The Phytogeography, Ecology and Conservation Status of Lechenaultia R.Br

Kinsia 1(1): 85 133(198?) 85

The phytogeography, ecology and conservation status of Lechenaultia R.Br. (Goodeniaceae)

David A. Morrison

John Ray Herbarium, Botany Building (A12), University of Sydney,N.S.W. 2006.Present address: Depart ment of Applied Biology.New South WalesInstitute of Technology,P.O. Box 123,Broadway, N.S.W. 2007.

Abstract Mor son, D.A. The phytogeography, ecology and conseNation stai:us of rec,,renaulfia R.Br. (Goodeniaceae).Kingia 1(1): 85-133 (1987). Distribution maps for each of the 25 known speciesof LechenauLtiaarcpresented for 0.5' lat. x 0.5" long.g d cells,along with discussionsofthe habitat, floweing period, morphology,and conseNation status. Nineteen of the speciesare endemic (or nearly endemic)to south westernWestern Australia. An isoflor map ofthis region indicatesa high node of species-dchnesson the northem sandplains,with a band of moderatespecies-richness running south eastwardsto the southem sandplains.This species-richnessband followsthe 400mm annual rainfall isohyet,and conespondswith the kwonganheathlands and the wheatbelt. Ordination, clusterand discriminant function analysesof the distribution of these19 aechenaultiaspecies suggestsihat there are five biogeographicregions: one area on the iar nolthem sandplains,one on the nofhern sandplainsand northern part of the wheatbelt,one in the jarrah woodlandsand karri forests,one on the southem sandplains and southem part of the wheatbelt, and one in the inland mallee area. Discriminant. function analyses of t]le dist bution of these 19 Lechenaultia species among the phytogeographicregions of Beard and Barlow indicate that they are both accurate reflections df the Leche nauLtia distributions. Similarity coefficientsof speciespresence among the phltogeographicregions of Beard indicate that the Irwin, Darling andAvon BotanicalDistricts are more sinilar to eachother than they areto the Eyre and Roe BotanicalDistricts; while cladisticanalysis ofendemic species indicates that the Irwin, Roeand Eyre Districts havea more similar history. This suggeststhat there has beenfairly \idespread dispersalof somespecies in recent times. LechenaultiaLaricinais considered to be an endangeredspecies, with L. juncea,L. longiLobaand L. puLuinnris vulnerable,and, L. acutiLobaand, L. superbararc; L. chlorantha and L. ouor@are too poorly known for the conservationstatus to be known. Only three of these speciesare confidently known to be representedin conservationreserves, with two of the others thought to be present.

Introduction The Goodeniaceaeare well-representedin the flora of WesternAustralia (Marchant 1973),and they are one of the commonly encounteredfamilies in the heathlandsof the south-west (Georgeet al. 1979,Lamont et al. 1984).Within this family, Lechenaulti,a R.Br. is oneofthe most interesting genera.It is restrictedto Australia and New Guinea, with all of the 25 recognisedspecies occurring in Australia (Morrison 1986).Nineteen ofthe speciesare endernic (or nearly endemic)to south-westernWestern Australia, with one specieson North-West Cape,two speciesin arid western Central Australia, one in arid easternCentral Australia, one restdcted to a small areaof the Northern Territory, and one specieswidespread across tropical northern Australia and extending to the southern coast and off-shore islands of New Guinea. 86 Kingia Vol. 1, No. I (198?)

Most of the speciesare small perennial sub-shrubsor herbs,with only Lechenaultia acutiloba,L. dtDaricqtqand, L.linarioides becomingshrubs over 1 m high. The plants are widespreadbut never dominant components of the flora of south-westernWestern Australia, occurringin woodland,forest, and especiallyheath (Georgeet al. 1979,Specht 1981.Lamont et al. 1984).

This paper provides previously unpublished notes on the ecology,morphology, and conservationstatus of all ofthe species,as well as presentingthe distribution data. The phytogeographyof the South-West Botanical Province of Western Australia is also analysedin relation to the distribution and phylogeny of LechenquLtia.

Methods The datapresented were obtained from the herbariumcollections housed at AD, BRI, CANB, CBG, LD, MEL, NT, PERTH and SYD, plus the type collectionsof CGE and S (codesas in Holmgren et al. 1981),supplemented by field observationsof some 70% of the species(all from south-westernWestern Australia). The taxonomic schemeused wasthat of Morrison (1986).

The distribution of each specieswas recordedon a map of Australia subdividedinto a 0.5" Iatitude by 0.5" longitude grid, correspondingto the 1:100,000topographic survey maps of the Division of National Mapping, Commonwealth Department of Natural Resources.

Within the Lechenaultiq distribution, only the South-West Botanical Province of WesternAustralia containsenough species for the analysisofphltogeographic patterns in detail. Consequently,the phytogeographicanalyses concentrated on this area.

Two species-richness(isoflor) maps of the South-West Botanical province were derived from the distribution data. The first presentedthe speciesdistribution based solely on validated herbarium records.However, as this is presumablyan incomplete record,due to sparsecollecting in someareas and land clearancein others,a secondmap presentedthe distributions as they could be conservativelyinferred from the recorded distribution. For most species,this secondmap differed frorn the first only in oneor two grid cells; and this second map was assumedto be superior to the first as a true representationofthe species'distributions. It wastherefore these inferred distributions that were used in the ph)'togeographicanalyses.

The pattern of variation among the distributions of the 19 Lechenaultia species present in the South-West Botanical Province was analysedsimultaneously using an ordination technique,detrended correspondence analysis (Hill 19?9a),on the presence or absenceof the speciesin each of those 130 grid cells that occur in the South-West Botanical Province. This non-linear technique is reported to be superior to the traditional linear ordinations (Gauch et al. 1977,Hill and Gauch 1980,Gauch et al. 1981),and this appearsto be true for the data setsfor which I have madecomparisons. The pattern of vadation among the 130 grid cells was analysedby ordinating the presenceor absenceof each of the 79 LechenauLtiaspecies in each cell.

The clusteringpattern ofthe 130grid cellswas investigated using a clusteringstrategy, two-way indicator speciesanalysis (Hill 1979b),on the presenceor absenceof the 19 Lechenaulti.aspecies in eachgrid cell. This polythetic divisive techniqueis reportedto be superior to traditional agglomerativetechniques (Gauch and Whittaker 1981),and this aDnearsto be also true for the data sets that I have used. D.A. Morrison, The phytogeography,ecology and conservationstal;s of Lechenaultia- 87 The predictability of the phJ.togeographicclusters formed by this analysiswas tested using discriminant function analysis (Klecka 1975). This technique derives a small number of linear functions that weight the original discriminating variables (in this case,presence or absenceof eachof the 19 Lechenaultiospecies) so as to maximisethe separation of the total scoresof a set of referencesamples summed over all of the variables.In this case,the referencesamples consisted of eachofthe 0.5" x 0.5" grid cells that make up a particular phltogeographic cluster. Each of the grid cells was then individually re-classifiedusing the discriminant functions derived from the original analysis,to seeif the they were correctly classifiedinto the particular phl'togeographic region to which they were originally assigned.This technique effectively weights the speciesby constancywithin groups (ie. minimal within-group variance and maximal between-groupvariance), and this approach to assessingclassifications has been suggestedby severalpeople (Farris 1966,Johnson 1982).

The relationships between the Lechena.ultia species distdbutions and the phltogeographicregions of Beard (1980d)and Barlow (1985)that coverthe South-West Botanical Province (see Figures 6a and 6b) were also analysedusing discriminant function analysis.For the Barlow (1985)regions, assigning each of the 131grid cellsto a particular region was straightforward, as this systemis basedon a 1" x 1" grid; while for the Beard (1980d)regions, which are basedon vegetationphysiognomy, those grid cells near the boundariesof the regionswere assignedto the region that occupiedthe greatestportion of the cell. Once again, each of the grid cells was then individually re-classifiedusing the discriminant functions to seeif the they were correctly classified into the particular ph].togeographicregion to which they were originally assigned.A compadsonwas also madebetween the original Barlow (1984)regions and the modified regions of Barlow (1985), to see if there was any significant improvement in the predictivity of the modified regions comparedto the original.

The relationships among the five south-westernphytogeographic regions of Beard (1980d),as related to Lechenaultia, were analysedin two ways. Firstly, the overall superficialsimilarity was assessedusing two similadty indices:-the JaccardCoefficient and the Simple Matching Coefficient (Hubalek 1982).

Secondly,the historical relationshipsamong the regionswere analysedby converting tineLechenaultia morphologicalcladogram of Mordson (1987)into an areacladogram. This was done by simply superimposingthe phytogeographicregions of Beard (1980d) onto the cladogram(cf. Nelson and Platnick 1981).Unfortunately, the cladogramas it stands (Figure 7a) is uninformative (or falselyinformative; Platnick 1981)as far as the phytogeographicregions are concerned,because too many ofthe speciesoccur in several of the regions.Therefore, the areacladogram was reducedto only those 11 speciesthat are endemicto one of the phltogeographic regionsplus those 3 speciesthat are more characteristicof only one region (Figure 7b). While this potentially resultsin the lossof information (Platnick 1981),it does allow the cladogramto be resolvedto a simple informative five-area cladogram.Those speciesexcluded from the analysis were t. biloba,L. floribunda,L. formosa,L. macranthaa\d L. tubif\ora,which areall widespread generalistspecies, while L. acutiLobawas consideredt).pical of the Eyre region and l. linarioi.desand L. stenosepdlatlpical of the Irwin region.

The predominant geographical modes of speciation shown by Lechenaultia in Australia were investigatedby studying the distributions of pairs or groups of closely related species,as defined by the taxonomic system of Morrison (1986). Kingia Vol. 1, No. 1 (1987)

Results and Discussion The distribution maps,and the noteson morphology,ecology, and conservationstatus are presentedin Appendix 1. The speciespresent or presumedto be present in eachof the 0.5' x 0.5" grid cellsof south-westernWestern Australia arelisted in Appendix 2.

Most of the south-westernspecies occur in shrubland of some sort, although several speciesalso occur in woodland or mallee. Only L. biLobais known from forests. The inland speciesusually occur in open grasslandor woodland.AII of the speciesoccur in freely-draining sand or lateritic gravelly soils, except for L. expansa,which occurs in permanently wet areas.All of the speciesapparently have woody rootstocks,some of which can spreadlaterally. This may allow them to regeneratevegetatively after a fire or a dry spell.

The main floweringperiod for the genusis Octoberand November,as shownin Figure 1. However,many of the speciesflower sporadicallythroughout the year, perhapsonly after rain or a fire.

16

14

12 .9 310

E8

O Eo z 4

Month Figure 1- Number of-Lecierault'iaspecies floweingperweek in south-western Western Australia. Dsta from Appendix1.

Table 1 summarisesthe conservationstatus for those speciesconsidered to be at risk. Six ofthe species(24%) arethoughtto be under threat, all ofthem from the South-West Botanical Provinceof WesternAustralia. A further two speciesare too poorly known for their conservationstatus to be accuratelydetermined. Three of the speciesare thought not to occurin conservationreserves, while two othershave not beenrecently confirmed as present in them. Rye et al. (1980)and Burgman and Hopper (1982)report only two of these speciesto be exploited in the wild by the wildflower industry, although 32% of the total number of Lechenaultia speciesare exploited.Nearly all ofthis exploitation is for the nursery trade. D.A. Morrison, The phytogeography,ecology and conservationstatns ol LechenauLtia 89

Table l. Summary ofthe conservation daiafor t\e Lechenaultia speciesconsidered to be at risk.

Consen'ation Category + Leighet al. (1981)+ This study +

Extinct (X) Endangered(E) L. Iongiloba(C) L. pulvinaris L. kric;na Vulnerable(V) L. juncea_ L. juncea L. longiloba (.C?) 1-.pulv,aans (C) Rare (R) L. aattiloba (C) L. acutiloba (C) L. superba(C) L. superba\C) L. chlonntha (C) PoorlyKnown (K) L. chlorantha (C1], L. ovata

* SeeLeigh et al. (1981)for a detailed explanation + C - speciesknown from conservationreserves. ? not recently confirmed to be present

The two maps of the species-richnessisopleths are shown in Figures 2a and 2b. The isoflor map of Figure 2b indicates a high node of speciesrichness on the northern sandplains,with oyer 44% of the speciesoccurring in a gdd cell south-eastof Eneabba. This is in accordwith the commentsof Georgeet al. (1979),who considerLechenaultia to be characteristicspecies of the floristically rich heathlandbetween the Moore River and Dongara. A band of moderate species-richnessthen runs south-eastthrough the wheatbelt, and then extendseastward along the southern sandplainspast Esperance' The kwonganheathlands predominate on both the northern and southem sandplainsof this species-richarea, with eucalyptand banksiawoodlands occupying the intermediate wheatbelt.The Jarrah, Mard and Karri forestsofthe extremesouth-west are relatively species-poor,as are the woodlandsand shrublands of the inland area.

