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DOI: 10.18195/issn.0313-122x.61.2000.175-199 Records of the Western Allstralian Mllsellm Supplement No. 61' 175-199 (2000).

The flora and vegetation of the seasonal and perennial wetlands of the southern Carnarvon Basin, Western Australia

Neil Gibson, G.}. Keighery and M.N. Lyons Department of Conservation and Land Management, PO Box 51, Wanneroo, Western Australia 6065, Australia

Abstract -A total flora of 265 taxa was recorded from 58 wetland areas in the southern Carnarvon Basin. The flora was dominated by arid species but also included 34 temperate taxa at the northern end of their range and 18 tropical taxa at the southern end of their range. Five species (DicllOpogon tyleri, Calandrinia sp. Coolcalalaya (GJK and NG 698), Myriocephallls gascoynensis ms, Lythmm sp. Towrana (RJC 2183), Psammagrostis wiseana) are endemic to the study area. The Calandrinia sp. appears not to have been collected previously. A collection of Rllmex crystallil1llS was the first recorded in Western Australia since 1885. Classification of the wetlands was undertaken at two different scales; at both scales heterogeneity was high. The major division in the data sets was between the saline and non-saline wetlands. Percentage of single species occurrences was high in both datasets, ranging from 46--55%; this component of the flora adds significantly to plant biodiversity but is not a predictable component of vegetation communities. In the seven-group classification no floristic groups were restricted to particular habitat types but at the finer scale sampling there is a stronger correlation between floristic community type and habitat type. At this scale the major habitat types (e.g. river pools) encompassed a range of floristic groupings. The patterns in low species richness and very patchy species distribution found in this study are similar to those reported for wetlands in the higher rainfall areas of the south west but the underlying causal factors determining these patterns remain unclear.

INTRODUCTION Other major wetland habitats found in the study The flora and vegetation of the wetlands of the area include those associated with permanent and southern Carnarvon Basin (Anonymous, 1975) have semi permanent river pools, and freshwater and been little studied (Figure 1). In the most recent and saline springs, wetlands of seasonally inundated comprehensive review of nationally significant claypans, and the widespread saline samphire flats wetlands only two non-marine wetlands are listed and birridas (gypsum pans) (Battye, 1915; Payne et from the study area (ANCA, 1996). These two large al., 1987). The flora and vegetation of Rocky Pool on wetland systems highlight the diverse nature of the Gascoyne River (Kenneally, 1978) and the wetlands in this area. Lake MacLeod covers some occurrence of Psilotum nudum in the lower 150000 ha formed on calcareous marine deposits and Murchison River (Smith and Butler, 1961) are lies 30 km north west from Carnarvon. In part, it is fed among the few accounts of wetland vegetation that by ocean water through 18 km of coastal limestone have been published for this area. which upwells in sinkholes in the central western Numerous classification systems for wetlands section of the wetland, but precipitation can also make have been published both at the regional (Paijamans a significant contribution to water supply in wetter et al., 1985; Lane and McComb, 1988; Jacobs and years (Scott, 1962; ANCA, 1996). The mangrove Brock, 1993; ANCA, 1996) and local scales community of Lake MacLeod is one of only two (Semeniuk, 1987), using geomorphic and physical inland occurrences (ANCA, 1996). In contrast the features, but none of these systems clearly McNeill claypan system is located on outwash alluvial differentiate the major geomorphic types recognised deposits near the mouth of the Gascoyne River. The in the study area; as a consequence the terminology soils have been described by Bettenay et al. (1971) as of Payne et al. (1987) is adopted here. saline-alkaline with high levels of exchangeable The aim of this paper is to describe the flora and sodium and a high content of soluble salts. These flats vegetation of the major seasonal and permanent are inundated with fresh water when the Gascoyne wetland types of the southern Carnarvon Basin and River floods. Both wetlands are important for determine how floristic units correlate with major migratory birds (ANCA, 1996). habitat types. 176 N. Gibson, G.]. Keighery, M.N. Lyons

113° 115"

24

I. ~ CB67a I/ ~ CB68 ( /~ ~ CB67b CB75c ( CB73 ~ ~ CB70b CB75a~ ~ I ~ \ I J CB54 CB75b (CB54a ~~ CB56/56a .. ~ CB62b CB44 ~ 'CB58b ~ CB62a1c Bemier I ~ /.C~58c CB58d o.\CB42 ~ CB62 qs. " co.): ~-.- Qe " .. CB38aAA CB43 DorreI

CB35a /a--CB30.. ---\ CB34-..... '.../\ ~ / "~j \ Dirk , Wooramel cB93g.... . / CB93f 1?l\?er·· "';.-j.iCB93d Hartog I CB93cl CB25 ~

CB27b ~ ·.... CB27c

CARNARVON BIOREGION ~ I N 27" I 27° 0 25 50 = Kilometres GERALDTON SANDPLAINS BIOREGION

!\

Figure 1 Location of wetland areas sampled and showing boundaries of major phytogeographic regions (Thackway and Cresswell, 1996). Wetland floraandvegetation 177

METHODS Table1 Wetland taxaendemic tothesouthern Fifty-eightwetlandareasweresampledacrossthe CarnarvonBasin. studyarea(Figure1andAppendix 1) inAugust Dlchopogontyleri andSeptember1995.Nocoastalsalinewetlandsor Calandriniasp.Coolcalalaya(GJKandNG698) mangroves were includedinthepresentsurvey. Myriocephlllilsgascoynensisms Severallocationswithinsomeofthelargerwetlands Lythrwnsp.Towrana(RJC2183) (e.g.LakeMacLeod) were sampled.Thesiteswere Psammagrostiswiseana thesameasthoseused fortheinvertebrate samplingbyHalseetal.(2000)withoneexception, genericlevelCalandrinia(8taxa),Eragrostis(8taxa), andtheadditionofasiteonMcNeillclaypan.As Cyperus (7taxa),Atriplex(5taxa)andGoodenia (5 clearvegetationzonationwas apparentaround taxa)werethemostspeciesrich. manyofthewetlandsaseriesof1m x1m quadrats Thirty-oneintroducedspecieswererecordedfrom was usedtosampleeachvegetationtype,running thewetlands. Thewetland vegetationwas heavily alongatransectatrightanglestotheshoreline.Lists dIsturbedaroundsomeofthespringsandriver ~ compiledofalltaxa(species,subspeciesand pools with eitherthecompleteremovalof vanetIes)foundinthesequadrats.Taxaseenateach w1derstorey, orwith taxasuchasCenchrusciliaris wetland butnotrecordedinquadratswere also andAsphodelusfistulosusbecomingdominant.The noted.Siteswereclassifiedaccordingtosimilarities vegetationofthesalinewetlandsandthewetlands inspeciescompositionusingtheCzekanowski oftheephemeralclaypanswasgenerallymuchless coefficientand'unweightedpair-groupmean impacted. average'fusionmethod (UPGMA;Sneathand A totalofsevennewpopulationsoffourtaxa Soka!,1973),whilespecieswereclassifiedusingthe (Bergiaauriculata,Goodenianeogoodenia,Rumex 'twostep'method ofAustin andBelbin(1982)and crystallinus,Wurmbeamurchisoniana) listedon theUPGMA fusionmethod. This analysisof CALM's priorityfloralist(CALM,1996)were vegetationpatterningwascarriedoutattwoscales locatedduringthecourseofthesurvey.Thesetaxa - alargerscaleusinglistsofspeciesfromeach requirefurtherinvestigationbeforeconsiderationof wetland area and thefinerscalebased on listingasthreatenedflora.Rumexcrystallinuswas individualquadrats.Treeswereexcludedfromthe collectedontheMcNeill claypan ~ Carnarvon; analysisofthequadratdata. thiswas thefirstcollectionofthisspeciesin a ~ chemistrydatawere collectedduringthe Western Australiasince1885whenitwasrecorded samplmgofthesesitesformacro invertebratesin byCareyontheLyndonRiversome150kmtothe August 1994and/orMarch 1995[seeHalse etal. north. (2000)fordetailedmethods]. Thesedatawereused Five taxaendemictothestudyareawere ascorrelatesforsoilchemistryinanalysisof recordedfromthefifty-eightwetlands(Table1).Of between ~ differences.Statisticalrelationships these,Calandrinia sp.Coolcalalaya(GJKandNG betweenSItegroupswere testedusingKruskal- 698)andLythrumsp.Towrana(RJC2183)appearto Wallis non-parametricanalysisofvarianceand berestrictedtoedgesofephemeralclaypans. Mann-Whitney U-tests(Siege!,1956). Mynocephalusgascoynensismsmayalsoberestricted Specieshabitatpreferenceanddistribution tothishabitat. informationwerecompiledfromcollectionsheldin Sevenofthetaxarecordedinthisstudyaretotally theWestern Australian Herbarium. Nomenclature restnctedtowater bodies while anothersix followsGreen (1985)andcurrentusageatthe generallygrowasemergentsaroundpermanentor Western Australian Herbarium. Selected voucher semipermanentwetlands (Table2).Two ofthe specimenshavebeenlodgedinthatinstitution.

