Aquatic Invertebrates and Waterbirds of Wetlands and Rivers of the Southern Carnarvon Basin, Western Australia

Recofdsof the Western Austtalian Museum Supplement No. 61: 217-265 (2000\.

Aquatic invertebrates and waterbirds of wetlands and rivers of the southern Carnarvon Basin, Western Australia

S.A. Halsel, R.J. Shiel'?,A.W. Storey3, D.H.D. Edward3, I. Lansbuql, D.f. Cale3and M.S. Harwef rDepartment of Conservationand Land Maiagement, Wildlife ResearchCentre, PO Box 51, Wanneroo,I{estern Aushalia 6946,Australia '?CRC for FreshwaterEcology, Murray-Darling FreshwaterResearch Centre, PO Box 921,Albury, New South Wales2640, Australia 3Depadmentof Zoology, The University of Westein Australia, Nedlands,Western Australia 6907,Australia aHope EntomologicalCollections, Oxford University Museum, ParksRoad, Oxford OX1 3PW United Kingdom sDepadmentof Teirestdal Invertebrates,Westem Australian Museurn, FrancisStreet, Perth, Western Australia 6000,Australia

Abstract - Fifly-si\ sites, representing 53 wetlands, were suiveyed in the southem Camarvon Basinin 1994and 1995with the aim of documentinethe waterbird and aquatic inverteblate fauna of the region. Most sites ivere surveyed in both winter and summer, although some contained water only one occasion.Altogether 57 waterbird specieswete recorded, with 29 292 waterbirds of 25 specieson Lake Macleod in October 1994.River pools were shown to be relatively important for waterbirds, while many ?reshwater .lavpanswere little used. Ai least492 speciesof aquatic invertebratewere collected.The invedebrate fauna was charactedzedby the low ftequency with which taxa occurred: a third of the specieswete collected at i single site on only one occasion. Patternsof occurence were not strongly seasonal.Many undescribedspecies were found and many range extensionswere recotded, reflecting lick of previous aquatic inveltebrate work in the region. The level of regional endemicity could not be assessedadequately, although it is probably comparativelylow. In terms of their invertebratefauna, Iive types of wetlands were distinguished:river pools,_rock pools and larger flowintstreams; seeps, springs and smaller cteeks;freshwater claypans; birridas; and Lake Macleod. Environmental factors to which invertebrates appeared to respond were ratio of calcium/ alkalinitr lotal dissolvedsolids, turbidity, colour, flow, longitude and nutrientt although somefactors were inter-corelated. Additional surveyi should find extra species of waterbird and, mote particularly, aquatic invertebrateusing wetlands of the southern Calnarvon Basin. For many invertebrates, occultences are too sparse for eflective protection of specieswithin a nature resetve system and other mechanisms will be required to ensute their conservation. Comparison of site classifications based on waterbird, aquatic invertebrate and Dlant data (Gibsonei al.,2000) showed pattems among sitesidentified using one element of Lhebiota djd not refle.t patternsshown by other element".ihi. .uggests that, until further work hal identified an element that reflects the iviole wetland communitt as many biotic elementsas possibleshould be surveyed.

INTRODUCTION Beginning with the extensive records of Tom The southem Camarvon Basin is located on the Carter, many casual observationshave been made mid-west coastof Westem Australia,in arrarea with of waterbirds in the southern Camarvon Basin and arid or semi-arid climate. The resion contains few a number of small surveysundertaken flohlstone et nature reservesor nafiondlparkq althoughShark al., 2000).Despite waterbirds probably being the Bay, in the centre of the regiorL is a World Heritage most studied faunal group in the Basin, these area (Anonymous 1995).In 1994,Environment historical data reveal only broad patterns of Australia commissioned a comprehensive biological waterbird occurrence and some of the more survey of the Basin, including its yegetation, important wetlands for waterbirds. They do not marrunals, birds, amphibians, tenestrial arthr:opods allow easy comparison of different wetlands. For and aquaticfauna (Burbidgeef al.,2000). aquatic invertebrates, even broad patterns of S.A. Hal6e,R.J. Shiel, A.W. Storey,D.H.D. Edward, I. Lansbury, D.J. Cale, M.S. Hawey

