll^lol{ uellv) oualloH pue snqluDlSold 1o sercedsur se psxegol ,{l8uoj1s ro Surppards,{leplr 8u[uoceq ueql ]eql"J 1mr; pauedu 'ouaqoH eql Sursolcue,{lunl?u] le lcora ul"rueJ seqol slr pu" Jo sercedssuo ur pelroder 'acueqle qlool ,fuossacc?eql s{cEI lr snqtuorSol4 3ql Jo ?Jaus8Jaq}o ur peuodor se :zlndnc 'J€lnqnl .ro alelnuedruec Js^ou pue cruotqo ro ot"urqJnl ,{lpeplcep s u)uahvl lo x[le.) eqt laql sr e:oq tueuodurr sI t"qi6 elJesep ,{eql uol}uelle eql ue,rt3 ueeq 1ou eleq f,eql 'oueqlrH ecuecuru8rscrreue8 ;o {1:eolc ore x,{lec eql Jo sJepereqt $uerlle snqtuo8Dld eql ur 'eee,tpyq equl oql ur (6961) seteg ,4q pe,lresqo ,(cuepual preue8 eql ol f,rerluo3 'sorceds qc?oJo uorlezuolr?r?qc 'su?8ro eql ur ecuzuodrur ?runss? leqlo Jo sJepEJ"qc eql uI uorl"r.rE^ qlr^\ uorlcunfuoc 'qrrq,r ur paJeprsuocuoq,r sacuereJJrpII€Je^o eDqns ur ,{po j1es1rslseyurru xf,pc eqt;o sarnl"el Jeqto pue eseqtur uor]erJerl q13uelaqol qlr,t orpr s1 pu? qfuol xdpt lelot aql ore oDuattvl ur sJep"r€rlc 1n3esn,{1e:eua? eJou eql lstuoruy ecuerlp snqtuotSold ?tll ur eleq,resle pe,rresqourelled eql ol surJoluoc,rruatutry KSoyorqdtourxfleJ Jrs?qs!1: ul xtloJ

'PaArJeps"A\ ecu"rTl" snLlluD!?Dldaql selelnlsod eq qctq.r tuo4 eaue\Ie uolunqv eql Jo slsqluelu 3o pcrd{} st uoBrpuoc slt{t lerlt sslou puB aqrJt at{l uI o,{rllurud xdl€clde u" Jo ecuesqeeql sJeplsuoc setsg oBe^l"I^troqrrl aqt ur (896I) seleg fq pazu?oceJ secueu" eql eqrJcsuncrrJ ot pu" sequlqns llturTep(]1 pesn ueeq s"q erntcnrls srrltJo ocuesq€ro ecueserdeqf 8ul{c€l fleJllua 'eJu€rlle 'a)uahell sr x.{pcrda ue srultuDtSoldeqt Jo e]eue8 Jeqto llB ur se u1 x,(1oc|dg 'D!)Uat^rrJ lo sarcodsSuruuulor eql Ip ur slun ecuacserouurcrseq se ellos qcq,\\ sJaA\oU,{repxz e13urs eql ol uorlrpuoc asorudc prlsecue ue ruorj Surprel serJesuorlcnper s ur e8els leuorlrsu?Jl e lueserder KlqBqoJd,aDquaq DDuaihtj Jo solJl'seJ ,{rEIIrye snopuoue dlSuruees eqI 'suose:sSurnol8 rraql ur ,{lree sre,rol} ,{J€llrx" ,{Jelrlosacnpotd sn4tuu3o14 pue Dt.tatloHqloq secueJsoJouureJnlBru rreql;o ,{lxelduroc oq1 elrdssq solclcs",JpuE seurfc'selclu"d 'seluec€rdr"firxE pu" 'sJa,,l\ou ,ft"lrx€ fElrlos sepnlcur puE elqeu"^,{HArq sr 3loq^\ " sB eJuallF snqluD!8Dldeql uqlr.r 'ra,\t,roH eJnFnJls 3cuacsorouur aea^pw aqul eql Jo EJeueBreqlo ur punoJ scrlsrJelc?Jeqc ecuocsaJogurur uoueu"d Jo eBuEJepr,r\ eql ,{eldsp }ou seopJleslr a)ua,lttq sacuatsatog[u1

olxuat$tl Io uo$l o).d'ropue'I S N NuytsiaVol. 5, No. 2 (1984)

The lack of auricles at the base of the petals in LQwrencia is a feature common to all members of the Plagianthus alliance, even in those genera with distinctly clawed petals, namely Hoheria and Plagianthus.

A greater range of colours is found in the corollas ol la\a'rcncia than in other genera of thJ Plagianthus alliance. In particular, the red- or purple-tinted corolla-sfound in several speciesof ltwrencl0 have not bcen reported in other generaof the alliance.only Plagianthus sirows much variation in the colour of its corolla which may be white, cream-coloured, yellow or green. The corollas of ,4rrerot richion, Gynatrix and Hoheria are white or cream- coloured throughout.

Several speciesof Lawrent:ia have simple-hairy or stellate-hairy corolla lobes Although this has noi previously b€en reported from any other genus in the Plagianthus alliarce' the petals of Hoheriq angustiJblia Raoul are densely stellate-haily, like those of certain sp""i-"n, ol L. tlensiflora. Many genera of the tribe Malveae have hairy petals, pafticularly in the region of the basal claw and auribles where they intermcsh serving to protect the delicatenictariferous sepalinepapillae below (Bates 1969,Hill 1982a).Protective hairs similar to these are found on the petals of male flowers ol L. be hqe, the only speciesin the genus possessingsepaline nectaries (see above). Such hairs are entirely lacking rn Gynatrix, but in iemale llowers their function seemsto have been assumedby the carpocrater which, although not previously reported, is lleshy and extelded, forming a nectarilerous collar [ke th;t desc;ibed lor Asterotrichion by Melville (1966). Curiously' although Hoheria apparently lacks nectaries of any kind, some speciesol this genus have petals which are sGple-hairy in the region of the basal claw, also reported for the first time here'

Apical insertion of the corolla lobes on the ovary cup appeals to be characteristic o1 all ginera in the p/agiqnthus allitL.'ce.Also constant in th e Plagianthus allance is the tendency for female flowers of dioecious speciesto be carinate and erect, forming a campanulate rather tharr rotate corolla. This tendency has been reported in Asterotrichion, Plagianthlts and Gynatrix(Curtis 1975,Moore & Irwin 1978,Salmon l98l)

As it Lawrencia, the corollas ol Hoheriq ar\d Plagianthus are releasedliom the maturing fruit by dehiscenceol the basal sutures of the ovary cup. The remaining two genera of the Ptagianthus alliar\ce,Asterotichion and Glnatrix, are unusual in that their corollas are regularly circumcissile(Melville 1966,Curtis 1975).

Anrlroecium. contrary to Heel's observationthat rn Plagianthus sensulato only hall:stamens leave the filament tube, {orked stamensand stamenswith closely paired (but neYer actually bilocular) anthers were commonly found in several species of I'qwrencia in the present study.

Although the five-ranked symmetry of the filaments often evident in the apical lobing of the filament luibeof Ittwrencia glomerqta, the frequently simple-hairy filament tube and occasionally pink anthers of L, berthae are exceptional in lQwrencia, similar variation in on" o, mori of theseSame characters has beenencountered in other generaof the Plagianthus alliance, namely Hctheria ar'd Plagianthus.