This band of speciesrichness roughly followsthe 400 mm annual rainfall isohyet,with the 800mm isohyetdefining the transition to the species-poorarea ofthe south-western forestsand the 300 mm isohyet defining the easternlimit of Lechenaultiodistribution. This distribution appearsto be a recurrent pattern at the genericlevel in the flora of south-westernAustralia, with many examplesappearing in a wide rangeof families (eg. Haemodoraceae,Proteaceae, Rutaceae, Mimosaceae, Myrtaceae; see Hopper 1979, Lamont et al. 1984).This has resultedin the recognitionof three species-richnesszones in south-westernAustralia:- one in the Eneabba-Mt Lesueur area, one in the Stirling Ranges,and onenear the FitzgeraldRiver (Georgeet al. 1979).It is thereforeinteresting to note that Lechenaultia does not appear to have the expectednode in the Stirling Ranges.

One olthe apparent reasonsfor the high speciesrichness in the kwonganheathlands is the high number of localised endemics compared to the wheatbelt and the species-poorareas. There are three extremely localisedendemics (L. chLorqntha,L. juncea and L. longiloba)in the northern kwongan and another three in the southern kwongan(L. acutilobq,L. heteromeraand. L. superba),whlle the wheatbeltonly has two localisedendemics (L. laricina and.L. puLDinaris)and the othet areasnone.

A comparisonof Figures 2a and 2b highlights severalapparently sparselycollected areasin the South-WestBotanical Province.The only specieswith any largeremaining disjunctions in its distribution is L. t'ormosa(Morrison 1986);the other gaps in the speciesdisiributions are therefore presumably the result of under-collection or the destruction of populations due to agricultural clearance. The most obvious under-collectedarea is the easternhalf of the Roe Botanical Distdct. This is an areaof mallee that has only recently becomeeasily accessibleto botanical collectors,and it Kingia Vol. 1, No. 1 (198?)

3 I't 3 r 3 2 3 22 3 22

SOUTH-WESTERN AUSTRALIA 5 5 o 300 5 E-- 5 5 kllometrea 5 5 o 43 43 ,| 2 34 ,| ?fa 2 - l.:_J 2 I JJ 2 2 I

Figure 2. Isoflor maps of Lechenaultiaspecies in 0.5' x 0.5'grid cells,based on validatedherbarium records (Figure 2a) and on inferred speciesdistdbutions (Figure 2b). Contour inteNal is 2 species.Data liom ADDendix2. D.A. Morrison, The phltogeography,ecology and consewation status of Lechenaultia. 91

' O brevifolia

O PaPillat a Oheteromera O suPerba Oacutiloba

o f ormosa

pulvinaris tubiflora o a o lar ic ina expansa a O biloba

stenosepala o f loribunda O a j uncea O linarioides

O hirsuta

O chlorantha macrantha ' a rongiroba

Figurc 3. Projection of the ,Lecienau.ltiaspecies onto axesrepresenting the filst two componentsftom the detrendedcorrespondence analysis nf the distribution data. The eigenvaluesassociated with these two componentswere 0.786and 0.360respectively. 92 Kinsia Vol. 1, No. I (1987)

a o oo a aao

o't' a

Fig!re 4. Projection ofrhe 0.5' y 0.5'grid cells onlu a\es reprejenling lhe tilst llro cumponenls rrom the detrended c, 'rrespondence anal) -iq ol r he di.r riburion dara. The eiqenvaluesassociarcd u ith these two components were 0.?86 and 0.360 respectively. The five ph]'togaographic groups frcm the clus ter analvsis are also shown, D.A. Monison,The phytogeography,ecology and conservation stat[a of Lechenaultia. 93 appears to be an area relatively rich in uncollected species. For example, the two newly described Lechenaultia species from south-western Australia are from this area (Morrison 1986). The other apparently poorly collected area is the northem tip of the Irwin Botanical District and the Shark Bay area, another area that has received only relatively recent botanical inspection. The area between Collie and Mt Barker in the south-west also shows gaps in many of the species distributions, but this is more likely to be a result of clearing for agriculture.

The ordination of the species distributions (Figure 3) indicates three main groupings of species. Those species that are restdcted to the Roe and Eyre Botanical Districts (-L. acutiloba, L. brevifolia, L. heteromera, L. papillata, and L. superba) ordinate together, as do most ofthose species common in the northern kwongan (-L. biloba, L. chlorantha, L. floibunda, L. hirsuta, L. juncea, L. linarioides, L. longilctba, L. macrantha and L. stenosepala). The two endemics ofthe Avon District (L. laricina and L- pulvraans) form the third group. The sole endernic ofthe south-western forests (L. expansa) and the two widespread species (L. formosa and L. tubiflora\ do not fit into any of the groups.

The first axis of this ordination accurately separates the north-western species from the south-eastern ones, while the second axis appears to reflect the rainfall gradient away from the coast. The distribution of the individual species thus also appears to be strongly influenced by rainfall, as well as the generic distribution.

The ordination ofthe grid cells (Figure 4) does not reveal any obvious clusters, indicat- ing that there are no clear-cut groupings of regions within the South-West Botanical Province as far as LecienauJrra is concerned. Once again, the first axis separates those gdd cells in the north-west from those in the south-east, while the second axis separates the coastal grid cells from the inland ones.

The clustering analysis reveals five main groupings ofthe grid cells (Figures 5a and bb); and the re-classification of the gdd cells in the discriminant function analysis of the clusters indicates that only 3.1% of the grid cells were incorrectly classified. This suggests that the cluster pattern has a high Ievel of predictability. However, while the projection ofthese clusters onto the ordination (Figure 4) shows that they are non- over- lapping, they are nonetheless somewhat arbitrary clusters in a continuum ofvariation.

These grid cell clusters correspond fairly closely to the biogeographic areas tradition- ally recognised in south-western Western Australia:- Groups l and 2 occur in the kwongan of the northern sandplains, with Group 2 extending into the northern part of the wheatbelt, Group 3 occurs in the kwongan of the southern sandplains and extends into the southern part of the wheatbelt, Group 4 occurs in the jarrah and marri woodlands and karri forests of the south-west, and Group 5 occurs in the inland mallee area. However, the cluster analysis fails to unite the wheatbelt as a single unit, and the nodhern sandplains are clearly divided into two areas. This latter pattern seems to be the result of a large number of localised endemics that are restricted to only one ofthese two areas. Finally, the analysis also suggeststhat the southern areas are more similar to each other than they are to the northern areas (Figure 5a).

The results of the re-classification ofthe grid cells in the discriminant function analy- ses (Tables 2 and 3) indicate that the ph5togeographic regions of both Beard (1980d) and Barlow (1985) are reasonably accurate reflections of the distribution of the Lechenaultia species.The discriminant functions correctly re-classified approxirnately 85% of the grid cellsin both cases(87.0% for the Beard regionsand 83.2%for the Barlow regions),and those grid cells incorrectly re-classifiedwere usually around the bound- ades of the regions (Figures 6a and 6b). KingiaVol. 1, No. 1 (1987) a

b -h\ia J \

L-, SOUTH-WESTERII aL ( ,AUSTRALIA

300 3Oo + -t kllomet.s s t L F-l J ( ( 4 I

^ -0 \ / I ll50 to0 .C Fisure5. Dendrosram 5b)ofthe five phytogeographic groups resulting from the 1il,'gJ:ii|:t1"T"iBJrl',Tre D.A. Morrison, The phytogeography, ecology and conseNation statns ol Lechenaultia 95

Table 2. Resutts of the re-classification of the grid cells of the Beard (1980d) phytogeographic regions using discrimiDant function analysis.

Phytogeographicregion Phytogeographic region of Beard Total no. predicted liom the of grid discriminant.function Avon Darling Ro€ Eyre cells

Irwin I 0 0,| 0 19 I 24 5 il7 Darling 0 I 32 0 0 33 Roe 0 0 22 0 22 Eyre 0 I I 0 15 t1

Table 3. Results of the re-classification of the grid cells of the Barlow (1984) and (1985) phytogeographic regions using discriminant function analysis.

Phytogeographicregion Phytogeographic region of Barlow Total no. predicted from the of grid discriminant lunction Bencubbin Leeuwin Esperance cells

Barlow(1984) Bencubbin 28 7 3 38 Leeuwin 2 32 2 36 Esperance 5 2 50 5'i (1985) Barlow ,7 Bencubbin 38 6 51 Leeuwin 3 32 2 37 2 39 43

For the Beard (1980d)regions, most of the incorrectly re-classifiedgrid cells were in the complexboundary area of the Avon, Darling, Eyre and Roe Botanical Districts to the north of the Stirling Ranges(Figure 6a). In particular, the analysisindicates that most of this area should be correctly classifiedas part of the Avon Botanical District. However, such a result is not unexpected,as the regionswere originally basedon veg- etation physiognomyrather than floristics (Beard 1980d),and there is thus no a prlorl reasonto assumethat they shouldaccurately reflect the distribution ofany oneparticu- lar genus.

On the other hand, the incorrectly re-classifiedgrid cellsfor the analysisofthe Barlow (1985)regions are around the boundadesof most of the regions(Figure 6b). In particu- Iar, the analysisindicates that most of the boundadesare displacedby 0.5" of latitude or longitude.This is in fact a very suggestiveresult, asthe regionswere originally based on a 1' gdd, and the analysisthus indicates that the regional boundariesare actually correctlyplaced at this scale.The Barlow (1985)regions appear to be slightly lesspredic- tive than the original Barlow (1984)regions, as the original regionswere 84.47ocorrectly re-classifiedby the discriminant function analysis (Table 3).

The similarity coefficients among the phytogeographicregions of Beard (1980d) (Table 4) indicate that the Irwin, Avon and Darling Botanical Districts are superficially very similar, asare the Roe and Eyre Botanical Districts. On the other hand,the reduced areacladogram (Figure 7c) indicatesthat the Irwin Botanical District has a more similar historv to that of the Roe and Eyre Botanical Districts. The history thus unites the northern and southernkwongan areas, even though they are now spaciallyseparated,

Such a relationshiphas beennoted for other generaas well, basedon the distribution of vicadous speciespairs and disjunctions in the distributions of species(see Nelson 1981,Lamont et al. 1984).The current disjunction betweenthe kwongan vegetation areashas been postulated to be the result of disruption ola previouslycontinuous range, Kingia Vol. 1, No. 1 (1987)

I \ Y I l,

\i\'*uY'*I 300 \ \ ME- qvoi \, I \ t\ t\/\ ' -'l -l '1 l) \ |.---t rE >-- q \l .= v &.+ EYRE DARLING 350 - t I 1 -0 r zloo t)s' NJ

I SOUTH.WESTERN

AU9TRALIA \ I I BENCUBIilN 300 !+ -...---...... - L. L'' kllomatro6 \t I +L: -r \

/\ \ J ESPERANCE + I t-\ I LEEUWIN --v 350 J I I 1150 1200 125c figure 6. The Soulh Wp.r Boraniral Province phy'ogeographi. regions of Beard !1980d) tFislre 6a, and Barlo$ {1985) lFiglre 6b). .howjng rhose grid rells incorreerly re classilipd b} rhe diseriminanr runcnonS. D.A. Morrison, The phytogeography,ecology and conservationstatns of LechenauLtia. 97

o Avon, Ey re, lrwin, Boe trwrn

Darlin9, lrwin

Eyre

Eyre, Roe

Avon, Darling, Eyre, lrwin

lrwrn longiloba

Avon, lrwrn

AVon

Boe

Avon, Darling, lrwin

AVOn

Da I ting

Avon, Darlins, trwin, Roe

Darling, lrwin

*

Jt

Eyre * *

o

Eyre

EYre

Eyre

Darlrng

Darling

Eyre

Figure 7. Area cladogramof all of the l-ecfieraultia species(Figure 7a), those ,ec,henaultiaspecies that are endemicto only one Botanical District of the South West Botanical Province (Figure?b) and the reducedarea cladogram from Figure ?b (Figure7c), showingthe Botanical Districts of the South West Botanical Province,in which they occur. * Not in South-WestBotanical Province. Kingia Vol. 1, No. 1 (198?)