Table2 Listofsubmergedandemergentaquatictaxa recordedfrom58wetlands inthesouthern RESULTS Carnarvon Basin (*indicatesintroduced species). Flora The floraof thewetlandswas foundtobe Submergedorfloating Emergentaquatic dominatedbyaridzonespeciesOessop,1985).Two aquatic hundredandsixtythreetaxawere recordedfrom *Callitrichestagna/ls Alternantheranodiflora thefifty-eightwetlandareas(Appendix1).Thebest *Crasslllanatans Avicenniamarina representedfamilieswere Asteraceae (32native,4 Elatmegratioloides Ballmeaarticulata introducedspecies),Poaceae (25native, 10 Najasmarina EleocharisaCli/a introducedspecies),Chenopodiaceae (24natives), Potamogetonpectinatlls Eleochllrisgeniclllata Cyperaceae (17native,1introducedspecies)and RlIppUlmegacarpa Schoenoplectllssllblllatlls Scrophulariaceae (10natives). This composition RlIpplapolycarpa Typhadomingensis clearlyreflectsthearidnatureoftheflora.At the RlIppiatuberosa 178 N.Gibson,G.].Keighery,M.N. Lyons

0.0 0.2780 0.5560 0.8340 1.1120 1.3900 1 1 1 1 1 1 WETLAND COMMUNITY TYPE 1 CB15 CB29a CB16 CB78 CB79 1----- CB9 1 CB9a 1 1 __

CB76 1 CBn 1 1__ CB25 1 ~ ~ 1 CB36 1 -;-

1 WETLAND COMMUNITY TYPE 2 1 CB20 1 CB4 1 CB49 1 CB93c 1 CB30 1 CB34 1 1 CB5 ------1--1----1 1 1 1 WETLAND COMMUNITY TYPE 3 1 1 CB35a 1 1 CB93d 1 1 CB43 1 1 CB56a 1 1 CB58b ------1----1- 1 1 1 WETLAND COMMUNITY TYPE 4 1 1 CB27a 1 1 CB56 1 1 CB93 a 1 1 CB93a 1 1 1 1 1 CB44 1 1 1 CB54 1 1 1 1 McNei11 1 1_ 1 1 1 CB75b 1_ 1 b 1 1 CB54a 1 1 1 1 CB73 1 1 1 1 CB82 1__1 -,- 1 1 1 1 WETLAND COMMUNITY TYPE 5 1 1 CB27b1 1 1 CB27c ------1- 1 1 CB6a1 1 1 1 CB6a2 1- 1 1 CB58e ------1- 1 1 CB58d 1 1 1 CB6b 1- 1 1 CB27b2 1 1 1 CB75e 1 1 1 CB62b 1 1 1 CB70b 1 1 1 CB75a 1 1 1 CB68 ------1-1 1 1 1 CB38 11 1 1 1 CB51 ------11--1-1-- 1 1 1 WETLAND COMMUNITY TYPE 6 1 1 CB67a _ 1 1 CB67b 1 1_ 1 1 1 WETLAND COMMUNITY TYPE 7 1 1 CB42 1 1 CB62 1 1 CB62a 1 1 CB62c 1 1 1 CB93g -,-- -;- ---; ---;- 1_1 1 1 1 1 1 1 0.0 0.2780 0.5560 0.8340 1.1120 1.3900

Figure2 Dendrogram showingtheUPGMAfusionofthewetland vegetationof57wetland areasinthesouthern CarnarvonBasin. Wetland flora and vegetation 179 submerged aquatic taxa are widespread weed Vegetation species across southern Australia (Aston, 1973). The habitat preference of other taxa generally restricted Large scale pattemillg the wetlalld classification to wetland areas is listed (Appendix 1) where such Amalgamation of some species was necessary as information could be determined from herbarium material could not always be positively identified labels. This infonnation is lacking for some genera to species level (e.g. Centipeda spp.). One of the where the material is on loan. claypans was completely bare of vegetation and The poor state of knowledge of the wetland flora was excluded from the analysis. Of the 259 taxa of the southern Carnarvon Basin is highlighted by included in the final dataset some 142 taxa (55';;,) the first collections of one apparently undescribed occurred only at one wetland area. The dataset was Calandrillia sp. and major range extensions of 15 analysed with and without singletons (i.e. taxa that other taxa (Appendix 1). occurred at only one site). The results of both Thirty-four taxa reach the northern limit of their analyses were similar with clearer patterning seen range in the study area and another 18 primarily in the analysis with the singletons excluded; these tropical taxa reach the southern ends of their range. are the results presented below. Species richness The study area straddles Beard's (1990) major varied from one to 25 taxa per wetland (with phytogeographic boundary between the temperate singletons excluded), with individual taxa occurring south west and the Eremaean zones (Figure 1). at between two and 22 wetlands. In this analysis

Lake MacLeod

Boolathana. A A Mooka Station AA ?1 ;::I S. 6 ~ / pnng A .. /' Gascoyne Camarvon .. Junction McNeill Claypan Salt I Creek

@$ Wooramel

~ N Wetland types I 1111 02 ", Kilometres A4a 4b

Figure 3 Distribution of the seven wetland community types across the southern Carnarvon Basin. .... Table 3 Two way table showing species distribution across the seven wetland community types identified by the UPGMA fusion. QC <:) WETLAND COMMUNITY TYPES SPECIES 1 2 3 4& 4b 5 6 7 GROUPS