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Figure I Map of the survey area and boundary of the southern Carnarvon Basir; showing the distribution of the aquaticsampling sitesand placesmentioned in text. SeeAppendix 1 {or site codes. Aquatic fauna of the southern Carnarvon Basin 279 occunenceare unknown becausestudies have been Table 1 Winter and summer rainfall (mm) associated restricted to a few collections of particular groups with sampling periods in the southern for taxonomic purposes (e.g. Lansbury, 1969; De Catnarvon Basin (see Table 2 for sampling dates). Deckker, 1978). Long-term median values are shown in parentheses (data from Bureau of This lack of biological data is hampering Meteorology). assessmentof conservation Driorities. Other than parts of Shark Bay, only two wetlands in the Wintel Summer Wintel southern Carnarvon Basin were identified as 1994 7995 1995 nationally significart in a recent review, namely Lake '1.996) larnarvon /J 162) 56 (11) 123(82) Macleod and McNeil Claypan (Lane et al., Gascoynelunction 39 (47) 109(2s) 56 (47) (Figure 1). Both are large claypans.Lake Macleod Kalbarri 786 (204) 2 (5) 773(204) receives marine groundwater, which upwells 20 km irLland in sinlholes on the western side of the lake, as well as swface inflow via the Minilva River. smaller creeksand sheetflow from the GasioyneRiver aJter Wooramel, Gascoyne and Minilya Rivers. The very hear.y rain. It has been shoum to support very southernmost part of the Basin has a semi-arid high numbers of waterbirds, especially migratory climate with predominantly winter rainfall (Gentilli, shorebirds (Smith and Johnstone, 1985;Jaensch and 1972). North of Shark Bay, tropical systems Vervest, 1990) as well as containing some of the few influence rainfall patterns and result in significant inland stands of mangroves in Westem Australia. summer precipitation in more inland areas,which The values of McNeil Claypan are less well are arid. Median annual rainfall is 379 mm at documented but it fills when the Gascoyne River Kalbarri, 230 mm at Yallalong statio& 206 mm at floods and contains one of the more extensive Camarvon and 190mrn at GascoyneJunction (data MuehlenbaeckiaI Sesban ia shrublands in north-westem on rainfall preceding the sampling periods are Australia, as well as supporting a diversity of given in Table 1 and Figure 2) (see Figure l for waterbirds, includhg freshwater crakes, rails arrd locations). shorebirds(Lane et a1.,1996). Major rivers in the Basin flow intermittently, with The southern Camarvon Basin contains a much significant dry intervals being more common in the greaterarray of wetland types than are represented north. Although flow patterns had not been studied by Lake Macleod and McNeil Claypan. The aims long (8-25 years), data collected prior to 1982 of this suryey were (1) to inventory the waterbird suggested there was significant flow in the and aquatic invertebrate fauna of the Basin" (2) to Murchison River nearly every year (flood events > identify the maior wetland types and faunal 90 per cent of years)(Anonymous 1984).In contrast assemblagesoccurring in the regiory (3) to relatethe flood events occurred on the Minilya River less thar occurrencesof thesefaunal assemblagesto physical every secondyear. Betweenflood events,riverbeds ald chemicalcharacteristics of waterbodies,and (4) dry and surfacewater recedesto a few pools. to examine whether the biological community at Many sections of river floodplain support each wetland may be characterizedby surveying a extensive networks of intermittently flooded single element of the biota. Fis[ amphibians and claypans, especially on the Gascoyne River. tortoises were not included in the survey, although Intermittently flooded interdunal claypans, often they occur commonly in the Basin. with well-defined lunettes, occur throughout the Implications of the data collected, in terms of Basir; although they are less common in the south- adequacy of existing nature reserves and east. There are also claypan-like pools on poorly formulation of a nafure conservation strategy for defined watercoursesthat, it appears,often fill from the Carnarvon BasirLare dealt with bv McKenzie e/ local rain but form part of the regional drainage aI. (2000\. system only after exceptional rainfall events. Examplesinclude Coollilee Pool (C849) and Tirigie Claypan (C856) in the northern part of the study STUDY AREA AND METHODS area (Figure 1). Close to the coast, birridas (evaporite pans) occur in interdulal depressions, Study Area especially around Shark Bay. Most of the birridas The area in which wetlands were surveyed contain gypsum and, although they may dry extended from the Murchison River in the south to intermittently, anecdotal information suggests their the Minilya River in the north and inland to water levels show subdued response to oceanic Gasco)me Junction, covering all but the northern tides. Lake Macleod, which was open to the sea portion of the Carnarvon Basin (Figure 1). The during the Last Interglacial,is arr example of a very physical environrnentis describedbyWytwoll et aI. large birrida. (2000).Essentially, the Basin containslow gradient Surface water is scarcein the Carnarvon Basin, alluvial plains that are traversed by the Murchisory except after major rainfall events. Permanently 220 S.A. Halse, R.J.Shiel, A.W. Storey,D.H.D. Edwatd, L Lansbury, D.J. Cale, M.S. Harvey