Table 4. Percentage similarity of the phytogeographic regions of Beard (1980d).The upper half of the similarity matrix contairs the simple vr'hile the lower half contains rhe U?:"i:1cfrrT:r"T*"t

Phylogeographic Region Phytogeographic region Irrvin Avon Darling Roe Eyre

lr\\,in 57.9 63.2 36.8 36.E 38.5 63.2 57.9 5?.9 Darling 50.0 27.3 52.6 52.6 Roe 14.3 20.0 20.0 68.1 Eyre 14.3 20.0 20.0 25.0

this continuous range itselfbeing the result ofcolonisation ofthe whole areaby laterite- tolerant species after the area was subjected to lateritisation during the Miocene (Marchant 1973,Lamont et al. 198,1).The causeof the disjunction is postulated to be the onset of more arid conditions during the Holocene (Nelson 1981, Hopkins et al. 1983,Lamont et al. 1984).The expansionof the drier regimeswould have resultedin the extinction of the kwongan species, leaving the upland areas as refugia and subsequent centres of speciation. The current superficial floristic similarity of the northern kwongan with the forests of the south west and the woodlands of the wheatbelt could thus be the result of very recent dispersal of species into these regions, particularly from the north.

Figure 7a indicates that L. subcymosa, which occurs on Nofth West Cape and the Shark Bay islands, is closely related to the arid zone species (particularlv L. divaricata), and is only more distantly related to the south-western species.Thus, this does not sup- port Burbidge and George's(1978) use of this speciesas an example of the closetaxo- nomic affinity of the Shark Bay area ivith that of the South-West Botanical Province rather than with that of the Eremaean Province. This area should indeed. as thev "transitional suggest,be part of a zone".

The analysis of the geographical distributions of the groups of closely related species reveals that within the South-West Botanical Province the congeners are frequently allopatric, sometimes sympatric, and rarely parapatric (Table 5). This suggeststhat, in this area, geographical isolation has probably played a major role in speciation within Lechenaultia; and a similar pattern has been found for Acacia (Hopper and Maslin 1978).Outside this region, most of the congenersare allopatdc, which is in contrast to the finding of Maslin and Hopper (1982) that sympatry and allopatry are about equally common in central Australia.

Table 5. Geographical relationships of the closely related species gro'rps ol Lechenaultia.

Speciesgroup Geographicdistribution Allopar.ric Parapatric Sympatnc

Sect. Patentes L. bilLtba and L. stenasepala l. erpar.sa and L. pulvinaris L. floribtnda and L. papillata Sect- Latouria L. hetenmera and, L. lutes.ens L. divaricaLa and 1,. subcyrrosa 1,. brcvilblia, L. juncea and L. striata L. lilitormis and L. ovata Sect. I-echenaultia L. hirsula, L. laricina and L. superba L. longiloba and L. macnntha L. acutiloba anc\ L. tubiflora L. chloantha and L. formosa D.A. Morrison, The phytogeography, ecologf and conseNation stat:us of Le.:henaultia 99

Acknowledgements Thanks to the Directors and Curators of the herbaria consulted for the use of their specimens;to Chas Chapman, Neville Marchant, Ken Newbey, Sue Patrick, and Rae Paynter Ibr all their help in Western Australia; to Ken Newbey for his time and help in improving the distribution data; to Malcolm Ricketts for help with the figures; to Kerri Gallagher for helping throughout; to the A.S.B.S. Sydney Chapter for helpful dis- cussionson this work; and to Tony Auld, Roger Carolin, Peter Myerscough and Peter Weston for kindly commenting on an earlier version of this manuscript. The research was funded by the University of Sydney, the Linnean Society of N.S.W. (through the Joyce W. Vickery Scientific ResearchFund), and the Australian Biological Resources Study.

References Anon.(1981). More rare flora illustrated. Swans 11(2): 12 13. A.P.S.G.(r980). Grow What Where. Thomas Nelson, Melbourne. Baglin,D. and Mullins,B. (1969)."Australian Wildflowers in Colour".Reed, Sydney. Baird, A.M. (19?7).Regeneration after fire in King's Park, Perth, WestemAustralia. Journal ofthe Royal So .iFrt I \\'e:rernAJsrralia A0: | 2- Baker, K.F. (19?1)."lVildflowers o1 Western Australia". Rigby, Perth. tsarlow,B-A. (1984)-Proposal for delineationof botanical regionsin Australia. Brunonia 7: 195'201. Barlrw, B.A. (1985)A revisednatural regions map for Australia.Brunonia 8:387-392. Beadle,N.C.W. (1981)."The Vegetationof Australia". CambridgeUniveNity Press,Cambridge. Beard,J.S. ( 1969).The vegetati(nrofthe Boorabbinand Lake Johnstonareas, Western Australia. Proceedings of the Linnean Society of New South Wales 93: 239-269. Beard, J. (ed.) 1970."A Descdptive Catalogueol West Australian Plants.'' 2nd ed. for Growing Australian Plants,Sydney. Beard,J.S. (1972)."The Vegetationofthe HydenArea, Western Australia". Vegmap Publications, Pefth. Beard. J.S. (197:la). 'The Vegetationof the RavensthorpeArea. WesternAustralia". VegmapPublications, I'erth. Beard,J.S.(19?3b).'TheVeg€tationofth€EsperanceandMalcolmAreas,WesternAustralia".VegmapPub lications. Perth. Beard,J.S. (1976a). "The Vegetationof the Newdegateand Bremer Bay Areas, Westem Australia", 2nd edn. \regmap I'ublications, Pefth. Beard,J.S. (1976b)."'l'he Vegetationof the Shark Bay and Edel Areas,Western Australia". VegmapPubli cations.Perth. Beard,J.S. (1976c)."The Vegetationofthe Ajana Area,Western Australia". Vegmap Publications, Pefth. Beard,J.S. (1976d)."The Vegetattunofthe DongaraArea, Western Australia". Vegmap Publications, Perth. Beard,J.S. (1976e)."The Vegetationofthe PerenjoriArea, Weslern Allstralia". Vegmap Publications, Perth. Beard,J.S. (19?9a)."The Vesetationofthe Moora and Hill River Areas,West€rn Australia". VegmapPubli- cations.Perth. Beard,J.S. (1979b)."'l'he Vegetationot'the Pefth Area,Western Australia '. VegmapPublications, Perth. Beard,J.S. (19?9c)."The Vegetationofthe Albany and Mt. Barker Areas,Westem Australia". Vegmap Publi- cati(ins.Perth. Beard, J.S. (1980a)."The Vegetationof lhe Kellerberrin Area. Western Australia". VegmapPublications, Perth. Beard,J.S. (1980b)."The Vegetationofth€ Corrigin Area,Western Australia". Vegmap Publications, P€rth. Beard, J.S. (1980c)."'l'he Veeetationof the Dumblenrng Area, Westem Australia". Vesmap Publications, Perth. B€ard,J.S. and Burns. A.C. (1976). "The Vegetationofthe GeraldtonArea, Westem Australia". Vegmap Pub- licati{)l1s.Perth. Bell, D.T- Loneragen,W.A. and Dodd, J. (19?9).Preliminary vegetationsurvey ol Star Swampand vicinily, Western Australia. Western Australian Herbarilrm ResearchNotes No. 2: 1 21. BLrmberv.A.M. (1973)."What Wildfloweris That?" Paul Hamlyn,Sydney. 100 Kingia Vol. 1, No. I (1987)

Boyland.D.E. (1910).Ecglogical and fl,,nsticsrudies in rhe SimpsonDesert National Park, south western Q r,pr.tldnd. Pro. eedirgs uf rh. R,,lal50. i"rt uf Q,.eFn-la,d B : I tC. Bridgewater,P.B. and Zammit, C.A. (19?9).Phytosociology of S W Australian limestoneheaths. Phytocoenologia6: 32?34:. Brown,J.M._and Hopkins, A.J.M. (1983). The kwonsan(sclerophyllous shrublands) ofTutanntug Natur€ Re- serve.Western Australia. Australian Joumal of Ecology8: 63 ?3. Buckle1,,R.C' (1981a).Soils and vegetatirrn ,r1 central Australhn sandridges IL Sand dgevegetation in the Uluru NationalPark area,Northern Territ,,rt. Ausiraha.Ausrratian ,Iournal of Ecotogt6: 391,d04. Buckle,v,R.C. (1981b). Soils a]rd vegetation ol centralAustralian -cndridges lll Sandridgevegetation of the s.m,,.,'nDFcerl Auq "alian .l,.urnel ot F.ulu$ h 1^ ,-4'n. Buckley, R.C. (1982).Central Australian sand ridge flora 18,000years ago:phytogeographic evidence. In: "Evolution of the Flora and Fauna of Arid Australia". Barker, W.R. and Greenslade,P.J.M. (eds) PeacockPublications, .A.delaide. pp. 107117. Buckley.R.C. (198:l).The flora and vegetationof Barrow Island.Westem Australia.Joumal of the Roval So cietv of Western Australia 66: 91 1{15. Burbidge,AA. andGeorge. A-S. (1978). The floraand fauna ofDirk HartLrgIsland, Westem Australia. Joumal oI ihe RoyalSocietv of WesternAustralia 60: ?1,90. Bur{man, M.A. and-Ho_ppe\S.D. (1982).'fhe WesternAustraljan Wildflower Industry 1980-81.Department Fisheriesand WiidliieWestem Australia. Report No. 53. ',Ptants Cunningham.G.M.. Mulham, W.E., Milthorpe, P.L. andLeigh, J.H. (1981). of WestemNew South Wales". GovernmentPrinter, Sl dney. I)odd.J.. Heddle.U. \1.. Iratc,J. S. andDixon, K. W. (1981).Rooting patterns of sandplainplants and their iunctn)nalsisniiicanc€. ID: Krvongan. Irlant Life (rftheSandplain. Pate, J.S. and Beard J.S. (eds) Unn,er- sitv of NesternAustralia Press. Pcrth. pp. 1.161i?. "Encyclopaedia Elliot, W.R. and Jones,D.L. ( 1982). of Australian Plants Suitablefor Cultivation,,.Volume 1. Lothian.Melbourne. Erickson,R.. George, A.S., Nlarchant, N.G. and Morcombe, M.K. (1979)."Flowers and Plants ofWestern Aus tralia , 2nd edn. Reed,Sydney. "West Fairali. A.R. (1970). Australian Native Plants in Cultivation". PergamonPress, Sydney. Farris.J. S. (1966 ) . E stim atio n of conseIvationof characteIs by constancv.!'! ithin biologicalpopulations. Evo I ui,ion20:58?-591. C,rdn-r..4.A.'l9alrThcreee'arion,lWp-r"rn{rsrrali€!\'irhspe,.ralretFrpn.p.,,.li.na,p anos^i1.. Jo-fnal ,t rrrFtt^\':rt 5u, r,'\ ,. w..,er4 Au- ratir,.8 ri lj,xr\ii. "\tildflowers Gardner,C.A. (19i8). of WesternAustralia", 13th edn. (ed. J. Edmonds).West Australian Nervs papers,Perth. Gauch.H.C. and Whittaker, R.H. (1981).Hierarchial classification ofcommunity data. Joumal ofEcology 69: 5:l7557. Gauch,H.G., Whittaker, R.H. and Singer, S.B. (1981). Acomparative study ofnonmet c ordinations.Jou rnal of Ecoloev69: 1351;2. Caucl,.H.C., Wh;ttaker, R.H. and Wentworrh,T.R. (19??1.A comparativestudy of reciprocalaveraging and ''rheror,l rari,,n e"hriqun..J,.-fnil or E.^1,,$bt.. t5, t:4. George,A.S.. Hopkins,A.J.M. and Marchant,N.G. (1979).The heathlandsof WesternAustralia. In: Ecosystemsofthe World, Volume 9A. Heathlandsand RelatedShrubtands. Desc ptive Studies.Specht. R.L. (ed.),Elsevier. Amsterdam. pp.211,230. "How Grieve,B,.J.and Blackq4,!'.Fj. (1q75). to Know WesternAustralian Wildflowels, part IV,,. University of WesternAustraiia Press. Perth. Griffin-E.A. gnd Hopkins. A.J.M. (19851 Thefloraand\egetationolMtLesueur.WestemAustralia.Journal .lth. R"1.r Surrpry,,r\\'e.rc. r A.-rralirrr. ,15j;. Criffin, EA.,.HopkiA., 4.:IM. and Hnatiuk, R.,L.(1983). Regional variation in meditenanean-rype shrublandsnear Eneabba, south-western Australia. Vegetati;52: 10: 12?. Hartley, W and Leigh, J. (19?9).Plants ai Risk in Austratia.Australian National Parks and Wildlife Sewice OccasionalPaoer No. 3. Hil1.M.O. (1979a).tIECORANA - A FORTRAN ProgramIur Delrcndedaorrespondence Analysis and Re- ,ipr,.alAro.adrng r-,.rrrel I rl\"r,irr. \"w \.r1. Hill, M.O. (1979b).TWINSPAN A FORTRAN Prrgram tor ArrangingMuttivariate Data in an Ordered Two-way Table by Classificati{)not lhc Individuals and Aftribuies. tj"rnell Universitv. New york. Hill' Nl o and Gauch,H.G. (1980).Detrended correspondence analysis: an improved ordination technique. Vegetatio42: .17-58. "A Hodgson,M. and Paine, R. (19?1). Field Guide to Australian Wildflorvers",volume 1. Riebv. perth. "Australian Holliday,I. (197:l). Shrubs".Risbv. Pedh. D.A. Morrison. The phltogeography.ecologl, and conseNation status of Lechenauliia 101