CCCCCCCCCCCC Iccccccc 1ccccc Icccc 1CCMCCCC Iccccccccccccccclcclccccc BBBBBBBBBBBB IBBBBBBB IBBBBB IBBBB 1BBcBBBB IBBBBBBBBBBBBBBBIBBIBBBBB 1217799772331244933513945512599145N75781226655627677635166146669 WETLAND AREAS 59689 a6758610 9304 15336817633144e5432177aa88b75205881177122223 a a I clad abla al iba Ibc12cd bcbba labl acg 1 I 1 I 11 2 I 1 IIII 1 I 1 I 1 I I 1 IIII I 1 Actinobole uliginosum I I *II II A Isoetopsis graminifolia II*I I II Peplidium sp. C (Burbidge and Kain 8152) I 1 * I 1** * II Calotis hispidula I * 1 * I 1 I 1 ------i------+-----+----+------+------+--+-- Calandrinia granulifera Gnephosis eriocephala Gypsophila australis B Crassula natans Pentaschistis airoides Melaleuca uncinata ------+------+-----+- -+------+----- +- +--- - Atriplex holocarpa Myriocephalus gascoynensis MS Eragrostis leptocarpa Trichanthodium skirrophorum Calotis multicaulis Cyperus rigidellus Calandrinia ptychosperma Bergia auriculata C Bergia perennis subsp. obtusifolia Calandrinia pumila Goodenia neogoodenia Eragrostis pergracilis Peplidium aithocheilum ** Glossostigma drummondii Marsilea angustifolia Isoetes sp. * ------+------+-----+-- -+------+------+--+-- Calandrinia sp.Coolcalalaya (GJK and NG 698) Pogonolepis muelleriana Crassula colorata D Eragrostis dielsii ** * * ***** Triglochin calcitrapum * ------+------+-----+- -+------+------+--+---- Asphodelus fistulosus I 1*****1 I Myriocephalus guerinae II ** I * I Atriplex lindleyi * II * I 1 Dysphania rhadinostachya Sclerolaena diacantha E Diplachne muelleri Atriplex semilunaris Cenchrus ciliaris Centaurium spicatum * Lythrum sp.Towrana (RJ Cranfield 2183) Sclerolaena recurvicuspis Tetragonia diptera ------+------+--- -+- --+------+------+--+----- Alternanthera nodiflora ***** * Centipeda sp. * ** ***** * *** * ****** Myriocephalus nudus F Ammannia multiflora Eriochloa procera ** Marsilea drummondii Ranunculus pumilio Pseudognaphalium luteoalbum ------+------+-----+-- -+------+------+--+- Chenopodium auricomum Mimulus gracilis Eleocharis pallens Goodenia corynocarpa **** Eulalia aurea *** Sporobolus mitchellii G Cyperus vaginatus *** Stemodia viscosa Emex australis Eragrostis australasica Sonchus oleraceus Muehlenbeckia cunninghamii Polycarpon tetraphyllum ------+------+--- -+-- -+------+------+--+--- Brachyscome ciliaris Eriachne flaccida * H Goodenia berardiana Calandrinia polyandra Lotus cruentus * Paractaenum novae-hollandiae * * ------+------+-----+----+------+------+--+----- Apium prostratum Tetragonia cristata * Medicago polymorpha Wurmbea murchisoniana Heliotropium curassavicum I Typha domingensis Ruppia sp. cyperus gymnocaulos Eucalyptus camaldulensis var. obtusa Polypogon monspeliensis Lotus australis Samolus junceus ------+------+-----+----+------+------+--+----- ....QC Table 3 (cont.) N

WB'l'LAND COMMtmI'l'Y 'l'YPES SPECIES 1 2 3 4a 4b 5 6 7 GROUPS

cccccccccccclccccccclccccclcccclcCMcccclccccccccccccccclcclccccc BBBBBBBBBBBBI BBBBBBBI BBBBB1 BBBB1 BBcBBBB1 BBBBBBBBBBBBBBBIBBIBBBBB 1217799772331244933513945512599145N75781226655627677635166146669 59689 a6758610 9304 15336817633144e5432177aa88b75205881177122223 a a I clad abla al iba Ibc12cd bcbba labl acg IIII 11 2 I I

1 1 1 1 II I 1 1 1 II I I 1 1 1 Avicennia marina II 1 1 Halodule uninervis I I 1 1 Halosarcia pterygosperma subsp. denticulata * *** II 1 Ruppia tuberosa II 1 Frankenia pauciflora ** II 1 Muellerolimon salicorniaceum ** * *** II I Sarcocornia quinqueflora ******* II 1 Halosarcia halocnemoides ** *** *** II I J Halosarcia indica **** *** ** 1** I I Cotula cotuloides * 1* I I Triglochin centrocarpum 1 1 I Lawrencia viridigrisea ** 1 I I Neosciadium glochidiatum 1 1 I Samolus repens 1 1 I Triglochin mucronatum I * I I Rostraria pumila I 1 I Senecio glossanthus ** II I ------+------+-----+----+------+------+--+----- Anagallis arvensis * ***1 1 1 1 * Schoenoplectus lateriflorus * **1* 1 1 1* Eucalyptus victrix *** **1* 1 I 1 Daucus glochidiatus *** I 1 I 1 Glossostigma diandrum * **1 1 I 1 Setaria dielsii ** 1 1 1 1 Isolepis congrua * **1 1* *1 1 Centrolepis eremica * * I 1 1 1 * Hypericum japonicum * * 1 1 I I Brachyscome iberidifolia *1 1 * 1 1 Cyperus aff. cunninghamii (GJK and NG 592) **1 1 1 1 Lipocarpha microcephala **1 1 1 1 Oldenlandia galioides **1 1 I I Schoenus elegans **1 1 1 1 Drosera indica **1 I * I I Eragrostis basedowii **1 I * 1 1 Elatine gratioloides **1 1* 1 1 Wahlenbergia tumidifructa **1 1 1 1 WetIand flora and vegetation 183 site groups are discussed at the seven group level is the weed Asphodelus fistulosus. One species poor (referred to as 'wetland community types' or riverine site also belongs to this group. The 'community types') which best reflected the scale of occurrence of A. fistulosus indicates moderate levels patterning seen in the field. of grazing. Whether the low species richness of The wetland areas sampled showed a high level these sites is also related to disturbance or local soil of heterogeneity in species composition (Figure 2). or moisture conditions is not clear. Several The primary division in the classification separates chenopod species were also recorded from this the vegetation of the saline flats (wetland wetland community type. community type 1) from all other sites. This can Community types 4 and 5 encompass riverine also be seen in the sorted two-way table (Table 3) wetlands and ephemeral claypans and swamps where species group J is largely restricted to this characterised by the occurrence of Alternallthera community type. Most taxa in this group are highly llodifiora, Celltipeda spp. and to a lesser extent faithful to it and species such as Halosarcia Myriocephalus nudus. Community type 4 differs halocllemoides, H. illdica and Sarcocornia quillquefiora from type 5 by the reasonably high frequency of have a high level of constancy. All other species other taxa in species group F while largely lacking groups are completely or almost completely lacking taxa in species groups A, B, C, 0 and E which from this wetland community type except for low further characterise community type 5, although frequencies of species in species groups 0 and E. constancy of species in these groups is generally Species richness in this community type ranges only moderate to low. Community type 4 was from three to 16 taxa/wetland (average 6.8). This found both in riverine vegetation and ephemeral community type is largely restricted to the near wetlands of the claypans and swamps while coastal samphires and birridas between Shark Bay community type 5 was largely restricted to claypans and Lake MacLeod (Figure 3). One outlying site and swamps (Table 4). was found at Salt Gully near Gascoyne Junction. Two distinct subtypes are discernible in wetland The inland mangrove stands from Lake MacLeod community type 4. Subtype 4a was the most are members of this group. While coastal samphires species-rich of the wetland groups sampled (mean and mangroves were not sampled in this study they species richness of 17.0/wetland), while community appear very similar to this community type. subtype 4b was much less rich (mean species Wetland community type 2 is generally associated richness of 10.6 taxa/wetland). Both subtypes are with deeper river pools; one wetland formed from characterised by the occurrence of taxa in species an uncapped artesian bore also belongs with this group F, while the occurrences of species in species group. This community type is characterised by the groups G and K define the subgroupings. occurrence of species group I. While a number of Three of the four sites in community subtype 4a species are restricted to this community type, are riverine and this subgroup is characterised by a constancy values rarely exceed 50%. Species in moderate to high fidelity and constancy of taxa in species groups A, B, E and H are completely species group K. The river pools tended to have a lacking, while species frequency in other species variety of microhabitats generally with a dense groups is low to very low. Average species richness band of sedges (Schoenoplectus lateriflorus and / or was low (6 taxa/wetland). This community type Cyperus vaginatus) along the water's edge. This was associated with the deepest pools which would community subtype was found on upstream areas not dry out in any but the driest years; it occurred of the Wooramel River and its tributaries and from the Murchison River to north of Carnarvon around a large semi-permanent water body on and east along the Wooramel River (Figure 3). Boolathana Station (north of Carnarvon) (Figure 3). Community type 3 are typically species-poor Community subtype 4b is restricted to claypans, claypans (mean species richness 5.6 taxa / wetland). ephemeral swamps and two river pools in the north The only species constant and faithful to this group west of the study area (Figure 3). Wetlands in this

Table 4 Comparison of the occurrences of the seven wetland community types with a broad habitat classification.