Another number variesmore widely in Icwrencia thanin any other genusof the Plagianthus alliance. obviously there has been a marked trend towards reduction of this figure in I4wrencia. The same trend may be observed i\ the Plagianthus alliance as a whole: only Lawrenciq alrd Plagianlhus have specieswith as few as 5 anthers in each flower, the other generahave a minimum of l0 and, llke Itwrencia, a maximum of 30. ,{lrlrueJpees ur uorlcnpor spr",l ot puert eql '(9961 se}?B'g961 ellr^lel/,\tr<1961 uellv) ocu€rlle snqluDlSDldeql lo eJaua8Ip ur punoJ sr Jequnu lad-recut uorpnper spw.rol puell eqJ '(s16lsurnc '9961 e1p,r1a11)ssrntns lurx"p" pu" prx"qu Suop {1.re1n8eruedo trTdssd:ecueru aqt qcrq,l.r ur xutDutg sr srql o1 uortdecxe elos eql dluntetu le luecsrqop,{1:zlnBer.rr Jo tuecsrqepur ,{1ensn erz eauerlle snquntsqd or{} ur "Jeue8 reqlo ;o sdrecrreru eql'mJuaLt\D1 e:1ll '(9161 .{e:9 d) o8prq:ng) s:ruq oldrurspere}t"rs seq r.?.{lrutr, Jo 1rn4 oqt :(1961u€llv) '(9/61 srultuDtBDldur puE (araq eurt tsrg aql roJ payodat) ouaqoll ur srunJ '9961 ellr^lel,{) uolq)upJa$V ur luelxe Jassel.ro raleer8 E ol punoJ oslu s aDquaq u)ual&rr7 Jo 1rn{ aql uo podlesqo slrcq elEJlalsJo ernlrlsJ^ lEulJtxJ Jrll :Juet|re snqtuDtSoldaql ur erlA\oslA 'pe8ur,n ro pe8prr1ou eJee)uErIIe snquafzp eqt.1oereue8 reql6 (0681 uu€urnqcs ur ggl e:n8rg aos)ouaqoH ur pe^resqosd:ecrreu pe8ur,tr-peo:q^lJ"rxeqe eqt eltpn elrnb e.re uJua tal Jo sarcedsp;e,las ur pedJesqosd:ecueu pa8ur,r Jo po8pr:-mor8u ,{Ipre1e1eq1 'uorlruSoceJ rueueS JoJ srs€qB s" pesneq ]ouu"J pu" o"o,\IBtrtr equt sql ur e:eua8 reqto ur punoJ sr eJuecsrqappuE ,{Soloqdjour trnrJ ur flrTrqeue^ JelrlulS erntdnJ ssoJB,{q ecuecsrqepSur,rollE uor8ar les"q eql ur sseulee,^/\Jo euoz 'uorleluerueu.ro 'urel-prur e 3o luerudole.tep drecrrad go luerudole,tep l€rxeqe pu" ernlns 'ornlrlse^ 'sseulcrql pu]rxo:d eql Suop ecuecsrqepJo uorsserdns Jo ssol drecrredut uorlcnpe: 'sd.rEJueru e1ue1s;o luerudole^ep erll Jo uorsse:dnspuE ,{lrlruoJ pu" requnu djrcrreru ut uortcnpeJ:spueJt l".re^es ur petsaJru"rusr srql eee^lEI,^,trequl aqtJo I€Jrd^l dr€JozrLlJsJrsEq Jq] Jo uortecgrpour se^lo^ur ,{U€slc DDuakAUT;o ,{3oloqd:oru trn4 elqerJe^JeqleJ eql l?tl.U' 'ecuq'rlle snq,uDlSDldeqt Jo uox"t rer{}o ou JoJ papJoceJ ueaq snq Durapos,{tqt oouatuttl Jo seqcuBJqe1,{1s eq1 uo pe^JesqoSurqol IeJrde aqA '(9/6I srunJ '9961allr^lot{'1961 upllv) seqru"rq rlBAElJ 'polrun ,{1poqs :o e3{ uo s?er? orleuiSrlselelrdec .ro ale}rdec ,{1enbr1qo'luurncep ,{nqgrlsrllr,r\ sal,,tts snojqelS ,{ll€urolxosE\DuaqoH :seqJu"Jqe}EA"lJ pelun flelalduroc lsoruqe uo see.rEcrleu8rls luarJnJep ffipurpn]r8uol qtr,r sal,{lssnorq€13 ,{lI"urelxe szq snqluotSold :seqcuuJqele^"lJ-qns Jo .reeurl'ruloJrTU palrun ,{11roqs;osuorgod :eddn oq1uo seo:r crluur8rls luer.rnJep,{llpurpntr8uol qtl,!\ sal,{ts,{rreqelellels ro snorqelSdleu:e1xe seqrrtfiutg :saqcueJq elr,,,\Blr'ee{ uo se B crleru8rtsluarJncep dlpurpnll8uoy qlvr se1f,1stueJseqnd ,{lleurslxa ssq uonpt4o.talsV exue\Ie snqtuotBDldeql Jo Ereue8reqlo ur pa,r.:asqoeq .{eruspu3Jl eul"s eseqJ 'Bere 'seqcueJq crl?ru8rls eql Jo luelxe eql ur uorpnpeJ pue el,{ls eql Jo uorsnJ SursBeJcur 'seqcuerq 'o.tnlrlsel el,{ts eql Jo sseulf,rql ur uorlcnpa: Furalxa Jo ssol 01 Surpeel spue:1 luepuedepur ueeq elerl ol uees aJerJlDDua.t^LlI uI seqcueJqol,{ls peue{crql uo }uelJncep seor€rrt€ru8rls e8r?l pEq oes.r1r1,1.{1:ee oqt Jo sal^ls eqt t"rll petseSSnssEq (996J) srteg 'suorgodreddn eq1 ,{1uo:e,r.oc{eur 1r:o ateldruoceq,{eru 1r :serre,rseqcueJq eql uo eele crl€urgrts}uoJrncep 'oee^py{ ,{lesJoJlureql Jo tuaxe eql equt eq1 Jo Erou38 rsq}o ur sV peqol ,{lecrde uelyo 'sepeds 'pue 'peue{crqt euo ur pauep"oJq dlpeprcep er€ sercadslBro^os Jo seqcuerq eytr1s 'sodft 'l€rauog eql lnq osoql Jo lsru eql ot urroJuoJ DDuatMoT Jo seru8qs aqt ul sruroJ etelrdeJ ,{llErluasseJo dteue^ € sopnlcur od,{l prrqt eqf qcueJq e1,{1spesserdruoc ,{1p:a1e1 'pouolcrql 'Surddec "'Jo eBpeJarllre uo lueJJnceppue 1eq1s ad,{1puoJos orll qtuerq ey,{1s uuo]rl$ € uo tuernJep ,{1:s:o:1urpue ele8uole sr eseqtJo lsru eql sed,{teur8rls crs€qoerql 'JelncrlJed peUrtusprseq (6961) saleg uI EJaueAJo Surdnoj8 pu€ uort"tllurlep eql ur serl 'srseqdtua scrlsuelJErEqJel,{ls Jo 3nl"A cltuouoxBJlseluer8 eqt leql Sureer8esseleqleuou srql '8961) 'fltueceu 'slo^rl Jo luetxe erll peuorlsenbsEq (696I satEg crreue8 pue Fqrrl-qns l" e€e^lehi equt oqt Jo ,{urouoxrl eq1ur srseqdrueqcmu ueat3uoeq el€q secuereJJrpoldrg sa7,f15

'jopu€.I mruatA t l lo uolst1..'d SN -

260 NuYtsiaVol. 5, No 2 (1984) is recorded only in Asterotrithlon (Melville 1966, Curtis 1975) and I'qwrencia brt may be more widespreadin the alliance (seeBates 1968).

Of the speciesconsidered in this paper known to him Melville (1966)relegated those he believedto have reticulate mericarpsto the genusZ4luencla and thosewith unornamented mericarp walls to the genus Selenothamnus.Ornamentation of the pericarp probably had its origin in the venation still evident in ce ain pluriovulate members of the t be Malveae Such reticulation of the mericarp walls occurs in many seeminglyunrelated or onl!' distantly related taxa in the tribe Malveae and is generally thought to have evolved independently severaltimes. It has not been recorded for other generain the Phgianthus alliar'ce.

On the basis of fruit morphology alor'e, I'awrencia berthae rs clearly peripheral to other speciesin the genus by virtue of its 9-11 indehiscent mericarps which have a stellate-halry, coriaceous pericarp with little or no reticulation. Further support for division of the genus Lawrencia is provided by the axillary fascicles of l-5 conspicuously pedicellate unisexual flowers, the sepaline neclaries, and the stellate-hairy style basesfound only in L. berthae In the present treatment L. berthae is placed in the subgenusPanifex; the remaining species form the subgenusLawrencia.

ln the subgenuskwlercra rcticulation of the pericarp is always associatedwith indehiscent mericarps all ol which are fertile. On the other hand, pericarps without reticulation are associatedwith dehiscent mericarps in which there is a strong tendency towards reduction in number and fertility. These two syndromes provide the basis for the segregationof the seetions L@Nrcncia and Selenothamnut respectively,in the present treatment. lt should be noted that the section Selenothqmnusrecognized here is considerably broader in concept than Melville's genusSe lenot hamnus.

within the sectiol. Selenothamr?a.rtwo further syndromes involving lruit characters are apparent. Fruit in which all mericarps are well developed are associatedwith plants of suffrutescent habit which have a vestiture of stellate hairs and which are hermaphroditic or dioecious with female flowers completely lacking anthers. Fruit in which only fertile mericarps develop are associatedwith plants of a distinctly shrubby habit which have a vestiture of peltate tdchomes or stellate hairs and which are dioecious with fernale flowers often possessingstedle anthers. These two syndromes have prompted the recognition of the serresHalophyton ar'd Selenothqmnus,respectively.

See.,/.Bates (1969) argues that in the tribe Malveae the uniovulate state is derived lrom the pluriovulate condition. A11species ol Lawrent:iq have uniovulate carpelswith pendulous ovules as do all taxa in the Plagianthus alltancewith the possibleexception of P divqricqtus which rarely has 2 pendulous ovulesin eachlocule, accordingto Melville (1966).The complete lack of any trace of an endoglossum in lawrencia is a major barrier to speculation on whether its uniovulate condition has been arriyed at by loss of the upper or lower ovuies in the carpels of the presumed pluriovulate ancestor. The fact that the solitary ovule ls pendulous may indicate the latter process,but this condition might conceivably have arisen by reorientation of the basal ascendentovule following loss of the ovules above it.

Sculpturing such as that found on seeds of speciesin the series Halophyton has r.ol been recorded for other generain the Plagianthus al]i'ance.