Holmgren,R.K., Keuken. W. andSchofieid, E.K. (1981)."Index Herbariorum", part 1.The Herbadaolthe World, Junk, The Hazue. Hopkins,A.J.M. and Hnatiuk, R.J. (1981). An eLologicalsun e) of the kq ungansouth of Eneabba.Western Au"tralra.\'ildl.le Re-earch Brlle 'r \\e. err Au.rral,d9: I l,l. Hopkins,A.J.M., Keighery, G.,I. and Marchant. N-C. (l98:). Species chuplands of southwest€ rn Australia. Proceedingsof the EcologicalSociety of Australia12: 15-26. Hopper,S.D. (19?9). Biogeographical aspects ofspeciation in the southwestAustralian flora. Annuat Review of Ecologyaud Systematics10: 399 422. Hopper,S.D. and Maslin, B.R. (1978). Phytogeography of Acaciain WestemAustralia_ Australian Joumal of Botany 26: 6:178. Hubalek, Z. (1982).Coefficients of associationand simila ty, basedon binary (presence-absence)data: an evaluation.Biological Reviews ii: 669-689. "Flora Jessop,J.P. (ed.)(1981). of CentralAustralia". Reed, Sydnev. "Flora Jessop,J.l'- andToelken, H.R. (eds)(1986). of SouthALrstralia",4th ed. Government Pinter. Adelaide. Johnson.R.W. (1982).Effect of rveightiDgand th€ size of the attribute set in nume cal classification.Aus_ tralianJ(rurnal of Botany30: 1611?4. Klecka, W.R. (19?5),Discriminant analysis-Inl, Sratistical Packagefor the Social Sciences.2nd edn. Nie. N.H., Hull. C.H.,Jenkins, J.C., Steinbrunner, K. and Bent. D.H. (eds).Mccraw Hi , New york. pp. 4314tr;. Lairlont,B.B. (i984). Sp,ecialised modes ofnut tion.In: K$,ongan.plantLifeofrhe Sandplain.pate,J.S. and Beard.J.S. (eds).UniveNity of WestemAustralia Press;pedh. pp. 126145. I-amont,B.B., Hopkins, A.J.M. and Hnatiuk, R.J. (1984). The flora composition,diversity and origins. In: Kwongan. Plant Life of the Sandplain.Pate, J.S. and Beard,J.S. (eds).UniveNity of Western Australia Press.Perth. pp. 27 50. "Extinct Leigh, J-. Boden, R. and Briggs, J. (1984). and EndangeredPtants of Australia". Macmillan, Melbourne.

Leigh,J,Rriggs,Jand Ha_rtlev, w. (r9a1).Rare o r ThreatenedAustralian plan rs. Australian National parks and \Yildlile ServiceSpecial Publication No. 7. Maconochie,J-R. (1982). Regeneration of a d zoneplants: a floristicsuNey. In: EvolutionoftheFloraand FaunaofArid Australia.Barker, W.R. and Greens.NL (eds). Peacock Publicarions, Adelaide. pp.141 144. Maier,J.I).11381)A fl,'u.erinq ralendar lbr Karragullen,a DorthernJanah forestlocality. Westem Austratian HPrbrfiLmH."-dfch \oLe. 5: 19-28. Marchant,N.G (197J).speciesdiversit)inrhesouthrvesternflora.JournaloftheRoyalsoci€tyofwestern Australia56: 2:l :10. Nlarchant,N.G,and K€igherv. G.J. 119]9). Pooriv cotlected and presumably rare vasc'lar prants in western Australia.Kings Parh ResearchNotes 5: 1 103. Maslin,B.R. and Hopper,S.D. (1982). Phytogeography of Acacia(Leguminosae: Mimosoideae) in central Australia. In: Evolution of the Flora and FaunaofAdd Australia.Barker, W.R. and Greenslade.P.,LM. (eds).Peacock Publications. Adelaide. pp. 301-315. Melville, R. (1973).Relict plants in the Australian flora and their conservation.In: Nature Conservationin the Pacific Region.Costin, A.B. and Groves,R.H. (eds).Australian National Universitv Press,Oanberra- pp.8il-90. Milewski.A.V. and Davidge,O. (1981).The physicalenvironment, tloristics andphenology ofa Banksiawood land nearPerth, Westem Australia. Western,{ustralian Herbarium Research Notes No.5: 29 48. "ALrstralia Morcombe,l,,I.K- (196E). s Western WildfloweN". Landfall. Perth. "Australia's Morcombe,NLX. (1970). Wildllowers".Landsdowne. Melbourne. "Wildflowers Morcombe,M. and Morcombe,L (19?0). of WesternAustralia". Lansdowne. perth. Morrison, D.A. (1986).Taxonomic and nomenclaturalnotes on -Lecierault'raR.Br. (Goodeniaceae).Brunonia 9:128. Nlorrison. D.A. (198?)Notes on the fruits of Lec.herau/t]aR. Br. (Goodeniaceae),with a new speciesfrom northem Australia. Telopea3: in press. NIuir B.a,.ir9;6|Bil,1,rqiraIsrne1ofthe we stern,{usrralian $'heatb€lt. pa.t r. Tain ri.ockanc] Norh 'l'arin HockHes€r\es. \egetatiorr. Rec,,rds oi the West€rnAustralian Museum Supplemeut 2. trILrir,R.Ctr9?-n Br,,L,git_nl "ur.rer^,rithe weslern Australian wheatbelt. part2. Vegetationand habitar of BenLlenngHeser!e- Re(ords uf !he WesternAustralian Museum Supplement 3. (19i9) part yornaning Muir, a.G. Biological.suNey of the Wesrem Australian whearbett. 7. Vegetationoi '- Nature lleserve.Records of the Westem ,{ustralian Museum SupplementB. Mullins. B. and tsaglin,D. (1978)."Western Australian Wildflowers in Colour". Reed,Sydney. 102 Kingia Vol. 1, No. 1 (198?)

Nelson,E.C. (1981)"Phytogeography ofsouthern Australia.In: EcologicalBiogeography ofAustralia". Keast, A. (ed.).Junk, The Hague.pp. 733'759. Nelson, G. and Piatnick, N. (1981)."Systematics and Biogeography.Cladistics and Vicariance".Columbia Univelsity Press,New York. Newbey,K. (1968)."West Australian Wildflowers for Hofticulturc", part 1. Societyfor Growing Australian Plants, Sydney. Patrick, S.J. and Hopper, S.D. (1982).Guide to the GazettedRare Flora of WesternAustmlia: Supplement 1. Depatment Fisheriesand Wildlife Westem Australia, Report No. 54. Platnick, N.I. (1981)."Widespread taxa and biogeographiccongiuence". In: Advancesin Cladistics.Funk, V.A. and Brooks, D.R. (eds).New York Botanical Garden,New York. pp.223 227. Rye,B.L. (1981).Ceographically Restricted Plants of SouthernWestern Australia. Department Fisheries and Wildlile Westem Australia, Report No. 49. Rye, B.L. and Hopper, S.D. (1981).A Guide to the GazettedRare Flora of Western Australia. Department Fisheriesand Wildlife Western Australia, Repoft No. 42. Rye.B.L., Hopper, S.D.and Watson,L.E. (1980).Commercially Exploited VascularPlant Native in Westem Australia; Census,Atlas and Preliminary Assessmentof Consenation Status.Depaftment Fishedesand Wildlife Western Australia, Repoft No. 40. Scott, J.K. (1981).A Survey Method for Identilying RoadsideFlora Suitable for Conservationin South Western Australia. Department Fisheriesand Wiidlife Western Australia, Report No. 41. Speck, N.H. and Baird, A.M. (198,1).Vegetation of Yule Brook Reserve,near Perth, Western Australia. Journal of the Royal Society of Western Australia 66: 147-162. Specht,R.L. (1981)."Major vegetationformations in Austmlia". In: EcologicalBiogeography of Australia. Keast,A. (ed.)Junk, The Hague.pp. 163-297. Specht,R.L., Roe, E.M. and Boughton,V.H. (19?4).Conservation of major plant communitiesin Austraiia and Papua New Guinea.Australian Journal of Botany Supplement?. Williams, R.F. (1932) An ecologicalanalysis ofthe plant communitiesofthe "Jarrah" regionocculling on a small area near Darlington. Joumal of the Royal Society of Westen Australia 18: 105-124. Wrigley, J.W. and Fagg,M. (1979)."Australian Native Plants". Collins, Sydnev. D-A- Morrison, The phytogeography, ecologJ and conservation status of l€.henaultra. 103

Appendix1. Notes on the distribution,ecology, morphology and conservationstatus of Lechenavltia

The morphological notes are based on personal observation of the herbariurn collections, supplemented by field observations of the species. Distributional and ecological data are from details on the herbarium labels, also supplemented by personal field observations.References which contain more details of some of these aspects(eg. more detailed descdptions of specific habitats) are also cited. Distributional data were derived from the point distributions of the specimens examined, and are mapped for grids of 0.5' lat. x 0.5" long. The distributions are also recorded for the standard botanical regions currently used by each of the state herbaria, and also for the more recent Australian phytogeographicregions ofBarlow (1985).The conservationstatus of each speciesis expressedusing the schemeof Leigh et al. (1981),supplemented where necessary by more detailed observations; and the number of specimens examined is given, to provide some measureof speciesabundance. The most widely-used common name is also recorded, where known; and a list ofchecked illustrations is provided, to aid with the identiticarionol the species.

Lechenaultia acutiloba Benth.. Fl. Austral. 4:41 (1868)

Distribution (Figure 8) Western Australia: Roe and Eyre Districts, between Ravensthorpe and Ongerup.

Esperance Region.

Ecology A small shrub with a woody rootstock that may sucker. It occurs in sand or sandy gravel,usually in damp soil near dver banks or occasionallyswamps. It is found in open floodplains or heath, where it is never very common. Flowering is from mid September to late December.

Notes Closely related lo L. tubiflora, from which it can be readily distinguished by the shrubby habit, larger green flowers, and wingless corolla lobes.

S)ecimens Examined

ConseNation Status 2RC, as it is geographicallylocalised (Marchant and Keighery 1979,Rye 1981) and restricted to a specific habitat. The river banks along which it usually occuls are, in many cases,parts of water conservation reserves;but these reservesare all extremely small, and are thus of only limited use as flora conservation areas. The species may also occur in Fitzgerald River National Park, but this has not been confirmed. It is exploited for the nursery trade (Rye et al. 1980), and is known to be in cultivation (Rye 1981). Newbey (1968) and Fairall (1970) discussits cultivation requirements.