Highest on record

Decil;1 Oecilcs 8&9

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A B

Figure 2 RainIall decilesin WesternAustralia in winter 1994and summer 1995.A. Winter 1994.B. Summer 1995.The arrow indicatesthe southernCamarvon Basin.Data from Bureau of Meterology. flowing water occurs only where there is the sites were selected during recormaissancein groundwater dischargeat seepsor springs,usually summer 1994 with the aims of (1) including in the headwaters of small creeks associatedwith representatives of all common wetland t)?es in the the Kennedy and Carrandibby Rangesin the eastern region, (2) obtaining a geographic spread of part of the study area. There are no naturally wetlands, and (3) as far as possible, sampling permanent freshwater swamps or claypans; the wetlands that were in a natural condition. The only sites with moderately deep permanent water swamp near Hamelin Station homestead (C820) are river pools, which are often formed by ro&y was included to examine conservation value of amphitheatres. There are also a few uncapped artificial wetlands. artesian bores, which flow constantly, and have Geographicalcoordinates of each site were formed small artificial swamps. The bore feeding determined using a handheld GPS. Most of the the swarnp near Hamelin Station homestead (C820), wetlands sampled were comparatively small the example of this wetland type chosen for survey, claypans or swamps (< 5 ha), or were river pools, was subsequentlycapped and the swamp has dried. and the sampling area associated with the site effectively covered the whole wetland or river pool. Site Selection and Sarnpling When wetlands were large, the site constituted only A total of 56 sites were sampled acrossthe study a sampling point within the wetland. This was seen area, representing 53 wetlands. Lake Macleod most clearly at Lake Macleod, which has an areaof containedfour sites(Figure 1).Fifty-three sites were 150,000ha when fully flooded, and contained four sampled in wnter, 37 in summer, arrd 36 in both sampling sites(C876-79) (Figwe 7). winter arrd summer (Table 2, Appendix 1). Most of On each visit to a site that contained water,

Table 2 Dates of reconnaissanceand sampling of wetlands in the southem Carnarvon Basin. See Appendix 1 for exolaration of site codes.

Sampling period Season No, of sites Comments

24-30 April1994 reconnalssance Most sitesnorth of GascoyneRiver not visited 17-30 August 1994 wrnter 11-13 October wlnter 2 C878,CB79 and aerial survey of Lake Macleod 15-27 March 1995 summet 37 25-27 Jl.ly 1995 winter 4 cB27b, CB30 CB51,CB54a Aquaticfauna of the southemCarnarvon Basin 227