Wetland Birrida Samphire River Spring Ephemeral Claypan community (gypsum pool swamp type pan)

1 7 4 1 2 6 3 1 4 4 6 1 4 5 1 8 6 6 1 7 4 184 N. Gibson, G.}. Keighery, M.N. Lyons

group were characterised by taxa in species group yards. There are no understorey species left at these G while almost totally lacking taxa in species group two sites. The overstorey tree is Eucalyptus K, which are typical of community subtype 4a. cmnaldulensis subsp. obtusa. These two sites form Subtype 4b wetlands tended to have a significant wetland community type 6 based on the sole shrub component and the claypans and swamps occurrence of this species. One site was located at were quite extensive. the spring itself while the second was along the The majority of the ephemeral swamps and watercourse below the spring. claypans form wetland community type 5, and The final community type (type 7) is comprised of while Centipeda spp. and Alternanthera nodiflora were tall dense species-poor sedgelands that occur at the also common in this community type, the herbs of less modified springs and the streams draining the species groups A, B, C and D clearly differentiate it west side of the Kennedy Range and as far south as from the previous type. The sites in this type were a dammed section of the Wooramel River. These quite heterogenous, with few constant taxa, but stands are typical of permanent or near permanent none the less species rich (mean 10.7 taxa/wetland). water bodies. Cyperus vaginalis is constant in this The most constant taxon was Eragrostis dielsii, group but is also common in wetland community occurring in 75% of sites in this group and totally subtype 4b. Typha domingensis is also a common lacking from the related community type 4. Species component of this community type. Mean species group C is most typical of this community type and richness is 3.2 taxa/wetland. is composed of annual taxa that either germinate in While no soil sampling or analysis was very shallow waters or rapidly germinate as the undertaken during the present study, Halse et al. clay soils dry. This community type was very (2000) collected water chemistry data from all but widespread across the study area (Figure 3). One one of these wetlands in their study of invertebrate riverine site was classified into this group. That site patterning. This water chemistry data could be had rich alluvial soil on which suites of annuals expected to be related to aspects of the soil were growing. Typically the riverbanks in the study chemistry. Measures of electrical conductivity and area are either sandy or rocky. turbidity highlight patterns as discussed above A series of permanent springs are found along the (Figure 4). western foothills of the Kennedy Range. These have When the claypans fill, the water becomes red until recently long been used as watering points for brown in colour due to suspended clay particles. cattle. Some of the springs are fenced off and water Only wetland community types 3, 4, and 5 occur in piped from them but two have been used as cattle claypans, with the percentage of claypan sites

Turbidity (NTU) Kruskal-Wallis: 20.236, df: 6, Probability = 0.0025

0.5000E-01 0.1550E+05 0.3100E+05 0.4650E+05 0.6200E+05 GRP +------+------+------+------+ 1 *M==3------U 2 ** 3 L------l======M======D======u 3 4 LD======M======3------U 5 *======M======3------U 6 * 7 *

l Electrical conductivity (mSm- ) Kruskal-Wallis: 37.524, df: 6, Probability < 0.0000

29.00 0.5102E+05 0.1020E+06 0.1530E+06 0.2040E+06 GRP +------+------+------+------+ 1 L------1======D======M======3------U 2 *M3U 3 * 4 *3U 5 *U 6 **=* 7 *==*=3U

1 Figure 4 Turbidity (NTU) and electrical conductivity (mSm ) for the seven community types. L = lower limit, 1 = Mean - 1 standard deviation, M = mean, 0 = Median, 3 = Mean + 1 standard deviation, U = upper limit, * = more than one symbol at print position. Wetland flora and vegetation 185

10 2... 3 4

8

9 11

12

Figure 5 Distribution of the 14 site groups based on individual 1 m 2 across the southern Carnarvon Basin. Table 5 Two-way table derived from the UPGMA classification of the 124 wetland quadrats from the southern Carnarvon Basin study area.

SITB GROUPS SPIICIBS 3 • 5 6 7 10 11 12 13 GROUPS

1797222111372429999117923112124bI33177777126647166613345951292666554555577129924999393767766766222663555122226522125534669475588b5751355M 58_6999556860_0_aa_168a56169174r I 00 1667691 022_81222144_3_15_5aab113688855173374333538520522588777aa6488 I 7777b877 176682223934422r434 1514c QUADRAT IDENTIFER _Cl-aaa_a_Cl-lL-q 1__I_a I a_o 1_1__I_acCl-I_I_qqcq1_Cl-22__abbdbc I a_aa_a_aad_bbbabbc_cccll_acc Ibbbb_dbb I a_a_a_II_a_oa_1 a-y/ qql lL-qqCl-qq3q2qq31 qqqqq ICl-I_qw I qq I qqqqqIIL-lqIqqq I qq22_31 q4IL-qqqCL--I_IL-Cl-lL-

III I II Actinobole uliginosum III I II Calandrinia granulifera II I I I I Pentaschistis airoides III I II Crassula natans I I' I I II • Wahlenbergia preissii III I II Brachyscome iberidifolia III I II Isoetopsis graminifolia III I II ------+-----+--+---+--+-----+-----+---+------+------+------+------+------+---- Calandrinia sp.Coolcalalaya (GJK and NG 69B) III I' • I I Pogonolepis muelleriana III I' • II Maireana oppositifolia III I' • II b Rostraria pumila III I" II ------+-----+--+---+--+-----+-----+---+------+------+------+------+------+---- Sergia perennis subsp. obtusifolia I I I '1 I•I Calandrinia pumila III I II Goodenia neogoodenia III I II Gnepbosis eriocephala III I II WUrmbea murchisoniana I II I I I Isoetes Sp. III I I "I c Triglochin calcitrapum •I I I I' I I Elatine gratioloides III I• I' I Melaleuca uncinata III '1 I' I Peplidiurn aitbocheilum III I I' I Glossostigma drummondii III I •I....•I Peplidium sp. C (Burbidge and Kain 8152) III I I·..•• '1 Marsilea angustifolia III I '1 I ------+-----+--+---+--+-----+-----+---+------+------+------+------+------+---- Asphodelus fistulosus I I I II I I I I MYriocephalus guerinae I I I II I II I Atriplex lindleyi '1 II I I I I I I pysphania rhadinostachya I II I II I I I I Cenchrus ciliaris II 1"1 I I I I I I Eragrostis dielsii IIII I 1"'1 I•I I d Calotis hispidula IIII II .... I I' I I Atriplex holocarpa III I I I I I I I MYriocephalus gascoynensis MS II II I I I II I Trichanthodium skirrophorum I I II II I I I I Eragrostis leptocarpa II II II I I I I ------+-----+--+---+--+-----+-----+---+------+------+------+------+------+---- Brachyscome ciliaris III II I •II •I Lotus cruentus III II I .. II I Diplachne muelleri I II II I "I I I Eriachne flaccida III II I "I I I • Chenopodium auricomum I I I I I •I "'1 I I Calotis mul ticaulis I I I I" I I • I I '1 Cyperus rigidellus I I I II I "'1 I I Goodenia berardiana III II I •II I ------+-----+--+---+--+-----+-----+---+------+------+------+------+------+---- Avicennia marina I I I I .... I II I I I I Halodule uninervis III I" I I I I I I I Halosarcia pterygosperma subsp. denticulata I III I I I I I I I Ruppia tuberosa 1"1 II I I I I I I I Eragrostis australasica I II II II I I I I .... Ruppia sp. III I 1'''1 II I II I TY,pha domingensis IIII I'" 'I I I II I 1 1 Halosarcia pruinosa 1 f Heliotropium curassavicum 1 1 'I Samol us juneeus 1 1 'I Polypogon monspeliensis 1 1 'I Cyperus gymnocaulos 1 I"'" 'I Tetragonia cristata 1 1 1 Decazesia hecatoeephala 1 I" 1 Phalaris minor 1 I" 1 Bergia auriculata 1 1 1 Schoenoplectus lateritlorus 1 1 1 Calandrinia polyandra , 1 1 1