Common Name Wineless lechenaultia. 104 KingiaVol. 1, No. 1 (198i)

Lechenaultia biloba Lindley, Sketch Veg. Swan R. 2? (1839)

Distribution (Figure 9) Western Australia: Irwin, Avon, Darling, and Roe Districts.

Bencubbin,Leeuwin, and EsperanceRegions.

Ecology A sub-shrubor small shrub with a woody rootstock(cf. Majer 1981).It has been recordedfrom white, grey, yellow, orangeand light brown sand, or sandy loam (Muir 1976,1977), often over laterite (Griffin and Hopkins 198b),as well as fiom sravel. It occursin a widerange of heaths(Beard 1976d, 1929a, 1980b, 1980c, Muir f979,Beadle 1981,Hopkins and Hnatiuk 1981,Brown and Hopkins1988, Griffin et al. 19g3,Griffin and Hopkins 1985),scrubs (Beard 1926e,1980b), mallee (Muir 1976,19?i, Beard 1,280b),woodlands (Beard 19?9a,19?9b), and (mainly Jarrah and Marri) forests (Williams 1932,Majer 1981),usually occurringin open patchesor evendisturbed areas (Muir 1977).It is often locally common in the westernpart of its range,but it usually occursas scatteredplants in the inland areas.Flowering is usually from early August to mid November(cf. Majer 1981,Milewski and Davidge1981); and it hasbeen reiorded to flower in the first seasonafter a March fire (Majer 1981).In the Jarrah forests.the roots showa high levelofinfection by vesicular-arbuscularmycorrhizae (Lamont 1gg4).

Variability Flower and leafsizeas well asflower colour vary greatly (seeMorrison 19g6),but these charactersdo not appear to be correlated.

Notes _.Closely-related to L. expansa and L. stenosepala,from which it can be readily distinguishedby the (usually)darker blue flowers with much broaderwinss and a mor; hairy floral tube. In inland areas it can look similar Lo L. brevifolia, which has a distilrctly scapigerousflowering habit and much reducedwings on the superiorlobes of the flowers.

SpecimensExamined 343.

ConservationStatus Not at risk, as it is widespreadand common throughout South Western Australia. It is exploitedfor the nurseryand seedtrades (Rye et al. 1980,Burgman and Hopper l9g2).

Common Name BIue lechenaultia.

Illustrations _Aryn.(196q)Aust. Pl.3:141; Morcombe (1968) p.12; Newbey (1968) pp.112_3,12b; Baglinand Mullins (1969)p.89; Beard (1970)pl.xxxi; Fairall (1-920)p.18i; Morcombe and Morcombe(1970) p.80; Baker (1921)p.19; Hodgson and paine (i9Zt)'p.89;Anon. (1973) Aust.Pl.7:115;Blombery (1973) p.190;Holtiday (1973) p.16; Grieve and Iilackall (1975) pl.v; Green& Wittwer (19?6)Ausr. P1.8:829; Gardner (t9i8) pp.5,130;Mu ins andBaglin (1978) pl.75; Erickson et al. (1979)p.46; A.p.S.G. (1980) betwe"" pp. fZO_2. D.A. Morrison, The phytogeography, ecology and conseNation status of te.henduh;d. 105

Lechenaultia brevifolia D.A. Morrison,Brunonia 9:18 (1987)

Distribution (Figure 10) Western Australia: Coolgardieand Roe Districts.

Leonora,Bencubbin, and EsperanceRegions.

Ecology A sub-shrubwith a woodyrootstock that may sucker.It hasbeen found on deepyellow or shallowred sand,usually in low scrubor heath(Beard 1969, as l. sp.inedit.). Plants are usually scattered,but are sometimescommon in patches.Flowering is from late Octoberto earlyDecember.

Variability There is somevariablity in flower and ovary sizebetween populations, but this does not seemto be significant.

Notes Closely related to L. heteromera,from which it can be readily distinguishedby the distinctly scapigerousflowering stemsand much darker flowers,and to -L.juncea and.L. sr.riara,which have much longer leavesand paler flowers.

Specimers Examined 27.

ConservationStatus Not at risk, as it is widespreadand relatively common throughout the south-eastern malleeof South WesternAustralia.

Lechenaultia chlorantha F. Muell.,Fragm. 2:20 (1860)

Distribution (Fieure 11) Western Australia: Irwin District, near the mouth of the Murchison River.

BencubbinRegion.

Ecology A sub-shrubwith a woody rootstock that may sucker.Little data exist on the habitat of this species,but it has beenrecorded from exposedred sandstonebreakaways, where it is apparentlyuncommon. Flowering specimens have been collected in Augustand September.

Notes Closely related to L. Iormosa and to ,L. linarioides,from which it can be readily distinguishedby the greencorolla.

SpecimensExamined 7.

ConservationStatus 2KC, asit is very geographicallylocalised (Rye 1981) and not locallycommon, and it is currently known from only one small population. It has also been recorded from Kalbarri National Park (Beard 1976c);but it is not known how extensivethese

54609-8 106 KingiaVol. 1, No. 1 (198?) populations are, and the specieshas not been recently confirmed to be present. This speciesis very poorly collected(Marchant and Keighery 1979),and much more needsto be known before the conservation status can be accurately assessed.If a viable representationin a conservationarea is demonstratedthen it would be more accurately classifiedas 2RC. It is not known to be in wide cultivation (cf. Rye 1981),but seealso A.P.S.G.(1980).

Lechenaultia divaricata F. Muell.,Fragm. 3:33 (1862)

Distibution (Figure 12) Northern Territory: Central Australia North and Central Australia South Regions; Queensland:Gregory North and Gregory South Districts; South Australia: Lake Eyre Basin Region; New South Wales: North Far Western Plains Sub-division.

MacDonnell,Thomson, Simpson, Cooper, and Torrens Regions.

Ecology A small shrub with a woody rootstock that may sucker (Maconochie1982). The fruit is woody and often persistentfor severalyears. It has beenrecorded from deepred sand dunes (where it occurs on the upper slopesand crests; Buckley 1981b),sandplains, alluvial soils,or sometirnesin flood-plains (Boyland 1970),swales, or other periodically wetdepressions. It occursin opengrassland,low open woodland (Boyland 19?0), or open mulga (Cunninghamet al. 1981).It usuallyoccurs as scatteredbushes, but is sometimes locally common. The plants appearto have high drought tolerance (Buckley 1982).It apparentlyflowers sporadically throughout the year.

Variability Throughout its range,there is considerablevariability in flower colourand size,but the speciesnevertheless remains distinctive. However,specimens are often mis-identified (eg.Buckley 1983, who recordsthis speciesoffthe WesternAustralian coast), often in confusion with, for example,Scaevola depauperata or S. spinescens.Within any one individual plant the number of articles per pseudocapsule(and hence pseudocapsule size) vades greatly, unlike any other speciesexcept L. filiformis

Notes Closely related to L. subcymosa and L. lutescens.from which it can be readilv distinguishedby the divaricate almost leaflesshabit.

Specr'mensExamined 725.

ConservationStatus Not at risk, as it is apparently common and widespreadthroughout arid eastern Central Australia. Specht et al. (19?4)report it to be rare in South Australia, but this doesnot appearto be the case.However, Melville (1973) and Spechtet al. (19?4)list it as a relict speciesthat has retained primitive morphological features, and that is therefore worthy of particular attention.

Common Name Tangled lechenaultia.

Illustrations Cunninghamet al.(1981) p.636; Jessop (1981) p.360;Jessop & Toelken(1986) p.1407. D.A. Morrison, The phytogeography, ecology and conservation status of aechenoultro 107 Lechenaultia expansa R.Br., Prodr. 1:581(1810)

Distribution (Figure 14) Western Australia: Darling Distdct. Leeuwin Region.

Ecology A sub-shrubwith a woody rootstock. It has been recordedfrom sand or peaty sand, where it usually occurs in swamp heath, around the edgeof paperbark swamps,or in other permanently damp or seasonallywaterlogged areas (Speck and Baird 1984).Ii is sometimesIocally common. Flowering is from Iate Octoberto mid January.

Variability Flower and pseudocapsulesize are very constant, but leaf size can vary considerably betweenpopulations.

Nores Closelyrelated to L. floribunda and often confusedwith it, particularly in the area around Perth where the two distributions overlap.However, 1,. erponso has larger and often minutely pitted leaves,a more denselyhairy floral tube, smallerand almost equal corollawings, a morethickly hairy indusium, and smallerpseudocapsules with fewerand ovoidal articles.

SpecimensExamined 96.

ConsewationStatus Not at risk, as it is widespreadand apparently common throughout the Jarrah and Marri forests of South Western Australia. It is exploited for the nursery trade (Rye et al. 1980).

Lechenaultia filiformis R.Br., Prodr. 1:581(1810)

Distribution (Fie!re 13) Western Australia: Gardner, Fitzgerald, and Hall Districts; Northern Territory: Victoria River, Darwin and Gulf, and Barkly Tableland Regions;Queensland; Burke, Cook, and North Kennedy Distdcts. Also Papua-NewGuinea, along the south'eastern coast. Kimberley, Arnhem, Barkly, Carpenta a, Tanami, and Cape York.

Ecology A herb or sub-shrubthat probablygrows annually from a woody rootstock.It is usually found in sand or sandy loam near water-coursesor other low-lying areas, but it sometimesoccurs on sandstoneplateaux or granitic pebblehillsides. It is usually found in Triodra grasslandunder Euca-lyptusor Melaleucawoodland, where it usually occurs as scatteredplants. The plants apparently flower sporadically throughout the year, perhaps only after rain.

Variability Flower sizeand colour is very variable,and this apparently correlateswith longitude, asplants with the Iargerflowers predominate in the westernpart ofthe distribution and plants with the smallest ones predominate in Queenslandand New Guinea (see Morrison 1986). 108 Kingia Vol. 1, No. 1 (198?)

Notes This speciesand ,L. ovata canbe readily distinguishedfrom all of the other rnembers of the genus by the fruits, which do not develop articles at the ends, and by the distinctive prolongation of the superior calyx lobe comparedto the inferior ones. L. filiformis has longer and thinner leavesthan L. ovata.

SpecimensExaminecl 85.

ConservationStatus Not at risk, asit is widespreadand commonthroughout tropical Northern Australia.

Lechenaultia floribunda Benth.,in Endl., Enum. PI. 70 (1837)

Distibution (Figure 15) Western Australia: Irwin, Avon, and Darling Districts.

Bencubbinand LeeuwinRegions.

Ecology A sub-shrubor smallshrub with a woodyrootstock (Baird 1977,Dodd et al. 1984).It has been recordedfrom white or grey sand or loamy sand, where it occurs in heath (Beard 1976d,1979a, Beadle 1981),thicket (Beard 1979a),or Banksia/Eucalyptus woodlandwith a heathunderstorey (Baird 1977, Milewski and Davidge 1981). It is often locally common. Flowering is from late August to mid December.

Vaiability Leaf lenglh, flower size,and flowering habit are all very variable, apparently with a morphocline running from north to south, with many of the northern populations havingplants with smallerflowers, larger leaves, and a more densely-branchedflowering habit than the southernplants (seeMorrison 1986).

Notes Closelyrelated to L. expansa,from which it can be distinguisherias above,and to L. papillata, which has papillate leaves,sepals, ovaries, and pseudocapsules.

Slecr'mers Examined 108.

Conservation Status Not at risk, as it is widespreadand common throughout the northern sandplainsof South WesternAustralia. It is exploitedfor the nurseryand seedtrades (Rye et al. 1980, Burgmanand Hopper 1982).

Common Name Free-flowering lechenaultia.

Lechenaultia formosa R.Br., Prodr. 1;581(1810)

Distribution (Figure 16) Western Australia: Irwin, Avon, Roe, Eyre, and Eucla Districts.

Nullarbor,Bencubbin, Leeuwin, and EsperanceRegions. D.A. Morrison, The phytogeography, ecology and conservation status of ,Lecheno&lho. 109

Ecology A sub-shrub with a woody rootstock that may sucker (Gardner 1944).It has been recordedfrom sand and sandy soils, often over laterite or granite, as well as from clay and gravellyclay. It occursin a variety of heaths(Beard 1972, 1973b, 1980c, Beadle 1981),scrubs, mallee (Beard 1973a, 1973b, 1976a, 1979c), and woodlands (Beard 19?9c, 1980c),where it is often locally common. Flowering apparently occurs sporadically throughout the year, but it is most prolific from early August to late October.