inJomation was collectedabout waterbirds. aquatic 1989).Samples for nutrient analyseswere usually invertebratesard the wetlandenvironment {mostlv passed through a 0.45 pm filter in the field and water cher stry). Vegetationdata were collected stored at ambient temperafure unless water was separately (Gibson ef al., 2000). Binoculars or a very turbid, when they were frozen in the field telescopewere used to identify waterbird species. without filtering and ultracentrifuged in the Where available, waterbird lists from the laboratory.Turbid samplesfor measurementof TDS recomaissance,as well as two sampling trips, were were passed through a 0.2 pm filter in the included in analyses to provide as much laboratory (rather than the standard 0.45pm) prior information as possibleabout waterbird use of sites. to evaporation to minimize contamination by fine An aerial survey was made of Lake Macleod on 12 particulate matter. At least 200 ml of water was October 1994 to document waterbird use of the passedthrough a glassfibre filter paper in the field whole lake during a time when palaearctic to obtain a chlorophyll sample for analysis.MgCO. shorebirdswere likely to be present. was added to the algal residueto stabilizi Two aquatic invertebrate sampleswere collected chlorophyll, which was frozen until amount of at eachsite using D-framed pondnets with 250 and chlorophyll present was determined in the 50 pm sized meshes.The 250 pm sarnple was laboratory (APHA, 1989). collectedby 50 m of rigorous sweepingin all Data on ionic composition were converted to identifiable microhabitats < 1 m deep at the site. milliequivalents L 1 and, to characterize water including benthicsediment. submerged and chemically, ratios of calcium to bicarbonate and emergentmacrophytes, coarse organic material and carbonate(termed calcium/ alkalinity), calcium and open water, over a distanceof up to 200 m. The 50 magnesium to chloride, and calcium to sulphate pm samplewas collectedwith 50 m of lessvigorous were calculated.In total 14 environmentalvariables sweeping and included all habitats other than in winter, and 15 in summer, were used in analvses benthos.The 250 pm sample was preservedin 70% (Appendix 2). alcohol; the 50 lrn sample was preserved in 1-2% formaldehyde. Samples were sorted in the Analyses laboratory under a dissecting microscope and representative specimens of each taxon were Waterbirds retained for identification. Most animals were Siteswere grouped accordingto similarity of their 'rnorphospecies' identified to speciesor leyel but waterbird fauna using the PATN analysis package Nematoda were identified onlv to phvlum and (Belbin, 1993) and presence/ absence waterbird Polychaetaand some Crustaceawit-h marine data. Czekanowski's coefficient was used to affinities were identified only to family. Protozoa measure degree of associationbetween sites after were very much under-sampled, although they specieswith a single occurrenceand sites with a were identified when collected.Vouchers of most single species were removed from the dataset. taxa have been retained at the Wildlife Research Under-estimatedassociation values (>0.9)were re- Centre (Department of Conservation and Land recalculatedusing the ShortestPath option in 'Unweighted Management),Western Australian Museum or PATN and the Pair-Group Mean Murray-Darling Freshwater Research Centre. Average' fusion method, with B : -0.1,was used to Names of Chironomidaefollow Cranston(1994). group sites (Sneath and Sokal, 1973). Waterbird Marimum waterdepth at eachsjle wasestimafed species were classified into groups with similar eachsampling occasionand siteswere assignedto a patterns of occurrence using the Two-Step flow category (1, lentic; 2, seasonalriver, not coefficient(Austin and Belbin, 1982)and UPGMA. flowing when sampled; 3, spring-fed flowing or The discretenessof wetland groups produced by seeping water; 4, flowing river {ed by catchment the classifications were examined by ordination run-off). At each site, water samples and using'Semi-Strong Hybrid Multidimensional measurementswere taken within the area sampled Scaling'(Belbirl 1991). for invertebrates.Conductivitv and oH were measuredl5 cm below the watersurface usins fPS Ena ir onme nt al aar i abl e s ModelsLCSI and LC80Ameters; dissolved oiygen Sites were also grouped according to their was measurednear the surfaceand at the bottom of environmental characteristics.This was done the water column with a WTW OXI96 meter and separatelyfor sitessampled in winter and summer. the two readings were averaged. Water samples All variables were range-standardized(each value were collected about 15 cm below the surface for of a variable was divided by the maximum value subsequentmeasurement of total dissolved solids recorded for that variable) after those with strongly (TDS), ionic composition (including silica in skeweddistributions had been loe-Lra nsformid. summer, total soluble persulphate nitrogen and The Bray-Curtisassociation measure ard UPCMA phosphorus, chlorophyll, colour and turbidity in the fusion method, with B: -0.1, were used after laboratory using standard techniques (APHA, association measures >0.95 were recalculated. 222 S.A. Halse, R.J.Shiel, A.W. Storet D.H.D. Edwar4 I. Lansbury, D.J. Cale, M.S. Harvey

Differences in environmental characteristicsof classifications were examined by ordination using wetland groups identified using aquatic SSH. Maximum linear correlations between invertebrate data were examined by one-way transformed environmental variables arrd vectors in ANOVA" using the SAS statistical package (SAS ordination space were calculated using the PCC Institute, 1989), atter variables with skewed option in PATN after a varimax rotation of the distributions were 1og-transformed. Student- ordination axes (Belbin, 1993). Significance of Newman-Keuls tests were used to identify the correlations was determined by Monte Carlo testing groups contdbuting to significant overall variation (1000 randomisations). in environmentalvariables. Site classifications based on invertebrates and on environmental variables were compared using a Aqunticinrertebrqtes modified Rand statistic (Hubert and Arabie, 1985) Sites were grouped on the basis of their (sites CB25 and CBo2c were omitted from the invertebrate fauna using presence/ absence data winter comparison because of incomplete data). Site and the same methods as for waterbirds. Separate groupings based on winter and summer classificationswere derived {or winter and summer. invertebrate datasets were also compared after Association measures>0.95 were recalculated.The datasets for each season were reduced to a common discretenessof wetland groups produced by the set of sites.