B. Sexuality "a Melville (1966) noted in the Plagianthus alliance strong tendency towards dioecy which has nowhere reached the final stage of complete supression ol one sex." In the present sludy of Lawrencla, hermaphroditism, polygamodioecy and complete dioecy were all '(I86I qceeg4 e,reg) ps;edsrppuu uorlcnpord uo1lod pacnper uo.rJ pel?s socJnoseJol enp sseulu lleql es€o:cut sl€nplllpul ueq,r peqsitqetseeuocaq,{eru seleuraJ'flJelMIS uollelndod crllpojqdEuueq fllEulul u" uI psrlsrTqelsaeq plno,r,\selElu 'ecuenbesuoc e sy selndo3IA ueql uellod er.rseueS e:out ltrusut:1 ,{qereql pu" ,{ep aql ur JelF"e poIsln eq ro 'ue[od eroul es:edstp,{qoreqt pu" s]ol"ulllod eJour 1rer]te Jeqtr3 .{eur sdeldsp prog re3ftl qlt,t stu€ld esneceqsseutu epru uI 3s?elcur eleuorlrodo:dsrp z o1 peel ,{eur sluz1daltpo:qdeu:aq uo Jequnu Je.,t\ouuI ss?5rcuruV (0861 'E,r"g) 'soJrnosoJ Jaquinu:anog ur sa8ueqc ,{1:elncrged I€loU uI se8ueqcol dlPclleruelp 'se3q puodsej sercadssnorceorp Jo s:o1euqlodlueururopejd aql ele l€ql sltesul leqlo pu" seIU lsqe:aue81eus leql luecgru8rsst 1r's1uo8lrSuIteuITIodeI^ ele:edo lsnu uotlceleslEnxes sculS 'snuuDqrcuaps snue8qnseqt Jo crlsuelcereqcdlrl1reg pue reqrunu dJecrreurur uorlcnpu spr€,rol ,{ruepuel Suorls eql [q pue u)uat tn jo sarcads1r ur punol sd:ecr:eu etelnlorun eqt,{q pelecrpur$ slrn{ pu" spees'so,{JqueSurdola,rep rol elqelrel" sscJnossJIauorlulnu pu€ ,{8raue Jo luour^oldap leorruouooooql roJ paeu e qcns eJnl"u ,{1uur:ou plnoc 1r u"ql lrruJ erou qllr 1ue1deqt SupeoFe^o ur llnsar ro pe^rl-uoqs ool eq ,{Erx lr ro'uorlcnpoJd s.reqlo qJee lrrurTecuorl pu? secrnoserIEuorlulnu 'ssaccns pu€ ,{8reua erues eql ezrTrlnselnlo pus uellod 1"q1 osuesaql ur p;u:e1eruecrpnferd ,(eru sre,troglBnxssrq ur uollod eqt 3ur,{1ddng eurt Suol e ;e,lo sroleurlyodyo sraqurnu aql urctuuru pu? ,{lr^rlce eql ecu"que 01 .repJo ur uorl"zqrual JoJ pepoau leql Jo ssecxe ur uelod acnpord ol stuBld ureuec .ro; snoa8elue,tpeeq ,{"ul 1I slcasur8uu€urllod ,{u?ur Jo ,{ruouore oq} ot l€tr,r sr ue11odsV srolrsrl peJaJJop.te,ner,41uo eq1 eq ot ruaasppo,r uallod os 'r€lJou 'e"eJ"^lBlr'{ ou atnpotd onuatuu| lo sercadseuo lnq lle Jo sro,\\oueql lsoru Jo esoql elqun ,{3o1orquorleullod:reqt o1 palurl fllelueurepun; a:e s1ue1dgo surals,{slEnxes eqf '(1861 '0861 rlcEeg? e.r"g E,rsg) acuelrodrur lsoru Jo ecJoJe^rlcelas eql se Sutsso:clnoJo slgeuoqoueuaB eq13ur1o,rur 1noq1r,tr dceorp jo uorlnlo,\o eqt urcldxe o1 papunodord ueaq e^erl slepotu e^rt"uretle lere^f,s sj€o,{ lueaeJ uI DDuaJi4t/I ur pelurT e.r" ursrtr pn.r qdrrulaq pue (l rlrqrledruoc-jlesluatxe leq.r ol u,,nou11ef, tou sl lI e3?1srt€l slql 1€ uorlEurTlod-JlesJo pooqrle{rTaql SursEejJur,{ltua:eddr 'snoJpu"loJd snrll rolsnlc roqtu€ eqt olur xeueJ seqcuerq el,{ls Jreqt llr,r sre./r\oueqt se tre a)uah^rl go serceds crlrpo:qduurreq l1tl ur sro,t\ou pnxesrq q8noqlyy e8eluea.pe trreuorlnyo,reu" 3q ol poroprsuoJ ,{llereue8 sr qcrq,,$ 'uoDelndod oql ur ,{lrso3,{zor3leq elqeraprsuocJo ecu"uolurEru eql sernsue ,{ceorp onrl }Brl} p3runssEuoeq Suol s"rJ lI 'pelrosqo aro,r ,{caorporpuerou ,{caolpou,{3:eqlreu 1nq 'sJeqtu" oluets ssossoduego s1ue1deprue;;o sJa,\\oupue runrcaouf8 pr3r1se,lz ssassoduago stueld elelu Jo sJ?tdoIJ'D'.:u uDT u1 ,{caorpo:puu"r^ peqsrlqetseeq osp plnoJ ,{ceorpteql ,{lqrqrssodpcrleroeqt oq} Jo s^\oll€ oslEoq.r (0861)e,a\€g ,{q paluournrop uoaq suq .{1,{1sore1eq ',{ceorpouf8 :o ,{ceouoru en :o rusrlrporqdeurJeqluo4 f,Ilce;rp fceorp Jo uorlnlo^a '(9961el|^l3t\l '1961u"llv) snorceouorueq sourrloruos,{ptll 'uortrppe ur 'qcrq,r snqtuotSolJ;o sercedsqloq ur peprooar uo3q oslp s€q '9961 orDrautolS7 ur pe^resqo,{ceorporuvS.{lod oql (6/6 I 11ex{:g sa}Eg)serlruuJ" ureueJun Jo 'qxod snue8e :eurl aqt Dtpix pug 1DaodDN ur punoJosl? sr tr :(8/61 ur,rrl ? erool^{ r9l6J '9961 '1961 'uonpllo.talsv srlrnJ ellr^le]Al u"llv) Jlaslr snqtuotSold pue xutpuig ul acuerlle snqlunSol4 eql ur punoJ sr DDua.!&Irl Jo ]Bql ol ,(qdrourp TenxesIEJoU relnurs eae^l"l^Jequl ?q] ur oJoq,resl3(1861 tsrnbuoJJ) e"eJe^l€l{ eql ur tuanb?{ur sr dceorc 'stu"ld ol"uleJJo sre,rou ur sJeqtu" aqt;o uorsse.rdnselelduroc :o lerped pue .{rluets fq peluerueldruocs1ueld epu 1o s.rf,,ro0 ur runrceou,{8aql Jo uorsseJdnsIElol ro IErtJ"d dq ,{lleJr8oloqdiour posse:dxe st puerl srqf ,{ceorpeleldruoc o1 rusrlrporqderureqruorl Surpeelpuert E SurTeeAaJ'pelolunocue

lropue-I I9Z ,rrraJr{r7 Jo uorsr^od s N --

262 Nuytsiavol. 5,No. 2 (1984)

In thesecomplementary processesrepresenting a responseto pollinator-mediated selection of the kind hypothesizedabove lies a possibleexplanation for the evolution of dioecy in Lawrenciauhich mayrepal furtherinrestigation.

C. Dispersal

The singlespecies in the subgenusPanlbx, I-awrencioberthae, probably exploits hydrochory or epizoochory. Its persistent mericarps are densely stellate-hairy on their apical and distai surlaces. Such hairs may aid flotation by neans of air-bubbles adhering to the seed coat in the manner described for speciesof Salicorniq L. by Waisel (1972), or they may serve to alfix the disseminuleto an animal's coat.

The mericarps of all speciesin the subgenuslawrenciq sectior Lavrren,ia ha\'e persistent indehiscentpericarps which become light, dry and stiff at matudty. Their apical- and lateral- distal margins are often projected to form narrow wings, and there is a substantial air- space between the single seedcontained in each mericarp and the pericarp These bladder- like fruit would seem to be well adapted for both hydrochory and for amenochory (see Pijl 1982). The areolae which develop in the retioulate walls of speciesin this section may further aid hydrochory by trapping air-bubbles.

Species in sectiot Selenothamnus appeat to have no special aids to dispersal. On the aontrary, the early disintegration of the pericarp and the releaseof naked seedswould seem to favour rapid germination at the parent site. Pr€sumably, further dispersal occurs when seed is releasedby wind-action on th€ plant or tumbling of broken branches The seeds ol these speciesare small and triquetrous, or swollen and irregularly globular, and may be smooth, angular or rugose. The significanceof the shallow pits or depressionsobserved in several speciesin this section is obscure. Ridley (1930) suggeststhat such ridges and other emanationsfound on the seedsof aquatic speciesrnay act as anchoring devicessewing to fix them to their substrate, thus preventing them from being carried away by water currents. If such seedsdo indeed readily adherc to mud then this may also aid their transport on the feet of waterfowl and grazing mammals (see Stebbins 1974) Conversely, such sculpturing may assistthe flotation ol seedsby means of trapped air bubbles thus favouring hydrochoryrather than epizoochory(see Pi1l l9lt2).

Endozoochory is unlikely in Ldrtrenciq. The seedsare not particularly hard and would easily be destroyed by a mammal's teeth or a bird's gizzard. The persistent pericarp of speciesin the seclion Ldwrencia would easily be broken in like fashion.

D. Phytogeography In general,those speciesol Lawrencia w tth mericarps favouring epizoochory, amenoahory or hydrochory are of wide distdbution; those lacking specializedaids to dispersal, are of considerablyless widespread osculrence. There are two notable exceptions to this. Despite its bladderlike fruit, L. buchananelxsdis restricted to a landlocked and geographically isolated saltlake system in north eastern Queensland. Its taxonomic alfinities would seem closestto I. \,iridi-grisea, a speciesextending from the western coast of Western Australia across the Great Sandy Desert into the Northem Territory. These two may well represent a vicarious speciespair which have diverged morphlogically through long separation by unfavourable edaphic and climatic conditions in the intervening regron. L. squsmqtq, 'Nhrch releaseslarge naked, smooth seeds,is widely distdbuted acrosssouthem mainland Australia.