Variability Flower and leaf size as well as flower colour and habit vary greatly. Large-flowered prostrate plants occur throughout the geographicalrange, but the smaller-flowered more erect forms are confinedto the coastalstrip betweenAlbany and CapeLe Grande (seeMorrison 1986).

Nofes Closelyrelated to L. chlorantha,from which it can be readily distinguishedby the red and orangeflowers, andto L. linarioides,which is a much largermore tangled plant.

SpecimensExamined 302.

Conservation Status Not at risk, as it is widespreadand common throughout South Western Australia. However, Specht et al. (1974) recognisethe conservationimportance of the disjunct relict populations on the coastal sand sheets of the Great Australian Bight.

It is exploited for the nursery and seedtrades (Rye et al. 1980,Burgman and Hopper 1982).

Common Name Red lechenaultia.

Illustrations Baglinand Mullins (1969)p.91; Fairall (1970)p.182; Hodgson and Paine(197i) p.89; Blombery(1973) p.190; Holliday (1973)p.17; Grieve and Blackall(1975) pl.v; Gardner (1978)p.5.

Lechenaultia lieteromera Benth.,Fl. Austral.4:43 (1868)

Distribution (Figure 17) Western Australia: Eyre District, between Starvation Boat Harbour and West Mt Barren.

EsperanceRegion.

Ecology A sub-shrubwith a woody rootstock. It is usually found in deepwhite sand,where it occursin heath (Beard 1973a,Beadle 1981, as I-. stenosepala,Beard 1976a,as -L. slenomera),open scrub, or Banksiawoodland. It is usuallyinfrequent. Flowering is from late August to Late November.

Variability This is a relatively constant species,varying only moderately in flower colour, Ieaf density, and the degreeto which the leavesare reflexed. 110 KingiaVol. 1, No. 1 (1987) Notes _Closely related to L. brevifolia, L. juncea, and L. striata, from which it can be readily distinguishedby the non-scapigerousflowering habit and scattered,thicker, recurvei leaves.It is also sometimesconfused with,L. b oba,which has subequalwings on all of the petal lobes.

SpecimensExamined 45.

ConservationStatus Although it has a restricted distribution it is common throughout its range, and is therefore probably not at risk. It is representedin Fitzgerald River Nationa-lFark.

Common Names Claw lechenaultia(Erickson et al. 1929);Hook-leaf lechenaultia(Grieve and Blackall 7975).

Lechenaultia hirsuta F. Muell.,Fragm. 6:9 (186?)

Distribution (Figure i8) Western Australia: Irwin District. Bencubbin and Leeuwin Regions.

Ecology A herb or sub-shrubwith a woody rootstock.It occursin white or light brown sandor lateritic sand, where it is commonly found in low open heath (Beard 1976c,l9?6d, Hopkins and Hnatiuk 1981).It is often frequent,especially when regeneratingafter fire. Flowering is from early Septemberto mid December.

Variability This is a very distinctive and relatively constant species,but it doesvary somewhat within populations in Ieaf size and density.

Notes -. Closely_related to L. superba and to l. Iaricina, from which it can be readily distinguishedby the extreme hirsuteness.

SpecimensExamined 56.

ConservationStatus ,-Not at risk, as it is widespreadand common on the northern sandplainsof South Western Australia. It is exploited for the nursery trade (Rye et al. 1980,Burgman and Hopper 1982).

Common Name Hairy lechenaultia.

Illustrations Moryombe(1968) p.110; Morcombe (1970) p.98; Morcombe and Morcombe(19?0) p.81; Baker (1971)p.19; Blombery (1973)p.i91; Grieve and Blackall (19?5)pl.vi; Gardner(1978) p.5; Erickson et al. (19?9)p.98; Crafter (1988)Aust. pl. I2:b6. D.A. Morrison, The phltogeography, ecology and conservation status ol rach?uo<io. 111 Lechenaultia juncea E. Pritzel,Bot. Jahrb. Syst.35:553(1905)

Distibution (Figure 19) Western Australia: Irwin District, betweenThree Springs and Gunyidi. Bencubbin Region.

Ecology A herb or sub-shrubwith a woodyrootstock. It occursin sandor sandygravel, in heath (Beard1976d, Beadle 1981, as l.juircoides). It is usuallyinfrequent, and usually growing amongstthe branchesof other plants. It is now only recordedin fields, and along road margins.Flowering is usuallyfrom late November(sometimes earlier) to mid December.

Notes Closelyrelated to -L.brevifolia, which hasrnuch darker-blue flowers and shorterleaves, L. striata, which has much thinner more crowded leaves and larger flowers and pseudocapsules,and, L. heteromera, which does not have the scapigerousflowering habit. This specieshas also been confusedwith l. subcymosa,from which it can be distinguishedby the more upright junciform habit and the much Ionger sepals.

SpecimensExamined 8.

Conservation Status 2V, as it is geographicallylocalised (Rye 1981),and currently known only from a few roadsideverges and relativelyundisturbed fields in farmland. This speciesis very poorly collected(Marchant and Keighery19?9), and it is not known from any conservation reservesor knownto be in cultivation(cl Rye 1981).This specieswould be classifiedas 2E, except that there are a couple of good populations in protected areasin farmland that will hopefullybecome conservation reserves in the near future (cf. Beard 1976d).

CommctnName Reedy Iechenaultia.

Lechenaultia laricina Lindley, SketchVeg. Swan R. 27 (1839) Distribution (Figure 20) Western Australia: Avon District, between Meckering and Clackline, and between Kukerin and Moulyinning. Bencubbin and Leeuwin Regions. Ecology A small shrub with a woody rootstockthat may sucker.In the past,this specieshas been recordedfrom sandor occasionallygravelly loam, usuallyin woodland.It is now known only from disturbedareas in farmland. Floweringis from late Octoberto late December. Variability This speciesis relatively constant morphologically,but it does vary somewhat in flower colour,with some populationsbeing less scarletthan others.As well, some variation exists between the two disjunct groups of populations of this species,the southern group being much more prostrate plants, with greyish bark and somewhat smallerllowers.

Notes Closelyrelated to L. ir'rsuta, from which it can be readily distinguishedby the glabrous habit, and to L. superba,which has a more upright habit, longer leaves,and yellowish flowers with a broader tube. However, this speciesis apparently persistently and 112 Kingia\tol. 1, No. 1 (198?) irequently confusedwith other species(see Morrison 1986),usually resulting in the identification of other species(often -L.formosa) as L. laicina. This has, in the past, suggestedthat -L.laricr'na occurs over a much larger areathan it really does(cl the map in Rye et al. 1980) and that it has been collected much more frequently. It was apparentlythis confusionthat causedMarchant and Keighery(19?9) and subsequent workersto overlookthis speciesas an endangeredtaxon.

Spec.imensExamined

ConservationStatus 2E. This specieswas once apparently common in the area between Meenaar. Meckering,and Northam (Gardner1978), but in this areait is nowknown from onlytwo populations in farmland. Both of these populations are currently being protected,but this relies entirely on the goodwill of the peopleconcerned, and one of the populations appearsto havecontracted recently. The only other reliablecollection localities are near Kukerin, where it has not been found recently.It thus seemsthat this speciesis under immediate threat, especially as no populations have been reported in conservation areas.Gazettal as protected flora is therefore essential.The speciesis reported to be exploited for the nursery trade (Rye et al. 1980,Burgman and Hopper 1982),but the collection locality given by Burgman and Hopper (1982) suggeststhat many of these reports may be mis-identifications (seeabove). Fortunately, it is apparently in wide cultivation, as this would seemto be critical for the continued survival of the species. Fairall (1970) discussesits cultivation requirements.

Common Name Scarlet lechenaultia.

Illustrations Fairall (1970)p.183; Grieve and Blackall(1975) pl.v; Gardner(1978) p.5.

Lechenaultia linarioides DC., Prodr. 7:519(1839)

Drstributron(Figure21) Western Australia: Irwin and Darling Districts, and CarnarvonDistrict on the Shark Bay peninsulas.Murchison, Bencubbin, and Leeuwin Resions.

Ecology A small shrub with a woody rootstock that may sucker. It occurs in deep white or yellow sand, where it is found in open-heath(Beard 19?6c,19?6d, 19?9a, 19?9b, Burbidgeand George 1978, Bell et al. 1979,Bridgewater and Zarnmit 19?9, George et al. 1979,Beadle 1981), scrub (Beard 1976c, 1976d, Beard and Burns, 1976), or occasionally woodland (Bell et al. 1979). Ii is often locally common. Plants apparently flower sporadicallythroughout the year, but flowering is mainly from early August to early December.

Vaiability Stature,leaf and flower size,and flower colour are all very variable.The isolatedgroup of plants near Shark Bay are taller and more denselyshrubby with thicker branches than the southernones, which tend to be moresprawling. Flower size and colour varies considerablybetween populations, and the flowers themselvesalso apparentlv become redderwith aee. D.A. Morrison, The ph).togeography, ecology and conservation status of -Lecheno./lfid.

Nofes Similar to l. form osa and.L. chloranl.ha,from which it can be readily distinguishedby the thicker stemswith downcurvedbranchtips and more scatteredleaves, as well as by the larger flowers.

Specimers Examined 135.

ConsewationStatus Not at risk, as it is widespreadand common throughout the northern sandplainsof South WesternAustralia. It is exploitedfor the nursery and seedtrades (Rye et al. 1980, Burgmanand Hopper 1982).

Common Name Yellowlechenaultia.

Illustrations Morcombeand Morcombe(1970) p.80 (as L. Iinaroides);Hodgson and Paine(1971) p.91;Blombery (1973) p.191; Grieve and Blackall (1975)pl.v; Gardner(1978) p.131.

Lechenaultia longiloba F. Muell.,Fragm. 6:10 (1867)

Distibution (Figure 22) WesternAustralia: Irwin District, betweenMullewa and Dongara.Bencubbin Region.

Ecology A sub-shrubwith a woody rootstock that may sucker. It has been recordedin deep white or greyearthy sand,in openheath (Beard 1976d).It wasapparently once common in Banksia heath (especiallyif regeneratingafter fire), but it is now more commonly found alongdisturbed roadsides. Floweringis usuallyfrom late July to earlyOctober, but it sometimescontinues sporadicallyuntil early December.

Variability This is a relatively constantspecies, but there is somevariability betweenpopulations in flower colour (varying from red to yellowish) and flower size

Nofes Closelyrelated to -L..hirsuta, from which it can be easilydistinguished by the glabrous habit, and to L. macrantha,which has longer leaves(which are characteristicallyheld to one side of the branch), larger wings, and a shorter broader corolla tube.

SpecimensExamined 30.

ConservationStatus 2VC, as it is geographicallylocalised (Rye 1981),and is now known from onlv a few populations along roadsidesin larmland (see Leigh et al. 1984 for a more detailed "A discussion),and from one population in an Class"conservation reserve (Beard and Burns 1976).Gazettal as protectedflora would seemto be appropriate,and it may be necessaryto managethe roadsideverges specifically to maintain this species(cf. Scott 1981).Hartley and Leigh (1979)report it to havebeen subject to heavycommercial exploitation in the wild, but this has not been confirmed (cf. Rye et al. 1980).It is not known to be in wide cultivation (Leigh et al. 1984),and horticultural propagationwould seemto be essentialto ensurethe continued survival of this species. lt4 Kingia Vol. 1, No. 1 (1987)

Common Name Irwin Iechenaultia.

Illustrations Baglinand Mullins (1969)p.91; Grieve and Blackall (1975) pl.v; Erickson et al. (1979) p.10?;Crafter (1983)Ausr. Pl. 12:57;Leigh et al. (1984)between pp.192-3.

Lechenaultia lutescensD.A. Morrison & R.C. Carolin,Brunonia 9:15 (198?)

Distributi on (Fieure 26) Northern Territory: Central Australia North, and Central Australia South Regions; WesternAustralia: Mueller, Giles,Carnegie, and Helms Districts. Gibson,Tanami, MacDonnell, and Victoria Desert Regions.