0.0000 0.4020 0.8040 \,2460 1.6080 2 -0100 I l I I I cB04 1) cB05 2)_ | cB20 8)_l_ | B06a 3)_ I I 893d 34)_l_l_ I 14)_ I I cB44 15)_l l_l_

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cB09 5)_ cB15 5)_l_ 829a 10) '7)_ t_ cB15 cB?8 1 I I I 0.0000 0.4420 0.8040 r.2460 1.6080 2.olaa

Figure 3 ClassificationoI wetland sitesin the southem Carnarvon Basininto sevengroups (B1-7)based on waterbird use.See Appendix 1 for site codes. Aquaticfauna of the southemCarnarvon Basin 223

RESULTS area, occurring in all site groupings, although concentrated at sites of groups B1-3. Group BIII Waterbirds speciesoccurred mostly at the more species-rich A total of 57 speciesof waterbird were recorded and deepersites (81 and 2). Group BIV consistedof during surveys with the most commonly occurring the Banded Lapwing and Black-tailed Native-hen, being White-facedHeron (28records of one or more birds at a site on a sampling date) Grey Teal (27), White-necked Heron (16), Pacific Black Duck (14), iir AplerdirI ll]frtr codrl Black Swan (12), Australian Wood Duck (12), Hoary-headed Grebe (12) and Black-fronted Dotterel (12)(Appendix 3). All speciesrecorded are widespread outside the study area (BTakerset aI., ll ^ A 1.984). Lake Macleod (C876-79)was clearly the wetland €sfi t supporting greatest numbers of species and .f; nF individualsduring the surveys(29 292birds o{ 25 speciesin October 1994, including birds counted oo during an aerial survey of the whole lake). Moderate nunbers of birds were recorded on all visits to the bore swamp on HameLinStation (C820) (100 birds of 8 speciesin March 7994;95 of 12 in August 1994).Lake Julia (CB62b)had 87 birds of 10 speciesin March 1994. Minilya Pool (CB82) and Winnemia Pool (C842) were the river pools with most birds (87birds of 10 speciesand 42 birds of 12 species,respectively, in August 1994).The birrida a a north of Big Lagoon (CB29a) supported 1285 F"ls82] shorebirds of 4 speciesin August 1994,making it alo ^ the birrida with highest waterbird numbers. r oqg: tr lA83l AAA oe+l Waterbirds were not recorded at 11 sites and at "9, Errrrr I^ another 10 sitesonly one speciesoccurred. These 21 {a oes] sites were not included in multivariate analysis r86 (Appendix 3). ] ^87 ) Based on waterbird usage,seven groups of sites were identilied (Figure 3). Only sites in groups 81 and 2 supportedmany species,with averagespecies richness of 10.911.3and 18.314.9,respectively. 83 sites showed a considerablerange in species richness, with 12 and 10 species,respectively, at CB62band CB82but a mean of only 3.910.5at other sites. No site in groups B4-7 had more than four species (Appendix 3). In general terms, 81 sites were large river pools or vegetated swamps. 82 consistedof the three wetter Lake Macleod sites, 83 siteswere freshwater claypans,often with some emergent shrub or herbaceousvegetatiory as well as the river pool site CB82.Groups 84-6 contained turbid freshwater claypans.87 consistedof birridas and the driest Lake Macleod site (C878). Groups showed moderately clear separationin ordination space(Figure 4). Sevengroups of waterbirds were identified, based on their pattern of occurrence(Figure 5). Groups BI- III containedthe more commonl; occurringspecies. Species groups showed only loose relationships Figure 4 Ordination of wetland sites in the southern with site groupings but BI speciesoccurred most Carnarvon Basin based on waterbird use. consistently at river pools and vegetated swamps Wetland gloups lrom Figure 3 are (B1 sites)and were absentfrom saline sites.Group superimposed on ordination (thlee Bll specieshad the widest occurrencein the study dimensions,stress=0.08). 224 S.A. Halse, R.J.Shiel, A.W. Storey,D.H.D. Edward, I. Lansbury, D.J. Cale, M.S. Hawey two relatively terrestrial waterbirds with limited same speciesbeing identified at different levels of occurrencein the study area. Groups BV afld VI taxonomicresolution). Fifty per cent of specieswere consisted of shorebirds, with BV species being microinvertebrates,with many other species(e.g. restdcted to saline Lake Macleod arrd birrida sites nematodes, oligochaetes,watermites) also being (82 and 7), whereas some BVI species were very small. The major componentsof farma richness restrictedto freshwater and othersoccurred in fresh were rotifers (14% of species),dipterans (14%, of as well as salt water and wete present in most site which well over half were chironomids), groupings. Group BVII contained a mixture of cladocerans(14%) beetles (11%), ostracods (10%) speciesthat occurred at Lake Macleod; some were and copepods (9%). The most ftequently recorded roosting in mangroves, others occurred on the water taxa were the chironomid species complex and somewere aedal. Procladiusspp. CBT1 (62%of samples),beetle Eretes australis(43o/") and, chironomid Tnnytnrsussp. CBC3 Aquatic invertebrate speciesrichness ,41,%\. At least 492 speciesof aquatic invertebrate were Of the 518 taxa in Appendix 4, 423 were collected collected during the surveys (Appendix 4 contains during winter,327 in summer and 232 were 518 taxa but Drobablv some of theseresult from the common to both seasons.The major cause of