Cotula eotuloides 1 Lawrencia viridigrisea 1 Samolus repens 1 Neoseiadium glochidiatum 1 Triglochin mueronat'UlTl 1 Parapholis ineurva 1 Frankenia pauei t lord 1 Halosarcia haloenemoides 1 Muellerolimon salicorniaceum 1 Sarcocornia quinquef1 ora 1 Halosarcia indica ------+- ----+ --+- --+ --+-----+ -----+---+ ------+-1 ------+ ------+------+ ------+---- Alternanthera noditlora Cen t ipeda sp. Cyperus vaginatus Eleocharis pallens Goodenia corynOCdrpa Eulalid aurea h Myriocephalus nudus Sporobolus mitchellii Marsilea drummondi i Ranunculus pumilio Helipterum strictum Streptoglossa tenui t lora Paractaenum novae-hollandiae Pseudognaphalium luteoalbum ------+-----+--+---+--+-----+-----+---+------+------+------+------+------+---- Ammannia multiflora 1 Eriochloa procera 1 Emex australis 1 Mimulus gracilis 1 Muehlenbeckia cunninghamii 1 Stemodia ViSCOSd 1 Senecio glossanthus ------+-----+--+---+--+-----+-----+---+------+------+------+------+------+----I' Anagallis arvensis 1 1 Daucus glochidiatus 1 1 Setaria dielsii 1 1 Cyperus afL cunninghamii (GJK and NG 592) 1 1 ' Wahlenbergia tumiditructa 1 1 Drosera indica 1 1 Lipoearpha microcephala 1 1 Eragrostis basedowii 1 1 Glossostigma diandrum 1 I .. Isolepis eongrua 1 I .. ------+-----+ --+ ---+ --+-----+-----+---+ ------+ ------+ ------+------+ ------+ - - -- Atriplex semilunaris 1 Dysphania glomulifera subsp. eremaea 1 Tectieornia verrucosa 1 Tragus australiallus 1 k Tetragonia diptera 1 Sclerolaena recurvicuspis 1 Sonehus oleraceus 1 Ca 1andrinia ptyehosperma 1 Centaurium spieatum 1 Crassula colorata 1 188 N. Gibson, G.]. Keighery, M.N. Lyons increasing from 31% in community type 4 to 80'Yo in Table 6 Mean species richness of the 14 site groups community type 3. Mean turbidity values increase identified in the analysis of the 124 wetland in the same fashion. Only one wetland in the saline quadrats. community type 1 has turbidity values approaching Site Mean species Number of those of claypan groups. The high levels of electrical group richness quadrats conductivity seen in community type 1 clearly separates it from all other wetland types. Essentially 1 4.2 19 the same pattern was seen for other elements such 2 2.2 5 3 1.0 2 Ca, Mg, Na, K, Cl, and S04' The only other 4 1.7 3 community types with significant but much lower 5 1.5 2 levels of electrical conductivity, Ca, Mg, Na, K, Cl, 6 1.8 5 and S04 were the wetlands fed from springs along 7 1.4 5 the western edge of the Kennedy Range 8 2.0 3 (community types 6 and 7). 9 3.8 6 10 9.9 15 Fine scale patterning - the quadrat classification 11 6.6 29 12 4.0 8 In addition to the floristic analysis of the wetland 13 5.6 18 areas a second analysis was undertaken of the 1 m x 14 1.3 4 1 m quadrats which sampled the finer scale vegetation pattern. In all, 124 quadrats were established in 55 of the 58 wetland areas. Three of Other species-poor site groups include groups 4, the wetland areas had no shrub or herb layer. Two 7, 8, and 9 which are mostly fringing freshwater hundred and thirteen taxa were recorded from sedgelands or Typha stands, and site group 14, the these quadrats. Of these 213 taxa, 99 (46%) only cane grass swamps (Eragrostis australasica). The occurred in one quadrat; these taxa were excluded characteristic taxa of each of these communities are from the analysis as discussed above. With these different but all belong to species group f. The other exclusions species richness varied from one to 13 species-poor site group (group 5) occurred on the taxa/quadrat with individual taxa represented in Wooramel River flats which had been heavily between two and 31 quadrats. grazed. The sole species recorded in these quadrats Fourteen quadrat groupings (referred to as 'site was the introduced Buffel Grass (Cenchrus ciliaris). groups') and 11 species groupings were recognised The three remaining site groups are herblands (Table 5). The data set showed high levels of (groups 10, 11, and 12). Eragrostis dielsii is the heterogeneity with few quadrats in many of the site typical species of site group 10. Species groups a, b, groups and generally low species richness (Table 6). d and k are largely confined to this type. Site groups The first division in the classification again 11 and 12 differ from each other primarily in the separated the Halosarcia flats (site groups 1 and 2) proportions of species groups c and h (Table 5). Site from the rest of the data set. The two remaining group 13 is generally a sedgeland on riverbanks or saline groups, the quadrats containing Ruppia drainage channels usually dominated by Cyperus tuberosa (site group 3) and the mangrove quadrats vaginatus. Taxa in species group h are also well (site group 6), totally lack taxa in species group g represented in this group. which are faithful to site groups 1 and 2. This fourteen site group classification showed a

Table 7 Comparison of the number of quadrats of the fourteen site groups with a broad habitat classification.

Site group Birrida (gypsum pan) Samphire Spring River pool Ephemeral swamp Claypan

1 14 1 2 2 2 2 3 3 2 4 2 1 5 2 6 4 1 7 1 2 2 8 3 9 6 10 2 3 9 11 1 10 13 5 12 6 2 13 11 7 14 4 Wetland flora and vegetation 189 strong correlation with the broad habitat of site group 3 were restricted to Shark Bay and the classification, again with the wetlands associated mangrove dominated quadrats (group 6) were with major stream being the most diverse (Table 7). restricted to Lake MacLeod. However, as was noted Turbidity and electrical conductivity of the wet previously, the coastal mangrove communities were season water body showed significance differences not sampled in this survey. Site groups 5, 8, 9, and between site type means (Figure 4). The occurrence 14 were found to be quite restricted, with group 8 of claypans in site types 10 - 14 is highlighted by being restricted to Mooka Springs area, group 9 to high mean turbidity values, and some of the saline the Murchison and Wooramel Rivers and group 14 water bodies had slightly elevated turbidity (Figure (cane grass swamps) to the area around Carnarvon 6). The water bodies associated with site types 1, 2, (Figure 5). In contrast site groups 10 and 11 were 3 and 6 were moderately to highly saline. A lesser widespread in the study area and group 13 was saline influence can also be seen in site types 7, 10 widespread in the northern half. and 13. Many of the wetland areas had more than one site The highly saline site groups (groups 1, 2, 3 and group present and this concentric zonation of the 6) were largely restricted to the coastal belt except vegetation around the wetland was an obvious for a quadrat at Salt Creek near Gascoyne Junction feature of the study area. At five of the 57 wetlands (Figure 5). The hyper saline Ruppia tuberosa quadrats three different site groups were recorded, a further

Turbidity (NTU) Kruskal-Wallis: 45.819, df: 13, Probability < 0.0001

O.5000E-01 0.1550E+05 0.3100E+05 0.4650E+05 0.6200E+05 GRP +------+------+------+------+

1 * 2 *==M===3---U 3 * 4 * 5 * 6 * 7 * 8 * 9 * 10 L==D======M======3------U 11 *=====M======3------U 12 *======M======3------U 13 LD======M======3------U 14 *======D=====M======3------U

l Electrical conductivity (mSm- ) Kruskal-Wallis: 74.103, df: 13, Probability < 0.0001 29.00 0.5102E+05 0.1020E+06 0.1530E+06 0.2040E+06 GRP +------+------+------+------+ 1 L------1======D====M======3------U 2 L------l======DM======3 ----U 3 *======*======* 4 * 5 * 6 L1=M=* 7 *===D==M======3----U 8 9 * 10 *======M======3 ------U 11 ** 12 * 13 *M=3 -U 14 *

J Figure 6 Turbidity (NTU) and electrical conductivity (mSm ) for the 14 site types. L lower limit, 1 Mean 1 standard deviation, M mean, D Median, 3 = Mean + 1 standard deviation, U =upper limit, * = more than one symbol at print position. 190 N. Gibson, G.]. Keighery, M.N. Lyons