Ecology A herb or sub-shrubwith a woody rootstock. It has beenrecorded from deepred sand dunes,sandy loam plains, or around the gravellyedges oflateritic breakaways.It occurs among mallees,desert oak, open Triodia grassland,or Sprnr'Iexopen scrub,where it is often locally common. Flowering apparently occurs sporadicallythroughout the year, perhaps only after recent rain.

Variability This speciesvaries somewhatbetween populations in leaf size,and flower size and colour, but this does not seemto be significant.

Noles Closely related lo L. heteromera,from which it can be readily distinguishedby the yellow flowers and shorter calyx lobes.It has also been confusedwith L. striata, wlniclr. has a distinctly scapigerousflowering habit and ridged rather than groovedarticles.

Spec.imensExamined

ConservationStatus Not at risk, as it is widespread and common throughout arid western Central Australia.

Lechenaultia macrantha K. Krause,Pflanzenr., IV. 54:100(1912)

Di stributi on (Figure 23) Western Australia: Irwin and Avon Districts; and one isolated collection in Irwin District, nearNerren Nerren station, and onein Austin District, nearBoolardy station.

Murchisonand BencubbinRegions.

Ecology A herb or sub-shrubthat growsannually from a woodyrootstock (Ericksonet al. 1979). It occursin yellow sandor, more usually,red gravellysoil, in open areasnear heathland (Beard 1976e),or along road margins or other disturbed areas.It is usually locally common. Flowering is from mid August to late October.

Variability Leaf size,and flower colour and sizevary considerablybetween plants. Much of this variability appearsto be latitudinal, with the more northern inland plants having the larger organs. D.A. Morrison, The ph1'togeography, ecology and conseNation status of ,€c,henoultio. Notes Closely related to L. longiloba, from which it can be readily distinguishedby the wreath-like flowering habit, longer leavesand sepals,and larger petals with broader wings.

SpecimensExamined 40.

ConservationStatus Not at risk, as it is relatively widespreadand common on the inland red gravelsof northern South Western Australia. Both Specht et al. (1974)and Hartley and Leigh (1979)consider the speciesto have sufferedmarked depletionsin distribution, but as it apparentlyresponds well to disturbancethis is probably not asbig a problem aswas first thought. Hartley and Leigh (1979) record the speciesas occuuing in conservation reserves.They also report it to have been subjectto heavy commercialexploitation in the wild, but this has not been confirmed (cf. Rye et al. 1980).

Common Name Wreath lechenaultia.

Illustrations Morcombe(i968) p.31;Newbey (1968) p.125; Baker (1971)p.i9; Blombery(1973) p.191;Grieve and Blackall(1975) pl.v; Gardner(1978) pp.131-2; Erickson et al. (1979) p.118;Crafter (1983)Aust. PL.72:56.

Lechenaultia ovata D.A. Morrison,Telopea 3: in press

Distribution (Figure 26) Northern Territory: Darwin and Gulf Region. Arnhem Region

Ecology A herb that probably grows annually from a woody rootstock. It has been collected from short sedgelandin a sandy depressionon a sandstoneplateau. The specimenwas floweringin February.

Nofes This speciescan be readily distinguishedfrom all ofthe other membersofthis genus by the ovateleaves.

Specr'mensExamined 1.

ConservationStatus 1K, as it is known only from the type collecion(see Morrison 1987).The population is apparently fairly large, but the locality is just outside Kakadu National Park. Intensive searchesneed to be made in the area to correctly ascertainthe status of this specres.

Lechenaultia papillata D.A. Morrison,Brunonia 9:12 (i987)

Distribution (Fieure 24) Western Australia: Roe District. EsperanceRegion. 116 KingiaVot. 1, No. 1 (1987) Ecology A sub-shrub or small shrub with a woody rootstock. It has been recorded from yellow or white sand, loamy sand, and gravelly loam. It usually occurs in heath (Beard 1969, Beadle 1981,as L, expansa),lowopen scrub, or eucalypt scrub mallee,where it may be locally common. Flowering is usually from Iate October to late November.

Vaiabilitv Flower size is somewhat variable. but the snecies nevertheless remains distinctive.

Nofes Closely related to L. floribunda, from which it can be readily distinguished by the shorter more crowded leaves,the papillate leaves,sepals, ovaries and pseudocapsules, and the much more hairy floral tube.

Specimens Examined 74.

Conservation Status Although poorly collected, this species occurs in an area that is sparsely collected in general. Given the large area over which collections have been made, the species is probably not at risk. It is represented in Frank Hann National Park.

Lechenaultia pulvinaris C. Gardner,J. Roy. Soc.W. Austral.47:63(1964)

Distribution (Fieure 25) Western Australia: Avon District, betweenCorrigin and Wagin, and near Beverley. Bencubbinand LeeuwinRegions.

Ecology A sub-shrubwith a woody rootstock. It occursin deepwhite sand on plains or gentle slopes,sometimes near low-lyingseepage areas. It only growsin openpatches in Iow scrub,becoming absent as the vegetationbecomes more densewith age.It is often locally common.Flowering is from mid Octoberto earlyDecember.

Notes Closely related to L. expansa,frorn which it can be easily discernedby the dense pulvinate habit, Iike that of l. tubiflora.It canbe readily distinguishedftom L. tubiflora by the hispid foliage and calyx lobes.

Spec.imensExamined

ConservationStatus 3VC, as,although it is not geographicallylocalised (but seeRye et al. 1980),it occurs only in open patchesin the vegetation.It is known from only a few apparently disjunct areas(see Leigh et al. 1984),and, although only rediscoveredand describedin the early 1960s,it was in fact collectedlate last century. It has been recently reported to occur "A around the edgesof a number of small Clais" conservationreserves, but it appears to becomeabsent from the vegetationas the vegetationbecomes more densewith age. Much more needsto be known about its biologybefore it can be decidedhow widespread the speciesis and whether it is adequatelyconserved (and thereforemore appropriately categorisedas 3RC).Hartley and Leigh (1979)report it to havebeen subject to heavy commercial exploitation in the wild, but the speciesis now protected as gazettedrare D.A. Monison,The phytogeography,ecology and conservation status of Lechenaultia ll7 flora (Patrick and Hopper 1982). However, it should be introduced into cultivation to ensure the long term survival ofthe species,and Newbey (1968) discussesits cultivation requrremenls.

Common Name Cushion Iechenaultia.

Illustrations Anon. (1981) between pp.12 3; Rye and Hopper (1981) p.145; Leigh et al (1984) between pp.192-3.

Lechenaultia stenosepala E. Pritzel, Bot. Jahrb. Syst.35:552 (1905)

Distibution (Fieure 28) Western Australia: Irwin and Darling Districts. Bencubbin and Leeuwin Regions.

Ecology A herb or sub-shrub with a woody rootstock that may sucker. It is found in yellow or white sand or sandy gravel, where it is usually recorded from low open heath (Beard 1976d, Hopkins and Hnatiuk 1981), otten in damp depressionsor valley floors. It is often locally common, especially as regrowth after fire. Flowering is from early October to mid December.

Vaiability Leaf and flower size vary considerably, and this apparently correlates with latitude, lhe largest leaves and flowers only occurring on plants in the northern part of the distribution, and the smallest Ieaves and flowers only occurring in the southern part (see Morrison 1986).

Nofes Closely related to L. Iloribunda, from which it can be readily distinguished by the longer sepals, and to ,. b oba, which has flowers of a much deeper blue and with much larger corolla wings.

Specimens Examined 51.

Conservation Status Not at risk, as it is widespread and common on the nofthern sandplains of South Western Australia.

Common Name Narrow sepaledlechenaultia.

Lechenaultia striata F. Muell., Fragm. 8:245 (1874)

Distribution (Figure 27) Northern Territory: Central Australia South Region; South Australia: North-western Region; Western Australia: Giles, Canegie, Helms, and Austin Districts. Gibson, MacDonnell, Leonora, and Victoria Desert Regions. 118 KingiaVol. 1, No. (198?) Ecology A herb or sub-shrub with a woody rootstock that may sucker (Maconochie 1982). It is found on deep red sand dunes, where it is characteristic of the mid and upper slopes (Buckley 1981a). It occurs in open ?riodia grassland or Spurifex open scrub, where it is often locally common, especially after recent rain. Plants appear to have only a moderate drought tolerance (Buckley 1982). Flowering apparently occurs sporadically throughout the year (perhaps only after recent rain), but it is usually from late August to mid November.

Variability This species varies considerably between plants in leaf, flower and pseudocapsule size, and sornewhat less in ilower colour. None of this variation appears to be correlated.

Noles Closely related to -L.brevifolia, from which it can be readily distinguished by the longer leaves, and to L. lutescens, which does not have the scape-like flowering stems.

Specimens Examined 34.

ConservationStatus Not at risk, as it is widespread and common throughout arid western Central Australia. Specht et aI. (1974) report it to be rare in the Northern Territory, but this doesnot appear to be the case.

Common Name Striate-stemmedlechenaultia.

Illustrations Jessop& Toelken (1986)p.1407.

Lechenaultia subcymosaC. Gardner& A.S.George, J. Roy.Soc. W. Austral.46:134 (1963)

Distribution (Fieure 29) Western Australia: Carnarvon District, on North West Cape and on the Shark Bay islands.Murchison Region.

Ecology A herb or sub-shrubwith a woody rootstock. It has been recordedfrom sand or loam over limestone (Burbidge and George1978) or in red sand dunes,where it occurswith scattered low shrubs. It is often found in very dense stands regeneratingafter fire. Flowering apparently occurs sporadically throughout the year (perhaps only after a recent fire), but it is usually from late July to early October.

Notes Closelyrelated to L. divaricata, from which it can be readily discernedby the larger leaves,smaller flowers, and non-moniliformpseudocapsule.

Specr'mensExamined 15. D.A. Morrison, The phytogeography, ecology and conservation status of tectenaultia. 119

ConservationStatus Although poorly collected,this speciesoccurs in an area that is sparselycollected in general.As it is apparently common where it doesoccur, the speciesis probably not at risk.

Common Name Wide-branching Iechenaultia.

Lechenaultia superba F. Muell.,Fragm. 6:10 (1867)

Distribution (Figure 30) WesternAustralia: Eyre District, atthe easternend ofthe Barrens.Esperance region.

Ecology A small shrub with a woody rootstock. It only occurs in quartzite soils on rocky hillsides or in open gullies,where it is found in open patchesin thick scrub.Plants are sometimes locally common. Flowering is usually from September to October, but flowering plants have been recordedat other times.

Notes Closelyrelated to tr. .hr'rsuta,from which it can be readily distinguishedby the glabrous habit, and to L. laricina, which has a low spreadinghabit.

SpecimensExamined 19.

ConservationStatus 2RC, as it is geographicallylocalised (Rye 1981)and restricted to a specific habitat. Aknost the entire known distribution is within Fitzgerald River National Park, but the number of known plants is very small (Rye 1981).It is known to be in cultivation (Rye 1981,Rye and Hopper1981), and Fairall (1970)discusses its cultivationrequirements. It is also protected as gazettedrare flora (Patrick and Hopper 1982).

Common Name Barrens lechenaultia.

I llustrations Ericksonet al. (1979)p.92; Anon. (1981)between pp.12-3; Rye and Hopper (1981) p.747.

Lechenaultia tubiflora R.Br., Prodr. 1:581(1810)

Distribution (Fierrre31) WesternAustralia: Irwin, Avon, Darling, and Eyre Districts. Bencubbin,Leeuwin, and EsperanceRegions.

Ecology A sub-shrub with a woody rootstock that may sucker (Gardner 1944).It has been recordedfrom deep yellow or white sand, where it usually occurs in open patches in mixed heath (Beard 1973a, 1979c, 1980a, Beadle 1981), or Eucal;rytus or Banksia woodland.It is often locally common, especiallyon bare ground. Flowering appearsto occur sporadicallythroughout spring and summer,but it is mainly from late September to earlyDecember. I

120 Xingia Vol. r, No. I (1987)

Variability Apart from flower colour and leaf size,which can vary considerablywithin a single population, this speciesis apparently differentiated into coastaland inland forms (see Morrison 1986).

Notes Closely related to L. acutiloba, from which it can be readily distinguishedby the smaller flowers and winged corolla lobes.The prostrate habit is similar to that of -L. pulvinaris, which has hispid leavesand calyx lobes.

SpecimensExamined 153.

ConservationStatus Not at risk, as it is widespreadand common throughout South Western Australia.

Common Name Heath lechenaultia.