81 B2 1ll 1J4 u) uo ul ccccccc lccc lccccccccccc ccclccc ccciccccc BBBBBBBJBBB I BBBBBBBBBBBBBB I BBBI BBBI BBBBB A0t094417 7 105523685767 Il59 | I47lr59 1012I? 45A6324| 67 9 | 648182263 8s I 863| 59sI 883| 95968 ad bcbba ca aadef a I Australasian Grebe Australian shelduck t1 Ausirralian !'lood Duck t.- Black-f ronted Dotterel BI Pacific B]ack Duck ..1. Eur.asian coot: GreaE Egret I Straw-necked lbis "l Yellow-bi11ed spoonbill

Grey TeaI --"--t-.-1...---.1."-11 White-faced Heron -.--t-.-t..--...---" t" BII Hoary-headealGrebe I Pink-eared Drick I white-necked Heron .-.1

Australian Pelican Darter BIlI Black swan Little Pied Cormorant Little Black cormorant

BIV Bandeal Lapv,/ing t-tl Black tailed Native Hen tt lt..-tl Banded Stilt t--- BV Red-capped Plover curlew Sandpiper t" Red-necked sti-nt t. Conuron Gr:eenshank I-t BVI Red kneeal Dotterel t"l t- Comnon Sandpiper -.t"1 t-- Real-necked Avocel -1-l -ll --l LiLtle Egret Pied. Cormor:ant BVIl silver cull Caspian Tern GreaL crested Grebe

Figure 5 Two-way table of waterbird speciesgroups and wetland site groups in the southem Calnarvon Basin. See Appendix 1 for site codes. Aquatic fauna of the southem Camarvon Basin 225