Green (1964) mentions that several hundred species of flowering plants have marked disjunctions between south-western and south-easternAustralia; Beard (1969) records 280 such species.Itwrencia berthae is clearly a good example of a disjunctly distributed species pu? uorlecursJe^Ip f,&puoces ,{qeJeq,{\luslu?qceru e papt'lord s€q qcq',r^ Jo uoqc"JluoJ puelut eql ezluoloc o1 pos:edstpexel iun uoirunit" tuenoesqnseqt ser{€l-tps ;o -su€:eru icuaqauuor; sleltqeq pu"rls uI po!]lsJellppu" par"edde lsJtJmluakwl lr.qt slsoqlodfq 3qJ '1sed :qt ut Exellrqlo Jo sulJlteduollelpel IJJUJI pecu?^p" ,("lu seldru?xeoseql pesredslpf,1opln pue elqeu"^,{lq8q oq? sI'scl}suepBnqc 'opruonbs 'pu€I{ 3ql .{ueuj qlrr uoxet E 7 J3qlo eql uo-Das!8-!p!'t!^ ? Jo snrl $ otu"s plrs uollElp"l a Ilfi al"crpul 01 luelxo Jessele oI luoluuol\ua pu? aql u uotleterSos ppo,r^tr1roia3o:e1eqalqereprsuoc sll euoz pye oqt lnoq3norql pelnqulstp '{lept"rrst ureos 'olotatuo18 ,snue8oql urqI.,,\ sJepEJBqJe,rtltulud fllueuluope.rd q1r,nserceds e oQual"'It/I 'ool 'Dpuat(Atry 'lxetuoc slql ul lu",Le1ereq o1 sreedde '{ur1 " qcns electpull osl€ {eru eeece11f,qdof,re3oql ls8uorue Bx"l Jo uolleleplsuoc e xaldlltv puz (oeacelpodoueqJ) teqi'stse3-3nsp,rn iuoqerlrn11,alqrssod s" (o?acelpodouaqJ) poddns a"elulocrlss eql sellt aq :uorlceuuoc lEJollll e^Il€lnd srql 3o uI ecuepl^a (6961) '?II"rlsnV pue ,{qd€r3o.8o qd eq1uo sotpnls Jo ',lreller slq uI lu€cs spul1 uqo:e3 Jo 'slelrqeq '19461 priegj --ueldecce pea:dsepvvtqlltr 1aru s"q'.{llu.car Illun'qclq^\ ',vlell " pu"i1t ,t, p"gn.trntp pue penadde lslg qclq,,trsuorlzlndod ruor; elprlsnv Jo euoz pu€ sql Jo 'ellq,t '1861 '1861 Lop-guopJinpuirxa pirsaSSns(9961) a8prq:ng (z8ot elpeeg ',i'olr"g) u€ql JeqlEJ uorlerper ,{:euognlor\eleuJelur Jo sue3rudq erog ctlsuelcereqcu'trio s1t Sutcnpord f,q ,uoirpuo" "1q"*oie.l ripun Sur,Tro:3suollelndod luec"[p€-Iuor.;pelceles sedf,lotq snoJeurnu fllnJsseccnspazuoloc usoq s"q euoz plle eql l?ql lueureeJS"leleue8 eq 01't\ou surees0l0qJ

'wets,{seuo ,{u" ulqll^\ suollalndod Jo eloJeJeqlpue salsl FnpI^IpuI '1361 Sursnsc '(2861 rel.tog) slee,{ gg9'997 }s€l 3ql ra^o luapl^e suollel[cso Jo uorl€lo$ 'saurrl puarorordpuJ prdel orlt ol esuodseJuI sd"qJed luecer altnb uI poAIoAe3A€q ,,8ororotru 'DlJua 'su.ra1leduotlnqtllstp luasa:d rraql ql1,r,r:eqrSol u/I Jo selJeds "*o'r rfqr 1sr33is '296I snor:el eql uee,rlaq serluelurs pctSoloqdrour esolcfre,r oqJ'(296I u"udeqJ '{"uellog popualxe ,gfOt preeg) ..rn",{'1.^ djFuolrdocxa ul{Iuo ueql 8ul{ull rele,,r\q1t'tr spoued -,o]"p"inrori.q truur ruo1s,{sB urqll.t sa{el .{epol uaag 11961uouS) 1a't se surolsf,serpl 01 fwo sI u"""^t"q tnq suolsr^lp uJeA\lJq f,Iuo lou suotlulndod Jo uollglo$ eql a8esnue 1I uo€er srql uy 'aJu ttta[ ur uorletcedsro3: serlrunpoddo peqsturn; e^€q eII"JlsnV uJ3lseA Jo €x"1 suets,(s e{sl eql t€qt sr?eddB1l aJeq,\\rolrelul eql u snutuDllloualas uolpes eq1Jo Jo :.relncrged sll ,ronnr"lrloid snollqo ue uaaq sBq eJeql uI '{"po1-'{lISr3 Ip Jo erluec Joletu e'teq seq snu"8 oql ?roq,t\ ell?llsny urolsal6 ulqllr.. uollnlole pdtcur:d rteql euoSropun ureessoop pue ur pele8ir8es "t"N otrr"r^r1 Jo ex€l cuaus?"{ul snousl eq} lsql dler'lll 1I ;ur8uo jo serluec ,{;rluepr o1 suollnqlJlslp .{tp-lueserd Sutsn ut reSuep sr o:oqt q8noqlly ',(lt)tllJJoul sl JAll€ulallBJql'A\31\ '?lqels elE srql Suruoddns oJuepl^a acuasqeaql ul lnq '{lq8lq selcedseql pu? luelcu€ Jo 'esJnoc 'sI 'uoI33J suo4cunkrp esoqt l"ql elq?Alecuoc Jo U slql uI spllq"q olqEllns Jo {c"l pu" ur"ld loq!€llnN eql eql lceual ,{leJeurdEru lq8tg ueqerlsny l€eJD 3ql Jo lseoc 3ql .7 .tale,rd alerslol wntl oioclds pua aDquaq ? Jo ecuesqeeql snqJ €3s ur uorsJeruurr IIe,{\ peldepe d"ur speesrlaql suolllpuoc eulps luenber; sanadsonuatutl 1z ecurg ps:adsp :o; qloq 'eloqe_ pessnc-s1psy uoper8tu ueql leql€l 11e,,r,(puueddr sdrect:aru e,teq setceds :reql Sulendas relr'r€q ecu?lslp e^lsserdrul iir.r"alip ""unlrtp-auoi o1 slurod suorlelndod culuoclq o',rl1aseql eqr :suoucunfsp llot{1 Jo luarue,\elqceluecer ,{:ea " salEJIpuIselJeds Jo e qcea dq u.uoqi a"rr"-3:"ntp {ftuollnlole Jo {c"l oqJ 0961) Ilox'fuC dq pegoddns 'tlat'l 'lesredsip slulod e11 sa8ueqc ecuelsrp-3uolJo lnsal eql eq osp .{zrusuollcunlsrp qcns leql lno ',tleiqeurarp 'ro 'fts "rr""o1ti"14 eley iq ut€ld roqr"llnN eql Jo uollepunuul ouecoll\ aql '{le{ITun eJ" ',(q €rl?llsnv ulels3,t.\puB u.Ielwo Jo uotleredes el3urs-e;o ]lnser eql eq ol clp'rods roi,,jirnfrtp lunld qcns rltql senSr?,t"arg ,ute,' fqpeddorp peesuro'IJ ecuerrncto go ,ilqrssbii 'nueatool uro{ uerulcedseJnlelulul a18utse ulor; lmdz'olnrds 7 sI ool oS 'rrpuel Drrua./,lrr7Jo uolsr^ru S N 7-

Nultsia Vol. 5, No. 2 (1984)

colonization of the arid zone has occurred deservesfu her exploration. It seems likely that cytological evidenceand cladistic analysismight prove most useful in such a study

E. Specimensfrom Giles' Fifth Expedition willis (1981)notes that specimenscollected by w. ErnestP Gilesduring his Fifth Expedition of 1876are presumedto have been lost. Thus the discovery of Giles'specimensof Lawrencia cinerea preiuned to have been gathered in the Little Sandy Desert and of Z glomerata from the upper Ashburton RiYer and Mt Murchison is significant: ironically, thesespecimens were amongst"from those accidentally destroyed (s€e note p. 201.).It is possiblethat further botarical collections Giles'last long traverse are to b-efound at The Nationa.l Herbarium of Victoria (MEL). Acknowledgements It is a pleasureto record my appreciation of the painstaking assistancegiven in the cou$e of this study by technical staff at the Western Australian Herbarium. I also wish to thank Mr R. J. Cranfield, Dr A. V Milewski, Mr K. Newbey, Dr R. F. Parsons and Mr P de Rebeira for their collections and fie1dobservations. I am especiallygrateful to Univ.Prof. Dr K. H. Rechinger lor providing the Latin diagnoses for this paper during his recent visit to Australia, to Ms Sandra Bird for the illustrations, and to Prof. D. M. Bates and Mr P G. Wllson for their constructive criticism ol the text. I am indebted to the Curators or Directors of the following herbaria for the loan of specimensorfor making facilities available to me whilst visiting their institutions: AD, BM, BRI, CANB, CBG, CHR, HO, K, KP, MEL, NSW NT, UWA, WEL.