19 had two distinct site groups and a single group Gibson et al., 2000). This rare component of the was recorded from the remaining wetlands. floras adds significantly to the biodiversity but is not a predictable component of vegetation communities. The conservation status of this DISCUSSION component of the flora will continue to be difficult The present survey has improved on the poor to assess. state of knowledge of the flora and vegetation of The highly variable nature of wetlands in south- the seasonal and permanent wetlands of the west Western Australia has been reported both at southern Carnarvon Basin. During the course of this the landscape scale (Churchward et al., 1988) and work one apparently new species was collected, plant and animal community scales (Gibson et al., seven new populations of priority flora were 1994; WardelI-Johnson and Williams, 1996; Wardell- located and major range extensions were recorded Johnson and Horwitz, 1996). In the survey of the for a further 15 taxa (Appendix 1). The flora of the plant communities of the Swan Coastal Plain the wetlands is largely arid Oessop, 1985) but with both seasonal wetlands were found to comprise 16 of the a significant temperate component (34 taxa reaching 30 groups defined (Gibson et al., 1994), while in a their northern range limit) and a significant tropical study in high rainfall forest areas near Walpole component (18 taxa reaching their southern limit). wetlands comprised 22 of the 44 community types The composition and affinities of the wetlands defined (Wardell-Johnson and Williams, 1996). The reported here do not accord with the recently reasons for these high levels of heterogeneity are published review of biogeography of freshwater not clear but appear to result at least in part from plants, which concentrated almost entirely on wet differences in substrate and differences in period temperate and tropical regions Oacobs and Wilson, and depth of inundation. High levels of 1996). Our study area overlaps the northern edge of heterogeneity can be seen from the present study at their south west Western Australian region but the both scales examined. affinities of the freshwater plants (defined in Jacobs It could be argued that part of the reason for and Wilson's study as plants that grow in or near heterogeneity at the larger scale could be related to water) in our study are largely with the arid zone the obvious zonation of the vegetation around Oessop, 1985) and not with New Zealand. Jacobs many of the wetlands. Vegetation zonation is and Wilson's (1996) work does indicate the lack of commonly observed in wetlands across Western knowledge of the seasonal and permanent wetlands Australia (Hopper et al., 1996; Halse et al., 1993). in the arid temperate zone (south of the Tropic) is However when sampled at a finer scale the wetland not restricted to southern Carnarvon Basin but is classification still showed a high level of true across Australia. With an increase in heterogeneity, indicating factors other than knowledge of the wetland systems of arid areas of zonation are involved. both the temperate and tropical zones the very While some of the community types in the seven strong division between the tropical and temperate group classification generally occur in a particular regions reported by them can be expected to break wetland habitat (e.g. birridas/samphires or down. claypans/ephemeral swamps), none were restricted Detailed quadrat-based floristic surveys of the to these habitats (Table 4). At the finer scale Swan Coastal Plain, the coastal communities of the sampling there is a much stronger correlation Warren bioregion (Thackway and Cresswell, 1995), between site groups and wetland habitat (Table 7). and the Tingle forests of the high rainfall zone have Even at this scale, however, some site groups are all shown that there is a high proportion of recorded from several wetland habitats. The most naturally rare species in south west Western diverse wetland habitat was the riverine wetlands Australian ecosystems (Gibson et al., 1994; Gibson reflecting the heterogeneous nature of these areas and Lyons, unpublished data; Wardell-Johnson and with variation in substrate composition, texture, Williams, 1996). In all three studies approximately slope, degree of disturbance and salinity to which 25% of total floras (1485 taxa in 509 quadrats; 901 the floristic groups respond. taxa in 301 quadrats; and 857 taxa in 441 quadrats Examination of the distribution maps of the respectively) were only recorded at one site. Data floristic groupings indicate only moderate from the present survey shows the same pattern biogeographic patterning (Figures 3 and 6). In the with taxa being recorded at only one site seven group wetland classification, the saline comprising 55% of species at the individual wetland community type is generally restricted to the coastal scale (58 wetland areas) and 46% at the 1 m 2 belt but with an outlier near Gascoyne Junction and quadrat scale (124 quadrats). The regional type 7 is restricted to wetlands at the base of the vegetation survey of the southern Carnarvon Basin Kennedy Range south to Wooramel River. The other study area and a detailed study of the sandplain community types (with the exception of type 3 vegetation in the south of the study area show which are highly disturbed) are widespread across essentially similar trends (Keighery et al., 2000; the region, although community type 4 only occurs Wetland flora and vegetation 191 in the north of the study area (Figure 3). Department of Conservation and Land Management, At the finer scale of analysis of the 14 site groups, Perth. much stronger biogeographic patterning is seen. Churchward, H.M., McArthur, W.M., Sewell, P.L. and Eight site groups (groups 1, 2, 3, 5, 6, 8, 9 and 14 - Bartle, G.A. (1988). Landforms and soils of the south Figure 5) have a restricted distribution, while site coast and hinterland, Western Australia - Northcliffe groups 10, 11 and 13 are widespread. This increased to Manypeaks. CSIRO Division of Water Resources. Divisional Report 88/1. patterning results from the finer subdivisions Gibson, N., Keighery, B.J., Keighery, G.J., Burbidge, AH. possible in the larger dataset. and Lyons, M.N. (1994). A Floristic Survey of the While most wetlands visited were not obviously Southern Swan Coastal Plain. Unpublished Report for impacted by grazing, the use of two springs in the the Australian Heritage Commission prepared by Kennedy Range as cattle yards had resulted in the Department of Conservation and Land Management total removal of the understorey (wetland and the Conservation Council of Western Australia community type 6). Grazing impacts could also be Onc. ). seen in the claypans and riverine areas of wetland Gibson, N., Burbidge, AH., Keighery, G.J. and Lyons, community type 3. These sites were generally M.N. (2000). The temperate to arid transition of the dominated by Asphodelus fistulosus, a weed Irwin - Carnarvon phytogeographic boundary, indicative of over grazing (Parsons and Western Australia. Records of the Western Australian Cuthbertson, 1992), and were typically quite species Museum Supplement No. 61: 155-173. poor. Green, J.W. (1985). Census of the Vascular Plants of Western The seasonal and permanent wetlands of the Australia. Department of Agriculture, Perth. southern Carnarvon Basin have a diverse and Halse, S.A., Pearson, G.B. and Patrick, S. (1993). poorly known flora. The present study has shown Vegetation of depth-gauged wetlands in nature that patterns in species distribution and species reserves of south-west Western Australia. CALM richness are similar to those reported for wetlands Technical Report No. 30. in higher rainfall area of the southwest, however as Halse, S.A, Shiel, R.J., Storey, AW., Edward, D.H.o., Lansbury, I., Cale, D.J. and Harvey, MS. (2000). in the southwest, the causal factors operating are Aquatic invertebrates and waterbirds of wetlands and still poorly understood. rivers of the southern Carnarvon Basin, Western Australia. Records of the Western Australian Museum Supplement No. 61: 217-267. ACKNOWLEDGEMENTS Hopper, S.D., Harvey, MS., ChappiIl, J.A., Main, AR. Funding was provided by the Commonwealth and Main, B.Y. (1996). The Western Australian biota through the National Reserves System Co-operative as Gondwanan heritage - a review. In S.D. Hopper, Program of the Australian Nature Conservation J.A. ChappiIl, MS. Harvey and AS. George (eds), Gondwanan Heritage: Past, Present and Future of the Agency (now Environment Australia), and by the Western Australian Biota, 1--46. Surrey Beatty and Sons, Western Australian Department of Conservation Chipping Norton. and Land Management. Jacobs, S.W.L. and Brock, M.A (1993). WetIands of Australia - Southern (temperate) Australia. In D.F. Whigham, D. Dykyjova and S. Hejny (eds) Wetlands REFERENCES ofthe World. I: Inventory, Ecology and Management, 244- Anonymous (1975). Geology of Western Australia. 304. Handbook of Vegetation Science Vo!. 15/2. Memoir, Geological SunJey ofWestern Australia No. 2. Kluwer Academic Publishers, Dordrecht. Aston, H.!. (1973). Aquatic Plants of Australia. Melbourne Jacobs, S.W.L. and Wilson, K.L. (1996). A biogeographical University Press, Melbourne. analysis of the freshwater plants of Australasia. Austin, M.P. and Belbin, L. (1982). A new approach to Australian Systematic Botany 9: 169-183. the species classification problems in floristic Jessop, J. (1985). Flora ofCentral Australia. Reed, Sydney. analysis. Australian Journal of Ecology 7: 75-89. Keighery, G.J., Gibson, N., Burbidge, A.H. and Lyons, Australian Nature Conservation Agency (1996). A M.N. (2000). Flora and vegetation of the southern Directory of Important Wetlands in Australia. Second Carnarvon Basin, Western Australia. Records of the edition. ANCA, Canberra. Western Australian Museum Supplement No. 61: 77- Battye, JS. (1915). The History of the North West of 154. Australia. State Library of Western Australia, Perth. Kenneally, K.F. (1978). Notes on the vegetation and flora Beard, J.S. (1990). Plant Life of Western Australia. Kangaroo of Rocky Pool, Gascoyne River. Western Australian Press, Kenthurst. Herbarium Research Notes 1: 29-39. Bettenay, E., Keay, J. and Churchwaod, H.M. (1971). Soils Lane, J.A.K. and McComb, A.J. (1988). Western adjoining the Gascoyne River near Carnarvon, Australian WetIands. In A.J. McComb and PS. Lake Western Australia. csmo Division of Soils, Soils and (eds) The Conservation of Australian Wetlands, 127-146. Land Use Series No. 21: 1-24. Surrey Beatty and Sons, Chipping Norton. CALM (1996). Declared Rare and Priority Flora List Paijamans, K., Galloway, R.W., Faith, D.P., Fleming, 21/10/1996. Unpublished Report, Western Australian P.M., Haantjens, H.A., Heyligers, P.c., Kalma, J.o., 192 N. Gibson, G.]. Keighery, M.N. Lyons