Table3 Frequency of occurtence of aquatic strongly contracted in preservative remain invertebrate species from the southern identified to genus only and the number of ralge Carnarvon Basin in 90 samples from winter extensions may increase 1994and surnmer 1995. after they have been examined in more detail. About 20 cladoceran No. oI samplesin No. of species species from eastern Australia or the Northern which speciesoccurred Territory (Smirnov and Timms, 1983;Timms, 1988; Frey, 1997)were recorded from Western Australia 1 158 {or the first time. In many casesthe range 120 extensions have 4-6 80 considerable conservation significance; for 6-9 44 example, Celsinotumhypsilophum was previously 10-18 52 19J7 22 >27 10 Table4 Undescribedspecies of aquaticinvetebrate collectedto date only from wetlandsof the southernCamarvon Basin in 1994and 1995. differences in species lists between seasons Rotifera appearedto be the high proportion of taxa collected Keralellasp. nov.la[[. austrdlisgroup) infrequently. A third o{ taxa were recorded only Asplanchxasp. nov. (afI. sieboldi) once (althouShmany individuals may have been Euchlanissp. nov. collected on that occasion) and only 10 species Lecanesp. nov. occurred in more than 30% of samples (Table 3). Anostraca Seasonalpreferences were sometimes evident Branchinellasp. nov. (atf. Iyrifera)1 among more common taxa but they did not prevent Cladocera collection in both seasons;for example, the rotifer Alonine gen.nov. Kerolellaaustrqlis had a strongpreference for winter Alona spp. nov. A-E (13 out of 14 occurrences)but was recorded oncein Biaperturasp. ^ov. summer (Appendix 4). The shield shrimp Tdops Raksp. nov. australiensisaustraliensis showed the opposite Macrolhrixsp. nov. Neothrix (aft. pattern.being found at eight of 37 sitesin iummer sp. nov. superarmata) Il:!/ocryptussp. r.ov. and only two of 53 sites in winter. More taxa Daphniasp. nov. (aff.baabata) exhibited preference for winter than surnmer. Dqphniasp. r\ov. (att.gibba\ Many new species were collected during the Daphniasp. nov. (aff.projecta) sfudy, including four rotifers, one anostracan, 14 Ostracoda cladocerans(including an undescribed genus), 10 Paralimnocytheridgen.nov.l ostracods (including an undescribed genus), four Ampullacyprissp. nov. 469 copepods,one hemipteran ard two beetles (Table ?Ampullacypris sp. nov. 498 4). Three named specieswere recordedin Australia Bennelongiasp. ^ov. 4141 for the first time. The rotlferc Hexorthra brandorffi Cypericercussp. nov, 415 au'LdProales sigmoiclea were previously known from Cypericercussp. \ov. 422 the foothills of the Andes in South America (Koste. Cypericercussp. nov. 4M Heterocyprissp. nov. 489 1978)and from Europeand Canada(De Smet, Mytilocyprissp. nov. 426 1996),respectively, and the harpacticoid copepod Zonocypris sp. nov. 4662 Robertsoniamourel had been collectedonly from the Brazilian type locality (Nogueira, 1961). The Calanoida Calamoeciahalsei] oligochaeteNals sp. CB1 probably also representsa new record for Australia of a cosmopolitanspecies Cyclopoida (A.M. Pinder, pers. comm.) and coliections of the MesocycLtpssp. nov. ostracodZonocyprls sp. nov. 466 representthe first Neocyclopspetooskiia time this predorninantly African genus has been Harpacticoida found in Australia. Identification of the cladocerans Amondnriasp. nov. Daphniasp. nov. (aff. barbata)and D. sp. nov. (aff. Hemiptera glbla) requires DNA-typing but both are similar to PIeasp. nov. describedAfrican species(C. Wilsor; pers. comm.). Coleoptera Large range extensions within Australia were Parostergp. nov. recorded for many species. At least 27 rotifers Tiporussp. nov. previously known from easternAustralia (e.g.Shiel and Koste,1979) (Koste r Common in Basin or the Northern Terrifory '?First and Shiel,1983) were recorded in Westem record oI genusin Australia Aushalia 3Describedby Bayly (1998) for the first time. Severalother rotifer taxa that were aDesdibedby De Laurcntrlset al. (1.997) 226 S.A. Halse, R.J.Shiel, A.W. Storey,D.H.D. Edwar4 I. Lansbury, D.f. Cale,M.S. Hawey known only from the Paroo River region of New records from the Carnarvon Basin extend the range South Wales (Frey, 1991). Occurrence in the of the anostracanBronchinellq probiscidn 2000 km Carnarvon Basin of the calanoid copepod south-westwards(see Geddes, 1981). Menke (1960) Eudiaptomuslumholtzi, common in northern recorded the belostomatid hemipteran Lethocerus Australia lncluding the Kimberley (Timms and distictifemuras occurring only in easternAustralia Morton, 1988),is a 1200km southwards extension and its collection in this study representsa large of the western range of the species. Similarly, range exrensron.

a .0'762 A.2528 0- 5059 4.7824 0.9590 I I cB04 cB05 cB20 B06a cB30

EW1 cB34

89 3d cB4 4 B93a 893f -t 7l CB42 t__ l ( 31) B93c I 48) I EW2 | 32) I B67b ( 34) I -t

cB09 CB15 ?) | cB16 8)-ll- B09a EW3 CB76 cB1'/ 42) cB78 41)_ |

EW4 B05b B21b 10) t_ ( 14) CB36 ( 2r) I CB51 | 22) ( 23) I cB3 8 ( 16) EW5 CB43 ( 19)__l_t_l \ 26) I \ 2'-7) I 30)r_i i B75a ( 38)_l_t_tl B70b ( 36) ,iil CB73 -t-t-t-l I EW6 Bs4a 24t I I cB56 25)_ I B58d 2e)_l_ I 858c 2ar I I l EW7 CB68 35) l_L I 3e)_ I I 4o)_l_l _ I CB82 45)_ -l I I o.0't62 0.2528 a .4293 0.6059 D_'/A24 0.959 Figwe 6 Classificationof wetland sitesin the southernCarnarvon Basininto sevengroups (EW1-7)according to their environmentalcharacteristics in winter (sitesCB25 and CB62cwere excluded).See Appendix 1 for site codes. Aquaticfauna oI the southernCarnarvon Basin 227