References

The abbrevialions uscd below for tirles ofjournals are those Siven by Lawrence et al. (1976) or ale formed according to the principles codified therein. "Flora Allan, H. H. (1961.). of N€w Zealand. Volume 1." (Gov€rnment Printer: wellington.) Anonymous(1909). Report of Meeting,January 21s1, 1909. Proc. Linn. Soc. London 1908-9:9. Anonymo us ( I 972). U ntitled M ap ( P astoral Districts of Queensland).Contrib. Queensland H erb- I 0: rear endpaper' Anonymous (1977). Regions of Souih Australia adopted by the State Herbarium Adelaide. J- Adelaide Bot Gard. l: rear endpaper. "Australian Baines.J. A. (1981). Plant Cenera."(Soc.Gro\v. Austral. Pl.: Sydney.) Baker,E. G. (1892).Synopsis of genefaand spcciesof Malvaceae.J. Bot. 30:7l 78. "Lawencia." Baker, E. G. (unpublish€d). (Btitish Museum of Natural History: London.) A photocopy of this manuscripi is held at PERTH. "Flora Barker. W R. (1981). k\rlencio, Se[enothamnus.ln Jessop,J. of Central Australia." (Reed: Sydncy.) "Flora B. A. The Australian llora: its origin and evolution. In George, A S. (ed.) of Australia, Barlow. {1981}. '(Australian vol. I Inlroduciron. CovernmentPublishing Service: Canbeffa ) Bares, D. M. (1967). Kealnemulvastum, a rtevtgenus of Malvaceae. Brittonia 19: 221 236 Bates, D. M. (1968). Ceneric relationships in the Malvaceae,Tribe Malvea€. Gentes Herb. l0 (2): I 17-135 Bates, f). M. (1969). Systematics of the South African genus, Anisodontea Presl (Malvaceae). Ccntes Herb. l0 (3):315-183. Bates, D. M. & Blanchard, O. J., (19?0). Chomosome numb€rs in the Malvales lI New or otherwise noteworthy iounts relevant to classificationin the Malvaceae.tribe Malveae. Amer. J. Bot.51:927-934. Ba\ra, K. S. (1980).Evolution of dioecyin floweringplants. Ecol. Syst. l1: l5-39. Bawa, K. S. & Beach, J. H. (1981). Evolution of sexual system in flowering plants. Ann. Missouri. Bot. Gard. 68t 254-214. "The Beadle, N. C. W. (1981). Vegetation of Australia." (Cambridge Universily Press:Cambridge.) Beard, J. S. (1969). Endemism in the Western Australian llora at the specieslevel. J. Roy. Soc. Western Australia 52: l8-20. Beard, J. S. (1970). The penetration ofrhe western deserts of Australia. Geograph. J. (London) 136 (4): 549- 568.

-, r(6c8) '(c€r^t€y{ 60t- € 'lllH t8 lroporitJ l :aBar€^tet{)[e4) V un4sD^tDt[snuo8 .q] Jo qd€rSououV (rz86l) U S 't9z-69t:(8t8) '(aee^tpt,{ t8 BropoqdII :leote^pt ) ter|, v untsq^tDn snuo8aqt lo qd€rSouotuV (qz86l) ):t S luH 't8-t :u.€8) '(rBe^tEr,{ 'lllH t8 €ropoqd I :o?o.?^l€II) t"J|. v atuFD^txn snua8eql Jo qdtl3ouolu V 1ez86l) d S '(996t)'ue^ n6t LLI :Qt tI Batunlg sets^lehluI tunlcoorpu?aqlJo ,{Soloqdrot^{ V ,,!\ leoH :.cUIO.{reuoll€ls s.,{No[ew raH)., ,{et 'suep.I?Crlu€log 'unueqr.H .qI l3 Cuopuo'I 'uosuDIM ',{rorsIB,{oU '( 'oplel[ 'a ',{ep{pH palduroc suorleherqqvroqtnv Jo xepul U"rC,, 10861) H T' f U :rt ErrErisnv -cos zt-sz 'uaa:C urotsiy'd ,{od I Erleilsnv uroqtnosul sarcadssnorE.r^ pournsordpue snonulluo.sr(l (}9et) ,rl t 'uotsr€ '^{o't '(6881) (uopuo't:uol;u rU ? alrpas I,\l uo sdu€S) .. Z lo^ pesra,tz:1aot,l\l erte jlsny., A A,\l'sollo (-uortEls ' a8e oJ :ssard,{rrsra^run N T V s?x:I) .. eqr{ uorroJ lqr lo tuolsrH I€rn}€N aql, (646t) V A ex,ftC '0SZ-61.t -(eeac€^t€Ir'{) :9t prEC log unossrt\ uuv ^EJ otsoBanlua1snu!3 rqt (696t) V U ll.xIrJ nT-gtt :gI uoxpl suoqnqulslpl.unfslp Jo uoqtrardrarulaqf (f96t)'V a 1laxfiC 'tl-01 :l s^\oNqj.H s€lelsnv 'uort.ollo.uorllp.dxE replA aqrpuE surlrH uaqox suonto[oc1u€ld ueII€4snV IIrEe 3uos (0e61)t,{ J'Iolprea ('oSpuque] rss d {trsr.^un a8prquleJ) .. uorreta8:Aunrrrjlsnv., H )J\o^orDut ruop8uDtrllslrou utrtBrlsnv rqrJo trqdEl8ooSot.{qd 1t86l) H'8uIoC 'y{ 'srunJ ( u€qoH :retuudrusulurs^oC) .. 1u?d ?rur?u-Is{Jo €rolJ s.luapnlseqJ,, (9/61) /A ()iro,{oN :ss3rd,{rrsr!^ru{lerqlunto:)).. slul?td Surrar\\olC Jo uop€rUrsselJJo ua}sfs pol€dalul uV' 11861) V lsrnbuorJ L9Z-10<- (') 96 setel[ qlnos ^{3N 3oS ur.l cord sru€ldtrro}Irr3l uraquoN Jo lsll)t]aqJ 1ir6l)-t{ C el€purddrqJ ( qlid :uoD€zru?8roqrnased ieulsnpul PuErgrluons qlPar{uoruruo)) 8l rodad IPJIUqral,{o^rnS q"r""r"U puljt..iqeirrsny u.iotso,lieun|trl pue uelD Buro-I le {o^rns ISqorpIH,,-(2961) C I'u?urd€q) (e l^r{\0rc:suorlecrlqnd

'lopue-] ,lrrrrr{t7 Jo uorsl^o){ S N 266 Nuytsia Vol. 5, No. 2 (1984)

Hochreutiner. B. P G. (1920). Organes carpiques nouveaux ou meconnus chez les Malvacces Ann Conscrv' Jard. Bot. Gencve2l: 347-387. "lndex Holmsren. P K.. Keuken, W & Schofield, E. (1981) Herbariorum Part I The Herbaria of the World-" Se\enth Ldition. RegnumVe8. 106{ Bon. Scheltema& Hol[ema: t trechl) "Flora Hooker, J. D. (1855). Tasmaniae.Vol. L" (Reeve:London ) "The " Huichinson. J. (1967). Genera of Flowering Planls. Dicotyledons Vol. II (Clarendon Pfess:Oxford.) Herbarium Lander, N. S. (inpress). Collectiorisof lawrenciaHook. destroyedby fire, April 1984.western Austlalian Res. Notes. "Botanico-Periodicum-Huntianum " Lawrence. H. M. et al. (1968). (Hunt Botanical Library: Pittsburgh ) Leenhouts, P W ( 1968).A guide to the praclicc of herbarium taxonomy Reg Veg 58r l-60 "Rare Lcigh, J. er al. (1981). or ThrearenedAustralian Plants." Special Publication 7. (AusrralianNarional Parks & Mldlife Service: Clanberra.) "Diclionnaire l-eme6. A. (1934). Descriptif et Synonmique des Genres de Plantes Phanerogames Tom V: PAI-- SCL" (Brest: Pads.) Maiden.J. H. (1906).Records of WesternAustralian bolanists W Austral.Nat Hist Soc 3: 5-33 ..Poorly Marchant, N. G. & Keighery, G. J. (1979). Collected and Presumably Rare vascular Plants of western Australia" King's Park ResearchNotes 5. Melvile. R. { lq66). Conlriburions to ihe Ilom of Australia: Vll generic delimitalion in the P/agianlir.t complcx. Kew Bull. 20r511-516. "Flowering " Mitchell. A. (1983). Malvaceae. ln Morley, B. D. & Toelken. H R (edt Plants in Australia (Rigbyr Adelaide.) "The Moorc, L. B. & Irwin, J. B. (1978). Oxford Book of New Zealand Plants." (Oxford University Press: Wellington.) (t) Mulcahy, M. J. & Be(ena], E. (19?2). Soil and landscaPestudies in Western Australia The major drainage divi'ionr J. Ceol. Soc. AustraliaI8 (4):l4q-357 Mulcahy, M. J. (1973). Landform and soils of soulhwestern Australia J. Roy Soc westerD AustraLia 56: 16- 22. Mueller.F. (187J).Malvac€ae. Fragm. Phytogr. Austral.9: 126 13l. Pickard. J. & Dluqai. H. {1981). Map: Botanical Divisions and Subdivisions of New South Wales. In Jacobs, 'Plants S. W L. & Pi;k;rd, J. ot New South Wales." (Gov€rnment Printer Sydney) "Principles " PiJl, L. van dcr (1982). of Dispersal in Higher Plants (Springer-Verlag: Berlin.) "The " Ridlev, H. N. (1930). Dispersal of Piants Throughout the world (Reeve: Ashibrd ) Weslern Australia: Census, Atlas Rve.' B. L. et al. tl980r- Commerciallv Exploited Vascular Plants Native in and Pretrminarv Assessmcntof Conservation Slatus." Report 40. (Department of Fisheries& W-rldlifc,western ALrslralia: Perth.) "Thc Salmon. J. T. (1980). Native Treesof Ne\l Zealand." (Reed: Wellington ) "Western Sharr. F A. (1978). Australian Plant Names and Their Meanings." (University of Western Ausiralra Press:Pe|ih.) "Die schumann. K. (1980). Malvaceae. In Engler, Pranrl A. & K. (edt Naturlichen Pllanzenfamilien. Teil 3. Abteilung 6." (Englemann: Leipzig.) Short, p S. (1981). Pollen-o\ule ratios, breeding systemsand distriburion patterns oI some Australian Gnaphaliinae (Compositae: Inul€ae). Muelleria 4 (4): 395-418 "Evolution Smith White S. (1982).Summary and redintesrarionln Barker,W R & Greenslad€.P J. M (edt ollhe F'oraand Iduna ol Arid A.r.lralra (PeacockPublicalions: I-rew\rllc ) Sprague, T A. & Summerhayes, V S. (1926). The taxonomic poslt\on of Hoheria lyallii Kew B\tll 1926:214- 220. "Taxonomic " Stafleu,F. A. & Cowan,R. S. (19?6). Literature Volumel: A-G (Bohn, Scheltema& Holkema: Utrecht.) " Stebbins,G. L. (1974)."Flowering Planls. Evolution Above the SpeciesLevel (BelknapPress: Cambridge.) Stones,M. & Curtis,w (1973)."The EndemicFlora ofTasmania.vol 4."(The Ariel Press:London.) Waisel,Y (1972)."Biology of Halophltes."(Academic Press: New York.) W-rllis,J. H. (1949).Botanical pioneers in Victoria L Vict. Naturalht66 (5):83-89 "Flora Mllis. J. H. (19S1).The history of bolanicalinvestigations in Cenlral Australia ln Jessop,J. (ed.) oI CentralAuslralia." (Reed: Sydney.) 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L9Z ,rr&rl1r-I}o uo6r^el{ ropue.I s N Nultsia Vol. 5, No. 2 (1984)