and Loffler, E. (1985). Aspects of Australian wetlands. Sneath, P.H.A. and Sokal, R.R. (1973). Numerical CS/RO Division of Water and Land Resources Technical Taxonomy: The Principles and Practice of Numerical Paper 44: 1-71. Classification. Freeman, San Francisco. Parsons, W.T. and Cuthbertson, E.G. (1992). Noxious Thackway, R. and CresswelI, ID. (1995). (eds) An interim Weeds ofAustralia. Inkata Press, Melbourne. biogeographical regionalisation for Australia: a framework Payne, A.L., Curry, P.J. and Spencer, G.F. (1987). An for establishing the national system of reserves, version inventory and condition survey of rangelands in the 4.0. Australian Nature Conservation Agency, Carnarvon Basin, Western Australia. Western Canberra. Australian Department of Agriculture Technical Bulletin WardelI-Johnson, G. and Williams, M. (1996). A floristic No. 73. survey of the tingle mosaic, south-western Australia: Scott, T.C. (1962). A visit to the salt marsh north of applications in land use planning and management. Carnarvon. Western Australian Naturalist 8: 126-127. Journal of the Royal Society of Western Australia 79: 249- Semeniuk, c.A. (1987). Wetlands of the Darling system - 276. a geomorphic approach to habitat classification. Wardell-Johnson, G. and Horwitz, P. (1996). Conserving Journal of the Royal Society of Western Australia 69: 95- biodiversity and the recognition of heterogeneity in 112. ancient landscapes: a case study from south-western Siege!, S. (1956). Non Parametric Statistics for Behavioural Australia. Forest Ecology and Management 85: 219-238. Sciences. McGraw-Hill, New York. Smith, G.G. and Butler, R.J. (1961). Psilotum nudum at the Manuscript received 3 April 1998; accepted 29 January 1999. Murchison River. Western Australian Naturalist 7: 190. Wetland flora and vegetation 193

Appendix 1 List of taxa found in the 57 wetlands in the southern Carnarvon Basin study area, showing aquatIc taxa (taxa restricted or almost restricted to aquatIc habitats), habitat, the endemic taxa, range ends and geographic limits of the 265 taxa recorded. (Aquatic taxa EAq emergent aquatic; Aq submerged or floating aquatic. Habitat - EW ephemeral wetlands; HW heavy soil wetlands; R riverine; RIF riverine or fringing; S saline. Range ends N northern; S southern. * indicates an introduced species).

Aquatics Taxon Habitat Endemics Range Geographic ends limits

Family: Aizoaceae GUlllliopsis septifraga Tetragollia cnstata Tetragollia diptera

Family: Amaranthaceae EAq Altemallthera nodiflora Hemichroa diandra EW Ptilotus gaudichaudii Ptilotus macrocephalus Ptilotus murrayi

Family: Anthericaceae DicllOpogon tyleri endemic Shark Bay to Lake McLeod

Family: Apiaceae Apium amlllurn Apium prostratum Daucus glochidiatlls Neosciadium glochidiatum N Dirk Hartog Island Trachymelle glaucifolia

Family: Asphodelaceae * Asphodellls fistlllosus

Family: Asteraceae

Actillobole llliginosllm Allgiallthus tomelltosllS * Bidells bipillnata Brachyscome ciliaris Brachyscome iberidifolia Calotis hispidllla Calotis mlllticalllis Celltipeda cllllllillghamii HW N Rocky Pool, Gascoyne River Centipeda rnillima HW S Carnarvon Celltipeda thespidioides HW S Gascoyne River Chthollocephalus pselldevax N 40 km N Gascoyne Junction Cot ilia cotllloides HW N Dirk Hartog Island Decazesia hecatocephala S 93 km N Carnarvon Gllephosis arachnoidea Gnephosis brevifolia Gllephosis eriocephala N 56 km N Carnarvon Isoetopsis gramillifolia Milluria illtegerrima Myriocephallls gascoynellsis ms endemic Carnarvon area Myriocephallls guerillae N Gascoyne River Myriocephalus IllldllS HW Myriocephalzls pygmaea Pluchea mbelliflora HW S Rocky Pool, Gascoyne River Podolepis lessonii Pogonolepis mllelleriana Pogonolepis stricta N Three Rivers Station, Gascoyne R. Pseudognaphalium luteoalbllln Rhodanthe chlorocephala subsp. splendia Rhodanthe margarethae 194 N. Gibson, G.,. Keighery, M.N. Lyons

Aquatics Taxon Habitat Endemics Range Geographic ends limits

Rhodanthe stricta Senecio glossanthus * Sonchus oleracells Streptoglossa tenuiflora HW S Boolathana Trichanthodium skirrophorum * Urospermum picroides

Family: Avicenniaceae EAq Avicennia marina

Family: Boraginaceae Heliotropium curassavicllm EW Plagiobothrys plurisepalus N Coolcalalaya

Family: Brassicaceae * Brassica tournefortii * Coronoplls didymus Lepidium oxytrichum Lepidium pseudoruderale Lepidillm rotundum Menkea australis Menkea villosllla EW * Sisymbrillm erysimoides Stenopetalum pedicellare Stenopetalum sphaerocarpllm

Family: Caesalpiniaceae Senna sp. Austin (A. Strid. 20210)

Family: Callitrichaceae Aq * Callitriche stagnalis

Family: Campanulaceae Wahlenbergia preissii N Coolcalalaya Wahlenbergia tllmidifructa

Family: Caryophyllaceae Gypsophila australis * Polycarpon tetraphyllum * Spergularia rubra

Family: Casuarinaceae Casllarina obesa R/F

Family: Centrolepidaceae Centrolepis eremica Centrolepis hllmillima N Wooramel River

Family: Chenopodiaceae Atriplex holocarpa N Lake MacLeod Atriplex lindleyi subsp. inflata N Flats near Gascoyne River Atriplex paludosa subsp. moquiniana Atriplex semilllnaris S Atriplex vesicaria Chenopodium auricomum EW Chenopodium cristatum Dysphania glomulifera subsp. eremaea EW Dysphania platycarpa EW Dysphania rhadinostachya S Halosarcia halocnemoides S Halosarcia indica S Halosarcia pruinosa S Halosarcia pterygosperma subsp. denticulata S Wetland flora and vegetation 195

Aquatics Taxon Habitat Endemics Range Geographic ends limits

Mairealla aphylla Maircalla opposilifolia N Yaringa Mairealla slip/tata S Tamala Osteocarpllm acroptenzm var. acropten/m N Boolathana Salsola kali Sarcocornia qllillqllej70ra R/F Sarcocornia sp. Sclerolaella diaca Iltha Sclerolaena recllrvicllspis Tecticornia vernzcosa HW Threlkeldia diffllsa