Illustrations Beard(1970) pl.xxxi; Morcombe (1970) p.110; Walton (1970)Ausl. P1.5:248;Wrigley (1970)Aust.Pl.5:241; Blombery (1973) p.191; Holliday (1973) p.16; Grieve and Blackall (1975)pl.iv; Ericksonet at. (1979)p.84; Wrigley and Fagg(1979) pp.128-9; Elliot and Jones(1982) p.185. D.A. Morrison, The phytogeography, ecology and conservation status of lechenaultto.

Frgure L Distributionol L. aculiloba Figure 9. Distributionol L. biloba

Flgu re 10. Disiribution ol L. brevilolia Frgure 1 1. Disrributionof L. chloranlha

,4609-9 122 KingiaVol. I, No. t (1987)

Flgure 12. O'Btribution ol L. divaric.l.

Frqur6 !3. Dlslribulionol L. lililormll D.A. Mordson, The phltogeography, ecology and conseNation sLat:usof Lechenaultia. 123

Figure 14. Distributionol L. expansa Figure 15. Distribulionof L. flo.ibunda

Figure 16. Dislribulion ot L. lormosa Flgure 17. Distribulionof L. heteromera KingiaVol. I, No. t (198?)

Figur6 t8. Distributionol L. hirsula Figure 19. Distribulionol L. juncea

Figure20. Distributionof L. laricina Figure21. Drslr'bulionol L. Inarioides D.A. Mordson, The phytogeography,ecology and conservatiot statvs of LechenauLtia.

Figure 22. Distribulionol L. longiloba Figure 23. Distributionof L. macrantha

Figure 24. Distribulionot L. papillata Figure25. Dislributionot L. putvinaris Kingia Vol. I, No. (1987)

Fi9ure 26, Dislribulion ot L. lulescens and * L. ovata

FiCure27. Oislributionol !. striata D.A. Monison, The phltogeography,ecology and conservat\onstalDl ol Lechend.ultia. 127

Figure 28. Distributionol L. sienosepala Figure29. Dislributionof L. subcymosa

Figure 30. 0isrribuiion ol L superba Figure 31. Distributionol L. tubiflora t2fl Kingia Vol. t, No. 1 (r98?)

Appendix 2. Zecrrenaultia species recorded (or considered likely) to occur in O.5'lat. x 0.5.long. grid cells of South Western Australia and adiacent areas, Occurrences not represented by a her- barium specimen are in brackets. The grid cell numbers and names are those of the Division of National Mappi g's coding system for the 1:1O0,0O0 topographic maps.

1446QUOIN (L. Iinarioides) L. subcymosa 1547DORRE L. subcymosa 1546DENHAM L. linarioides (L. subcymosa) 1545EDEL L. linarioides L. subcymosa 1646SHARK BAY L. linarioides 1645PERON L. linarioides 1743COOLCURDA L. hilsuta (L.linarioides) 1?42KALBARRI L. chlorantha L. floribunda L. hirsuta L. linarioides 1741HUTT L. floribunda L. linarioides 1844WANNOO L. floribdnda 1843NERREN NERREN L. Iloribunda L. hirsuta L. linarioides L. macnntha 1842AJANA L. floribunda L. hircuta L. linarioides L. macrantha 1841NORTHAMPTON L. Iloribunda L. hilsuta L. Iinarioides r84OGERALDTON L. lloribunda (L. hirsuta) L. linarioides L. longiloba 1839DONGARA L- floibunda (L. linarioidesJ 1838BEAGLE ISLANDS (L. biloba) (L. floribunda) L. Iinarioides 1837GREEN HEAD (L. bilobal (.L. floribunda) L. linarioides 1830CLAIRAULT (L. biloba) (L. expansa) 1829TOOKER (L. biloba\ (L. expansa\ 1942COOLCALALAYA (L. macnntha) 1941MUNGO L. floribunda L. macantha 1940INDARRA L. floribunda (L. hirsuta) L. Iinarioides L. Iongiloba L. mactuntha 1939MINGENEW L. floribunda L. hirsuta L. Iinarioides L. longiloba L. macrantha 1938ARROWSMITH L. biloba L. floribunda L. hirsuta L. linarioides L. stenosepala 1937HILL RIVER L. biloba L. floribunda L. hirsuta L. linafioides L. stenosepala D.A. Morrison, The phytogeography,ecology and conservationstaLrs of LechenauLtia 129

1936WEDGE ISLAND L. biloba L. floribunda L. lina oides L. stenosepala 1935LEDGE POINT (L. bilobal (L. floribunda) L. l;narioides (.L. stenosepala) 1930BUSSELTON L. biloba L. expansa 1929LEEUWIN L. biloba L. expansa 2O4ITALLERTNC L. lloribunda L. mactuntha 2O4OMULLEWA (L. floribunda) L. macrantha 2039YANDANOOKA L. floribunda L. hi$uta L. linarioides (L. macrantha) 2038 CARNAMAH L. biloba L. floibunda L. formosa (L. hirsuta) L. juncea L. linarioides L. stenosepala 2037BADCINGARRA L. biloba L. floribunda L. formosa L. hi$uta (L. juncea) L.linarioicles L. stenosepala L. tubiflora 2036DANDARAGAN L. biloba L. floribunda L. linarioides L. stenosepaia L. tubillon 2035GINGIN L. biloba L. floribunda L. linarioides L. stenosepala L. tubiflora 2034PERTH L. biloba L. expansa L. florib nda L. linarioides L. tubillora 2033FREMANTLE L. biloba L. expansa L. floribunda L. Iinarioides 2032PINJARRA L. biloba L. expansa 2031 BUNBURY L. biloba L. expansa 2O3ODONNYBROOK L. biloba L. expansa 2029 DONNELLY L. biloba L. expansa 2028MEERUP (L. expansa) 2141WURARGA L. floribunda 2140 MELLENBYE L, macrantha 2139PERENJORI L. macnntha 2138CARON L. biloba L. juncea L. linarioides L. macrantha 2137 WATHEROO lL. biloba) L. floribunda L. juncea L. lina oides L. stenosepala 2136MOORA L. biloba L. floribinda (L. formosa) L. Iinarioides (L. stenosepala) 2135 CHITTERING L. biloba L. floribunda L. formosa L. linarioides L. stenosepala 2134 WOOROLOO L. biloba L. expansa L. floribunda L. formosa L. linarioides L. tubillora 2133JARRAHDALE L. biloba L. expansa L. floribunda L. linarioides 2132DWELLINGUP L. biloba L. expansa 130 KingiaVol. r, No. I (1987) 213ICOLLIE L. b oba L, expansa 2130BRIDGETOWN L. biloba L. expansa 2T29MANJIMUP L. biloba L. expansa 2128NORTHCLIFFE L. expansa 2244BOOLARDY L. macrantha 2238MONGERS L. macrantha 2237DALWALLINU L. biloba 2236WONGAN L. biloba L. floribunda 2235GOOMALLING L. biloba L. floribunda 2234NORTHAM L. biloba L. floribunda L. formosa L.laicina L. tubiflora 2233BE\ERLEY L. biloba (L. floribunda) L. formosa L. pulvinaris 2232CROSSMAN L. biloba 2231DARKAN L. biloba 2230DINNINUP L. biloba 2229TONEBRIDGE L. hiloba L. expansa 2228DEEP RIVER L. expansa 2227RAME HEAD L. expansa 2338MOUNT GIBSON L. macrantha 2336KOORDA L. biloba 2335DOWERIN L. biloba L. floribunda 2334CUNDERDIN (L. biloba) L. floribunda L. formosa L. tubiflora 2333BROOKTON L. biloba L. lloribunda (L. fomosal L. tubillon 2332NARROGIN L. biloba L. fomosa L. tubiflotu 2331WAGIN L. biloba L. lormosa L. tub;Ilora 2330KOJONUP L. biloba L. formosa L.tDbiflora 2329FRANKLAND (L. tubiflora) 2328DENMARK L. expansa L. tubiflora 2327PARRY INLET L. expansa 2434KELLERBERRIN L. biloba L. formosa L. tubiflora D.A. Morrison, The phytogeo$aphy, ecologyand conservationstatus of Lechenaultla. 131

2433CORRIGIN (L. biloba) (L. formosa) L. tubiflora 2432YEALERING (L. biloba) (L. formosa\ L. pulvinaris L. tubillora 2431DUMBLEYUNG L. biloba (L. formosa) L. Iaricina L. pulvinaris (L. tubiflora) 2430 KATANNING L. biloba L. formosa (L. tubillora) 2429TAMBELLUP L. formosa L. tubitlon 2428 MOUNT BARKER L. expansa L. formosa L. tubillora 242? ALBANY L. expansa L. formosa L. tubillora 2534BRUCE ROCK L. biloba (L. formoga) 2533NAREMBEEN L. biloba 2532 KULIN L. biloba L. formosa 2531KUKERIN L. biloba (L. formosa) L. tubiflora 2530 NYABING L. acutiloba L. formosa (L. tubifiora\ 2529 BORDEN L. formosa L. tubiflora 2528 MANYPEAKS L. expansa L. formosa L. tubiflora 2527 BREAKSEA L. expansa L. formosa L. tubillon 2634MUNTADGIN (L. formosa) 2633 HYDEN L, formosa 2632 PEDERAH (L. fomosa\ 263I BURNGUP L. biloba L. fomosa 2630JERRAMUNGUP L. acutiloba L. fomosa (L. tubillora) 2629 PALLINUP L. formosa L. tubillora 2628 CHEYNE L. formosa L. tubiflora 2735 SOUTHERN CROSS L. brevifolia 2?34HOLLETON (L. brevifolia) L. formosa 2733o',CoNNOR L. brevifolia (L. formosa) 2?32HURLSTONE (L. formosa) 2?31NEWDEGATE L. biloba L. formosa L. papillata 2730JACUP L. acutiloba L. formosa L. tubiflora 2729BREMER L. formosa L. heteromera L. tub;flora 2728CAPE KNOB (L. fomosa\ (L. tubiflora\ 132 Kingia Vol. 1, No. I (1987)

2835YELLOWDINE L. brevifolia 283,i CHERITONS FIND (L. brevifaiia) 2833 HOLLAND (L. brevifolia) L. formosa L. papillata 2832 IRONCAP L. brevifolia (L. formosa) (L. papillata) 2831KING L. bretilolia L. lormosa L. heferompra L. papiltata 2830 COCANARUP L. acutiloba L. formosa L. hetetumera L. superba lL. tubifjora) 2829HOOD POINT L. formosa L. hetetumera L. superba L. tubiflora 2935 BOORABBIN (L. brevifalia) 2934 LAKE PERCY (L. brevifolial 2933ROUNDTOP \L. brevifolia) 2932HOPE L. brevifolia L. formosa L. papillata 2931MOOLYALL (L. brevifolia) L. formasa L. papi ata 2930 RAVENSTHORPE L. formosa L. heteromera L. superba L. tubiflora 3035 WOOLGANGIE L. brevifolia 3034DIAMOND ROCK (L. brcvifolia) 3033JOHNSTON \L. brevifolia) 3032TAY L. brevifolia L. papillata 3031NORTHOVER (L. brcvifolia) (L. formosa) L. papillata 3O3OOLDFIELD L. lbrmosa L. heteromen L. tubiflora 3132PEAK CHARLES (L. brevifolia) (L. papillata) 3131LORT (L. brevifolia) L. fomosa \L. papillata) 3130STOKES INLET L. formosa L. tubiflon 3232DUNDAS (L. brevifolial L. formosa (L. papillata) 3231SCADDAN L. brevifolia L. lbrmosa (L. papi ata) 3230 ESPERANCE L. formosa L. tubillon 3332COWALINYA (L. brcvifolia) (L. papi ata) 333i BURDETT (L. brcvifolia) L. formosa \L. papillata) 3330 MERIVALE L. formosa L. tubillora 3432MOUNT ANDREW (.L. brevilblia) (L. papillata) 3431BEAUMONT L. brevifolia L. papillata D.,{. Nlorrison,lfhe phllogeographt.ecol;g1' and conservationslatus of aechena&laia. 133

3.130HOWICK L. formosa L. tubiflon ijI]3I BURANIINYA L. papillata 3530SANDY BIGHT [,. formosa L. tDbitlow 3630 \'l,ALCOLM L. lbrn)ota L. tobillon

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