s.n.,Nov. 1924(2-Al)), s.n...lan. 1926 (2-AD). s.n..5 Nov. I926(10'AD 96601963), s.n.,5 Nov. 1926(10 AD 96805867).s.n..7 Nov. 1926(10 AD), s.n.,I Nov. 1928(10-AD). s.n., 27 Nov. 1928(1o-AD). s.n.. .lune I933(3-NT). s.n.. Aug. 1939(3-AD), s.n., Nov. l94l (3 AD), s.n.,July I94l (l-AI)). s.n.,Nov. 1950(10-Ai)), s.n.,Dec. 1953(3-AD). s.n., Dec. 1953(l0AD), s.r., Nov. 1955(3 AD) s.n.,Nov. I956(l Al)), s.n.,Nov. 1959(2AD), s.n.,July 1960(3-PERTH), s.n.. Feb. 196,{(3-AD). s.n.. Feb. 1968(10-AD), s.n., Nov. 1968(3 AD), s.n.,s.dat. (3 NT). alle,re'?r,f' s.n.,s.dat. (4-K). Coleb.Williamx, .,i. ll7 (ll-PERTH). 118(ll-PERTH). Combet,H.F 2024(.2 KJ. Caoper, FI.M s.n.-March 194|(3-AD). C.Ukt, B.81'/(3-AD). 1020 (10-AD). II36 (2-AD), r235(ro-AD). r563(3-AD). l57l (3 AD). l60l (3 AD), 2198(10 AD), 2719(r0 Ar)), r67it(3,Ar)), 3843(3-AD), 4541 (10-AD). Catnwatt. G.C. 129(3-AD). Carrtuk. M.C. 5072(3 MEL), 6219(l AD. MrL). 6376( I -MEL), 6380( I MEL), 6656( I0-M f:l.). Covenr,, -ti. 8258 ( l0-NSW PERTH). 841I (I2-PERTH).ounfttd, R../.638 (s-PERTH), 1723 (7-PERTH), i856 (7 PERTH),2521 (4 PERTH).2584 (4 PERIH), 2595(4 PERTH). 4009(I0-PERTH). Ctaven, L.A. s033(12-BRl, MEL. NT, PERTH).C/dqbld, 39 (10M[L),48 (10MEL]. Crisp,B.C. 5'76(3-CANII). (ilv, M.D. 676(]-AD, CANB), l2:ll (10-MEL).3295 (1O-CBG. MEL). 3417(3- CBG,MEL).5469(IICANB),5630(I0CBC. N'I) C/,./.ea,R.L. s.n...lul), l9l9 (3-AD),s.n.. ug. 1939(3- AD), s.n.,Aus. 1939(10-AD), s.n., I939 (l0AD), s.n.,s.dat. (10 AD). Cronin,'l1. s.n.. 1890 (l MFl,). s.n., 1893(l-MEL). Cumminss,D.J.232 (10-CBG, NT\. Cunninshain,G.,l1. 3026 (3 NSw),4776(10 NSW). Carrrs, WM. s.n.,J^n. i944 (2-Ho), s.n...Ian. 1946 (2-Ho). s.n.,Dec. 1946(2-Ho). s.n.,Apfil l9sl (2-Ho). s.n., "Chorney Feb.1966 (2 HO). CzornijGs \, K. 1066(3-n D, BH. K, I'ERIH) /)atu.&i, s.n..s.dai. (2-MEL. NSW PERTH).Dallathy, J. 217(3 MEL), 218(3 MEt.). s.n.,s.dat. (10-K). D Alto'i, St f' I6 (l MEL), 26 (3-NSW).s.n., Oct. 1899(3-MEL). Daries, C.F 283(l-PLRTH). De MakL C. s.n..Scpt. 1964 (4 PLRTH). Dennis, T. 152(2-AD). de Rebeira,2 415 (6-AD. PBIL|H), 122 (6-AD. PERTH), 433 (l2l'ERTH). D"'ral7, l/. 4445(l-KP, PERTH).46t0 (10KP PERTH),4712 (1 AD, KP, PFR-fH),4587 (3 KB PERTH),4690 (3 KP,PERTH). 4E29 (4-PERTH). 4831 (7-KB PERTH).4894(8-KB PERTH),5292 (10 KP), s167 (3 KP,PLRrH). 60r5(2-Kp),63i0 (r0-KP. PERTH),7140 (7-KP, PERTH).8241 (7-KP),8995 (7-KR PERTH).9258 (2-PERTH), 9540 (4 PERTH). De.\eslise,t,. s.n., 1866(3-BM). Dieh, t,. 5204(8-PEI{I'H). Dlr.od, .t. s.n., Oct. 1893(i2- AD). Donnet,x. N. l78s (3-AD).24s7 (10-AD), 24s8 (10 AD). Dtunnnnd, 1 6 (3 K). 24 (9 BM. K). 55 (l BM, MEL, PERTH.K), 104(9-K). 106 (10-BM. K, PERTH).137 (3,K).208 (r0 BM, K. MEL. P|.RTH),2s2 (10-BM,K, MEI-),246(3-K),2?5 (l-K),302 (2-BM, K, MEL).s.n., 1848 (10-K). s.n., Jan. l8sa (2-MEr-), s.n.,s.dar. (l MEL). s.n.,s.dat. (2 K, MEL 98729,98710 98754), s.n., s.dat. (4-MEL 98692).s.n., s.dat. (10 K), s.n.,s.dar. (10-MEL 98771. 98772. 98781, 98785). Dunlop, C 1810(l AD), 2452(10 K. NSW).2541 (l AD, MEL, Nt.). Eckefl,J.P s.n.,1892 (10-MEI-). Ei.hkt, Hi 12536(3-AD). 12958(3-AD). 13001 (10- D), 14r86(10-AD), 15732 (10 AD), r6859(2 AD), r7629(3 Ar)), 18745(3,AD), r9417(r0 Ar)), 19433(r0 AD) 19434(10-AD), i944s (l-AD). 19550(3-AD), 19550 (3 AD), 20317(9 AD, PERTH).20121 (10 AD, PERTH). 20322(10-AD). Elkingtan, .t.330 (12-PERTH).Effoy, E.G.3228(7-NT). 3314 (12-NT). beri.\!, 5.L.9164 11- BRI). Fass, M. 376(3-AD). Fairul, A.R. 1825(12-KP). Fell 1-.678 (10-KP). Fi dgan,t.3 (2-MEL). Fitzgenld, lU L s.n.,1893 (2-HO), s.n., Ocl. 1898(8NSw, PERrH), s.n.,Jan. 1903(10'PEIUH), s.n.. Aug. I903(10 NSW, PERTH), s.n., Sept. 1903(7 PERTH), s.n., Nov. 1907(10 NSw). ri.)"nriclr, ,. 8 (3 AD). r.,/.ic. N. 479(3-AD, K, MEL). 1330(I0-CANB), 1355 (I2-CANB). Forrest, A. s.i., 1879(3 & 4-MEL).Faffest, J. s.11., 7 June I874(3-MEL), s.n., 28 June I874(3-MEr.), s.n., 30 June 1874(I-MEL). s.n.,1882 (3-MFll.). l-laJ?/, C. 27 (2 BM), 126(2 BM), 118(2 K). Fren(h,C s.n.,April 1923(3-MEL). Galbraith. J. t0 (t0 A,I)) Gardn?t, a:,4.r993 (l-PERTH),2094 (1o-PERTH),2r28 (12-BM, PERTH).2403 (7-PTRTH).2441 (7 BM. K. P|RTH). 2444(I0-PERTH),2545 (?-PERTH),3264 (t-BM, PERTH),3264A(3-K,PERTH), 6075 (.r-PERTH).6089 (?- PERTH),7532(1o-PERTH),7800 (3-PERTH),7814 (12-PERTH),8618 (1o-PF.RTH), I3352 (?-PERTH). s.n., Aug. 1927(12-PERTH). s.n.. Aug. l93l (7-PERTH),s.n., Sept. I935 (l-PERTH). Casteen, J.85 {s-BRl).Gauha, I] s.n..Oct. 1942(3-NSW. CANB). 6€orse,,4.S. 915 (I2-PERTH), 923 (I2-PERTH). 952 (7-PERTH), 1182 (7-PERrH), 2290( ro-CANB).229 r ( ro-CAN B). 2948 (3-PERTH), 4211 (8-r'ERTH), s692 ( l -AD. BH, K, PErt r H), Slll (3 Ar), BH, PE|TH),9r3r (lo,AD,PERTH), 9r32 (3 MEl.*,NSW, PERTH), 9r37 (6-PFTRTH, AD), 9765(3 BH, K, PERTH),10207 (7 PERTH),10214 (4 BH, K, MEL*, NSq \l PERTH).ll380 (Nl PERTH). 14783(4-PERTH), 14812 (6-BH. NT. PERTH),14813 (6-PERTH). s.n.. Dec. 1961 (1O-PERTH). s.n...lune 1970 (9-PERTH).s.n., Sept. 1972 (4-PERTH). Georye, D.N. s.n.,.htiy 1931(i-AD). Gilet, WE.P sn, 18721(3 MEL), s.n., 1875(|'-MEI-), s.n., 1876(6-MF.1,), sn. t876 (3-MEL 98695),s.n.. 1876(3-MFL 96731).Gi//, W s.n.,Feb. i903 (2-NSW). Glennon, P 109(4-PERTH). Goodal, D.W 2910(I'-PERTH),331i (7-PERTH. UWA). Cosr,C.li s.n.,April 1955(3-AD). Grutte, S. s.n.,Aug. 1970(3-AD). Grcen, A. s.n..No\. 1924(l- I'ERTH). c/egof), "/.l/. s.n.,l90l (l-BM). G/egolr',I'rof. s.n.,May 1909(12-MF-'.). Cfle'., AJ. s.n..Ocr. 1960(10 tJwA). GlilJin,E.A.301J4 (4 PERTH).Gril|ith, H.H.D. s.n.,Dcc. 1907(2 AD), s.n.,Oci. 1927(10- AD). Gulan, Pr( 384 (2-MEI-). Gunn, R. 746 (2 K). Ha , ,1."/.s.n., Feb. 1928(2 PERTH). Ilannafor.l, S.A. 9 (10-r-SW).HeaL M. s.n.,1889 (l-MEL). Helns, R. s.n.,Sepl. l89l (l-MEL), s.n..Nov. l89l (9-AD, K. MEL), s.n..Nov. l89l (12-AD.K. MEL. NSW,PERTH), s.n., 1899 (12-K, PERrH). Hendetson,w.2556 (lt NSW),s.n., Oct. 1946(i0-NSW). Henr!, N.M. 378(4-MEt.), 383 (3-AD, N I), 953(4-AD). Henshall, T s.n.. Sept.i966 (10-NSW). Herb?rt, D.A. s.n.,July l9l8 (12-PERTH).Ilerlisus, PC.79140 (I0-CANB), 80136 (10- AD, CANB). Hirr, F.L. 119(4-BM), 124 (4-BM. K), 116 (4-CANB, K). 6t9 (4-K). Hir, R. 54 (3-AD), 229 (3-AD), 1006(]o-AD), li73 (3-AD).Aolsr, ,V. O. s.n.,April 1896(12-ME:\). Holbut.t, s.n.,Oct. l9l7 (3 PERTH). Humphties,A. rl: P48 (ll PERTH). 1nce,tad s.n.,Jun€ 1903(12 K). Isins, E.H. s.n., Dec. 1922 (2 AD). s.n.,Jan. 1925(2 AD), s.n.,Aug. 1925(8-AD), s.n.. Scpt. 1936 (l-AD), s.n.,Jan. 1937(1o-AD), s.n., Nov. 1938(10-AD 96935429 & 97416003),s.n., Sept. 1938 (3-AD), s.n., Ocl. 1950(3 AD), s.n.,Scpt. l95l (3- AD), s.n.,Aug. 1952(3-AD), s.n., Aug. 1955(3-AD), s.n., Sept. 1955 (l-AD), s.n.,Oct 1955(3 AD). s.D.. Dec.1958 (10-AD), s.n., Sept. 1960 (1 AD).JuLkson, tN.S. 741(3 AD, PERTH),Il36 (AD). 1835(10-MEL), 2559(10-AD). Jacob, G.L. s.n.,Ma), 1968(3-NSW). Jd..,bs, S.WL. lll4 (10-NSW)../orrJ, (:.W s.r., 1941 (10-AD\.Johnsan. U. 30 (1O-NSW).Keishet),, G..t. 1956 (1o-KP, PERIH),2966 (8-KP),3883(2-KP),5541 (10-PERTH).Kenneally, K.F 7351(7-PERTH). Ke,r,zy, fi. s.n., Feb. 19,{6(2 UwA). ('ngs,'ri4 W 185 (12- BRl, NSW). 1(n/€p,ir'. s.n., 1937(l2-PERIH). Koch, M. 1161(10 K. NSW). Kraehenbuehl,r). N. 495 (10- AD). 9i8 (10-AD).Kuthet. R.H.271 (3-AD),5ls (10-AD).659 (3-AD). 681 (10-,^.D) 7i6 (3-^D)- 2147(8- 'reps'us lus rO'tols ('669 T €669 A\SN'WB-t) 6l ^oN J ?]r[o,S (H.Ll{gdZ) 8/6t ^oN 'us (CV-ot) '(CV-€) ('IgI,'t'SNVJ '(]V I'ut)sua^ats'L4\sN"tEht ') 'Ius6 9'(AV-Z\,ZZt'cv '(H-Lxado€tz'(CV-ot) r/9t ty'lauoads 'sNvf-t) -€) st 0t) 9"t ,rsN'gNVl'rua-zr) ttl'(HJdEd 0t/ '(HrdEd-r) '(H-Lxsd'sNvJ 'l)ds (dv-0r) '^oN 'u's '(cv-z) '(cv-Or) 'r{)rd.S €89 ' t\'lloqtnos nn9 H N 696r /zzz rz ?y '(cv-z)(CV t)'(cv-z) 096t .0(1 u.s'(cv-Or)7 d. '(qv-Or)(OH-d 0961'(cv-0r)racr'usT riiinrr'(cv-0r) ros1CV-0t)006t'(CV Z) I/81 r98l 098r 96/r rzzr rsrr Ezzr'(cv-z) 60| '(cv-Z) '(cv-ot) '(ov-0t) 'tttlus 'qt!111s(Hl.xE.l-r) 't,ru.r,s(v,\\n-ot) 960i 'u 08s'D't 'tlttruS6€n tJ 1'raN-zt)'u fls 'sllrisI a 'OSJ 'AV-t)ts a 'ir"ryg 8S6t 8nV s (CV-OI)696t 8nV s g? (tl^ S8 Sa iOO (HDlAd-d qoCr'usj ',{tua^lds(CV-OI) reps'us'(CVZ) '0dB0) nZZt 'CV-l)IIJN'pDrtotts 'u '()-Z) Z86l''u '(HJdado ') '(]V-01) leps,.u.s reps s Z/81 tl'rtt{ s Z8 0 U 1tunqtuo?S (d)-t) 116l t,pa,,s (H}{sd-rr) yN'Ia4lDS (Hrugd-z) '(Hrdsd-€)lro'uss4.1i 9r8'(HrdEd Zr) f08'(HrJEd,t) 199 /116 ss99'(Hrdsd/) 0€99'(Hrxad-or) 6z8s'(Hr)lEd-or)L9t9 ov art:av (^[sN-0r) 9r6r rro 'u.s y frl.{ 'rratoy (ASN-Z) lep s 'u s '(8L6ZI rPps 'u s '(t{ 'OH-Z) 'u s '(OH-Z) 'uel'us'(OH-Z) 'us OH-Z) 0t6l IudV '{'uosutqoY/681 t88t 7 tr,npo)' (/dSN')-Z)SSIZ ?'rr,{poy ((IV-01)9t6t rro'us '(r-Z) rep s 'u's 't't 'uosuaqo\ q.j'us (X 'rtg-/) g1 ',{r/pru 'u '(Tsl^l-t)I]gt,t-Z) 'u 0961 '('IEr{-t) l{'uowtit'us'(9a986t960t86'r:tNt)//8t'us'( t98t tdosJrrdv'us ('rsl^,t€) l€ps s 088t s 008t 896 'IEhl ' u s '(01886'IEt^l-ot) ' u s '(-lej sptDWtA 1ppsIus ('rett 0t)9t8l "u '(-IAI,{-0I) '(t3rnl-ot)Stsl -t t-t) t J klY-1t)'u OT'rrry'JeC '! t) S68tqrret\ s 8 9 tapoaYI'Igr,,t-t) 868t ^oN s (tan-Z) 968t u s '('rar,'t-01) !o ' s '(r{s-r) qr&hr 'u s '('rEW 'CIEW-C) 'tapDaa(H-LdEd-z) 'u s68r'dutoluDuDnA 968r 'altn] Z) 8 '(CV-0r)z Nr '(CV-Or) '(CV-0r)}16r rec s ? lrxs-0r) C 691,,r1V (Cv-or) tfr zzr 02, '(ov-01) 'uaqold (.I1n-z) 'sr]a.? 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692 Dlruat 'n Jo uotst^ad'rapu€'I sN Nultsia Vol. 5, No. 2 (1984)