Family: Clusiaceae Hypericllm gramille/lm HyperiCllm japoniclml

Family: Colchicaceae WlIrmbea mllrchisoniana EW N Coolcalalaya

Family: Crassulaceae Crassllla colorata Crassllla exserta Aq Crassllla natans Crassllla pedicellosa

Family: Cymodoceaceae Halodllle /minervis EW

Family: Cyperaceae EAq Balmlea articlllata R Cyperlls aft. c/mninghamii Cyperlls bifax R/F; s Talisker Cyperlls gymnocalllos R * Cypems hamllloslls Cypems rigidelllls EW Cypems sqllarroslls EW s 50 km S Carnarvon Cyperlls vaginatlls R EAq Eleoclzaris aCllta N Carnarvon EAq Eleoclzaris genic/data S Mooka Creek, range extension Eleoclzaris pallens EW lsolepis congma N Kennedy Range, Boolathana lsolepis cyperoides N Coolcalalaya, range extension Lipocarpha microcephala Schoenoplectlls laterij70ms EW S Cardilya Ck, Carey Downs, range extension EAq SchoCl/oplectlls sllblllatlls S Murchison River Schoen liS elegans SchoCl/lls Iwmilis N Wooramel River

Family: Droseraceae Drosera illdica EW

Family: Elatinaceae Bergia iIl/riclllata EW Bergia perellllis subsp. obtllsijr)/ia EW Range extension Aq [lat/Ile v:nlho'{o/,'tes EW Range extension

Family: Frankeniaceae Frankellia cillerea Frankema laxij70ra N Talisker Frankenia pallciflora Family: Gentianaceae Centallrillm sp/CIltmn 196 N. Gibson, G.]. Keighery, M.N. Lyons

Aquatics Taxon Habitat Endemics Range Geographic ends limits

* Family: Geraniaceae Erodium cicutarium

Family: Goodeniaceae Goodenia berardiana Goodenia corynocarpa Goodenia maideniana Goodenia neogoodenia Range extension Goodenia pinnatifida Vel/eia hispida

Family: Haloragaceae Haloragis trigonocarpa Myriophyl/um decussatum N Doorawarrah

Family: Hypoxidaceae Hypoxis occidentalis N Kalbarri

Family: Isoetaceae Isoetes muelleri Isoetes sp.

Family: ]uncaceae funcus aridicola N Cardilya Creek, Carey Downs * funcus bufonius funcus kraussii Family: Juncaginaceae Triglochin calcitrapum N Kennedy Range Triglochin centrocarpum Triglochin minutissimum N Boolathana Triglochin mucronatum N Dirk Hartog Island

Family: Lythraceae Ammannia baccifera S Wooramel River, range extension Ammannia multiflora S Cardilya Creek, Carey Downs, range extension Lythrum sp.Towrana (RJC 2183) EW endemic" Camarvon - Gascoyne Junction area Rotala diandra S Cardilya Creek, Carey Downs, range extension

Family: Malvaceae Abutilon sp. Lawrencia glomerata Lawrencia viridigrisea * Malva parviflora

Family: Marsileaceae Marsilea angustifolia EW Marsilea drummondii EW Marsilea sp.

Family: Mimosaceae Acacia anettra

Family: Myoporaceae Eremophila oldfieldii subsp. oldfieldii Myoporum acuminatum

Family: Myrtaceae Eucalyptus camaldulensis var. obtusa R/F Wetland flora and vegetation 197

Aquatics Taxon Habitat Endemics Range Geographic ends limits

El/calyptl/s coolabah R/F El/Calt/ptlls victrix R/F Melalel/ca glomerata R/F Melalel/ca IflIcadendra R/F Melalel/ca linophylla R/F Mclalel/ca l/ncinata Verticordia forrestii

Family: Najadaceae Aq Najas marina

Family: Nyctaginaceae Boerhavia coccinea

Family: Ophioglossaceae Ophioglossllm 11Isitanicllm

Family: Orchidaceae Prasophyllllm gracile N Tamala

Family: Papilionaceae Lotl/s allstralis Lotlls crllentlls Medicago polymorpha Medicago tnl11catllla MlIelleranthlls trifoliolatlls Rhynchosia minima 5 Cardilya Creek, Carey Downs Swainsona pterostylis Swainsona sp. Trigonella sllavissima

Family: Plantaginaceae Plantago drllmmondii

Family: Plumbaginaceae MlIellerolimon salicorniacellm 5

Family: Poaceae Agrostis sp. Aira caryophyllea Aristida holathera Al/strostipa tricllOphylla Avena barbata Bromlls arenaril/s Cenchrl/s ciliaris Diplaclme mllclleri Ehrharta longiflora Eragrostis allstralasica EW Eragrostis basedowii Eragrostis cllmingii Eragrostis dielsii Eragrostis leptocarpa Eragrostis pergracilis Eragrostis tenellllla Eragrostis xerophila Eriaclme aristidea Enaclme j7acCliia Eriaclme ovata Eriaclme plllchella Eriochloa procera 5 Cardilya Creek, Carey Downs ElIlalia allrea HordCllm leporinllm Iseilema eremaellm N Minilya 198 N. Gibson, G.J. Keighery, M.N. Lyons

Aquatics Taxon Habitat Endemics Range Geographic ends limits

Paractaenum novae-hollandiae ,. Parapholis incurva N Tamala, range extension ,. Pentaschistis airoides ,. Phalaris minor ,. Polypogon monspeliensis Psammagrostis wiseana endemic Lake MacLeod area ,. Rostraria pumila Setaria dielsii Sporobolus mitchellii Tragus australianus

Family: Polygonaceae ,. Emex australis Muehlenbeckia cunninghamii EW Rumex erystallinus EW Lyndon R. and McNeill Claypan, Gascoyne R. in WA ,. Rumex vesicarius

Family: Portulacaceae Calandrinia corrigioloides Calandrinia eremaea Calandrinia granulifera Calandrinia polyandra Calandrinia ptychosperma Calandrinia pumila Calandrinia sp. Coolcalalaya (G}K and NG 698) EW endemic Coolcalalaya area Calandrinia stagnensis Portulaca oleracea

Family: Potamogetonaceae Aq Potamogeton pectinatus Aq Ruppia megacarpa Aq Ruppia polycarpa Aq Ruppia tuberosa N Shark Bay

Family: Primulaceae ,. Anagallis arvensis Samolus junceus Samolus repens

Family: Ranunculaceae Ranunculus pumilio N Boolathana, range extension Ranunculus sessiliflorus N Eurardy

Family: Rubiaceae Oldenlandia galioides EW S Cardilya Creek, Carney Downs, range extension Synaptantha tillaeacea

Family: Sapindaceae Dodonaea petiolaris

Family: Scrophulariaceae Elacholoma hornii Range extension Glossostigma diandmm EW N Kennedy Range Glossostigma drummondii EW N Doorawarrah Mimulus gracilis EW S McNeill claypan, Gascoyne River, range extension Peplidium aithocheilum EW Peplidium muelleri EW Peplidium sp. C (N.T. Burbidge and A. Kain 8152) EW Wetland flora and vegetation 199

Aquatics Taxon Habitat Endemics Range Geographic ends limits

Stemodia flomlenta EW Stemodia grossa Stemodia viscosa

Family: Solanaceae Nicotiana occidentalis Solanllm lasiophyllllm

Family: Thymelaeaceae Pimelea trichostachya Range extension

Family: Tiliaceae Corchoms walcottii

Family: Typhaceae EAg Typha domingensis

Family: Urticaceae Parietaria debilis

Family: Zygophyllaceae Zygophyllllm ammophilllm

A A recent (August 2000) taxonomic review of the genus Lythrum in WA (Lepschi, 2000, Nlltysia 13: 273-282) has included this taxon in Lytlmtnl wilsonii which is widespread across the arid and semi-arid regions of New South Wales, the Northern Territory, Queensland and South Australia.