(2-MEL).Srrei.h, L s.r.. i895(12-MEL). Srrid, ,4. 20150(12 PERrH). Stuart,J.M. s.n.,1859 (3-K). .trror, C.S.I 174(10-K), Ii75 (3-K),s.n., Ocr. 190s(2-MEL). Swinbourne. n. 94 (3-AD),180 (3-AD). Slmon, D.E. 2338(3 AD, PERTH),3205 (r0-NSW), 325r (r0-CANB, K), 3268(3-CANB, K), 4300(i AD, CANB),4308 (3-CANB),5460 (12-AD, PERTH), IJI49 (3-K), 9945 (12'AD, PERTH). Tate, R. s.n.,1880 (3-AD), s.n., 1883 (2-AD),s.n., l88J (3 AD), s.n.,1889 (3 ADr. Telfard,LR. 5859(12'CANB). T(pper. J.G.o.82 (l-MEL), 444 (I-MEL), 756(2-MEL),880 (2-MEL), 916 (3-MEL),932 (2-MEL), 1209 (2-MEL), s.n.. oct. 1879(l-AD), s.n., Jan. 1880(1-MEL), s.n.,s.dat. (l-MEL). Tietkins,WH. s.n..I880 (3-MEL). Tilden,J.E. 797(2-BM, K).'l:indale, I,LD. 130321(3-NSW). Iirdalp, N.a. s.n.,8Aug. 1935(3-AD), s.n.,22 Aug. 1935(3-AD). Toelk?n, H.R.6093 (12-AD,BH, BM, CANB, PERTH),6094 (3-AD, BH, PER-|H),6129 (10-AD). ?iszira P 359(3 AD), 370 (.3-AD).Turner, 11. s.n.. Sept. 1960(3 AD). Tillner, J. s.n.,April 1977(2-MEL). Tyson,L IE (10-MEL), s.n., 1892(7-MEL). Vitkery,.t.W. s.n.. Oct. 1949(10-NSw). rara N.M. 9 (io-AD),45 (1o-AD),I36 (10AD), l,{6 (3-AD), 172(10-AD), 193 (3-AD), 2r2 (10-AD),2r9 (3-AD, MEL), 228(r0-AD), 255 (3,AD), 262 (10 AD), 270(10-AD), 285(3 AD). Wad?,R. s.r., Sept. I97l (10'MEL). trakefeLl N.A.2243 (2-MEL). Wahet,C. s.1., Oct. 1899(3-NSW), s.n., Oct. 1899(10-NSW), s.n., Oct. 1900(10-BRI, K). s.n..Feb. 1902(2-NSW\. Warbwton, P a'. s.n.. s.dat.(2-MEL); s.n.. s.dat.(3-MEL). Wames,K.B. 3l (3-AD), 34 (.3-AD). Webet,J.Z. 774 (3 AD, MED, r32l (3-,\D),r380 (3-AD), 208r (3-AD), 3568 (lo,Arf, BRr),4229 (10 AD), ,1251(10-AD), 4339 (10- AD), 503i (12-PERTH),5786(l-AD, BH, PERTH).Web\ter, G. L. 18666(8 NSW). Webstet,L.G. s.n.,1889 (8-Nsw). Weindor/ar,G. 222a(2 MEL), 222b(3'MEL). Weston,A. s.n., 1892(7-MEL). Wheattroft,C. 3618 (3-A.D).Whibkr, r.Lt 562(10-AD), 1234(3-AD), 3425(3-D), 5673(10-AD). Whinrc!, J. 17(2-MEl,). Whne, S.,4.s.n., Sept. l9l0 (3-AD),s.n., Sept. l9l2 (3-AD.K), s.n.,Sept. 19l2 (10-AD, K), s.n.,Aug. l91l (3-AD), s.n.,April l9l6 (2-AD),s.n., Oct. I9l6 (3-AD),s.n., May 19l7 (2 AD), s.n.,1936 (2-AD). mlcox, D.C. 31 (3 PERTH), s.n.,April 1954(l I PERTH), s.n.,Sepl. 1966(Io-PERTH). Mlcocks, M.C. 46 (10-AD). M iuns, C s.n.,s.dat. (10-AD). Wlhelm| C s.n.,Jan. i852 (10-MEL),s.n., 1854 (10-K), s.n., s.dat. (2-MEL 98719), s.n.,s.dat. (2-MEL 98738), s.n., s.dat. (10-MEL). Mllians, L. D. 4102(2-AD),6388 (3 AD), 7119(10 AD),7234 (1o-AD),7488(10-AD),8688 (10-Al)). Williatnson, H.B. l02l (2-NSW).Wi is,J.H. s.n., Aug.1947 (10-K, MEL 98782),s.n., Aug. 1947(10 MEI- 98780),s.n., Aug. 1955(.3-MEL). Wilson, H.M.22 (2-PER-IH).lnbon, PG. 602 (3-AD, K), il54 (2AD), 1s88(3-AD), 1604 (10-AD), 162l (10-AD),1762 (3-AD),2623 (ro-AD),2664 (2-AD),2793(ro-AD),5798 (10-PERTH),607s (r0-K, NSW, PERTH),6403 (10-BH, BRr, NSw, NT, PERTH), 6912(3-BH, NS\ PERTH),7i36(10-BRr, K, NSW,N',r, PERTH),7340 (7-PERTH),7393 (10-PERTH),7506 (r2-PERTH),7654 (ro-K, PERTH),7655(I0-PERTH), 8234 (4-PERTH), 8249 (10-BH, NSW, PERTH), 8604 (r I-PERTH).8635 (10-K, PERTH), 8636 (r0-K, MEL*. NSw' PERTH),8866 (r2-BH, BM, BRt, CANB, K, NSW,Nr), 8867(r2-BH, BRr, CANB, CBG, K, MEr-*, NT, PEnTH),8945 (7 PERTH),8964(7-BH, NSW, PERTH),9950 (3-BH, BRr, MEr-*,PERTH), 10163 (9-PERTH), rr582 (r2-PERTH), n583 (12-PERTH),s.n., Sept. 1966(g-PERIH), s.n.,July 1967(1O-PERTH). lntwet, E. 349 (l-KB PERTH), 97? (10-KP,PERTH), r 120(3-PERTH), 1395 (Io-PERTH), 1397 (3-KB PERTH),1429 (10-KP, PERTH), 1447 (3 PERTH),1734 (12- PERTH),1768 (12-KP), 1845 (1o-PERTH), I89l (2-KP).Wood-Jones, E s.n., Nov. 1920(l0,AD, NSW).r/is1el, J.ll 5802(2-CllG). Zinner, WJ.s.n., Oct. 1932 (l MEL).

Index to Names and Synonyms in Zawrencia Names of new taxa, new combinations, and names with new status appear in bold type, other acceptednames and epithetsappear in roman, and synonyms in italic. For an accepted name, referenceis made only to the page where the main entry begins;lor a rejected namc, only the page where it is listed as a synonym of an acceptedname is indicated.

Halo t hamnui F. M[elL, 211 micrcphyllus (F. Muell.) E Muell., 246 LawrenciaHook.,2l7 subgenusPanilex Lander, 221 subgenusLawrencia, 224 section Lawrencia, 225 section Selenothrmnus (Melvitle) Lander, 237 seriesHelophyton Lander, 238 series Selenothamnus, 2zl4 berthae (F Muell.) Melville, 221 buchanenensisLander, 236 chrysoderma Lander, 249 cinerer Lander, 238 densiflora (E.G. Baker) Melville, 239 diffusa (Benth.) Melville, 243 giomerata Hook., 229 helmsii (E Mueil. & Tate) Lander, 252 ,rca"l? (J.M. Black) Melville, 246 repens(S. Moore) Melville, 242 spicata Hook., 225 squamata Nees ex Miq-, 246 viridi-g sea Lander, 233 'f,ero ZIZ'IE^uI u.rou V elel'ue{{[ '3 9ZZ'[enVI ol2uaraol IZZ'l?^u tnou'[onI^I J aDquaq?PIS 'al[I^ley'{ ( bI}^I xe saeN)shtbtubnbs 9tZ 'eIII^lol ZSZ I (erel d' llenl,^{ I) llsulr{ ' ' 'eIJ]./'I.l{^I vrz Ltz Llz snuuoqtouaPs 9gz'llonll U snulnF^ sodlecoulclu 'qlue8 9tZ ( b4/'I xa seeN)snpwDnbs sunsaqtld&^ 6ZZ' \!'re(' SZZ qruo8ClooH) sntrrSdr ZIZ'Izzlud ! otPuDruadre^ ZrZ'erool L S suadar xa .utleH snclouoa 992'ue.i.!" glz l]ptryll f sqlh.l dor c! w snuD)ul '5$178wz'\celg I Lf z9z IIenI^I gl|stuptl "qrr\eg ( :|.Loo]]d)sntDDulolS ' 'o^eu5 6aZ 6eZ le^ul ulou ? eI.EIg xo lele{ C X FauP@8 tVZ' q\taq sns4liP re\eg g snn uaP 6tZ' '[onwD' l/is JZZ u ar{r./tq snqruel3Eld ' /lz qlueg OIooH) ,ptrarnatl'uopcos snqlueFeld

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