Male ostracodsthat morphologically appearedto through-flow during the survey and it is likely they be Sarscypridopsisaculeato were collected from were ecologically more similar to claypans than Minilya Pool (C882) both times it was sampled. If rivers. Sites in IW4 were more coloured, turbid this identification is correct, it is the first time males claypans with high nutrient levels and intermediate of this common, cosmopolitan specieshave been numbers of species (27.8t2.4). One 'river pool' recorded (seeDe Deckker,1981). (Bulgra Pool CB70b) was in this group but physiognomically it resembleda claypan.IW5 sites Invertebrate communities and environmental were more coloured and turbid claypanswith high Yariables nutrient levels and receding water levels, sometimes being close to dry. These claypans Winter containedfewest species(18.211.9). Group IW6 sites Based on their environmental characteristicsin were birridas arrd contained Iew species(11.0t1.5) winter, seven groups of sites were identified (Figure while IW7 contained the four Lake Macleod sites, 6). These were large river and rock pools (EW1) which averaged 16.010.9species. Sites in the latter small creeks and seeps (EW2), saline sites with a two groups were saline with high ratios of calcium/ marine connection (birridas and Lake Macleod, alkalinity (Table5). EW3) two very fresh claypars in the south-eastem Despite some superficial similarities in site part of the surveyed area (CB06b, CB27b: EW4), classifications based on invertebrates and highly turbid claypans with elevated TDS values environmental variables, concordance between (EW5), the single crab-hole swamp sampled (CB54a: them was poor. Using seven groupings, o y 26 of EW6) and lessturbid claypans(EW7). the 51 sites in the two classifications fell into the Seven groups of wetland sites were also identified same groups and the Hubert / Arabie Rand statistic on the basis of their use by invertebratesin winter was 0.3435(1.0 indicates identical classificatiory0 (Figure 7). IW1 siteswere river pools, rock pools in indicatestotal dissimilarity). river channels and the larger flowing streams and, Ordination of the wetland sites based on on average, had the highest species richness invertebrate data showed the same relationships $4.512.6 SE).Group IW2 sites were small flowing betweensite groupings as the classification(Figuie streams and seeps,which tended to be brackish 8). Salinesites (IW6 and 7) were strongly separated (Table 5), and supported 28.814.0species. The bore from rivers and claypans. There was minimal swamp at Hamelin Station homestead(C820) also overlap betweenthe three groups of claypans(IW3- belonged to IW2. Group IW3 contained the more 5) but the smaller streams and springs (IW2) did specioseclaypan and swamp sites (38.5+3.4 not separateclearly from larger river pools (IW1). species),which were less turbid and had less Eight variables were significantly correlated with 'river coloured water. Three pools' (Minilya Pool site positions in ordination space, including the C882, Coollilee Pool CB49 and Boolan Pool CB73) ratio of calcium/ alkalinity, TDS, turbidity and were included in this group but none received colour (Table6, Figure 8).

Table 5 Mean values (1SE) of environmental variables in winter for the wetland groups identified by LPGMA cluster analysis based on winter invertebrate data. The significance values of one-way ANOVAS for each variable are shown. **** P<0.0001,.."P<0.001, NS P>0.05.

Variable Wetland group IW1 tW 2 IW3 IW4 IWs IW6 tw7 pH 8.6!0.2 8.0r0.4 8.110.2 7.810.1 8.310.3 8.2X0.2 8.0r0.4 NS DO (% sat.) 1,1,2!6 134123 97!6 97!4 1,09!12 11519 132110 NS Colour (TCU) 77!2 12!4 3151152 879!265 4432!2490 8.2r3.8 8.511.4 Turbidity (NTU) 0.7t0.2 0.310.03 2587!1195 12620!5509 21796]172840.610.1 0.14r0.0s TDS (mg/L) 946\266 5950!1787 776!45 1.52!47 490r105 9420012276739920!2803 Ca / Alkalinity' 1.310.3 1.810.3 0.24!0.06 0.2110.04 0.10r0.0s35r3 8.8r0.4. Ca+Mg/Cl' 1.2!0.4 0.57t0.2 0.8710.3 0.92!0.3 0.42!0.2 0.26!0.02 0.2510.01 NS Ca/ SOn' 3.4+1.6 0.3810.1 3.011.3 2.7a0.9 2.6!7.6 0.5610.06 0.4510.05 NS Nitrogen (mg/L) 0.67X0.1,7 0.64+0.1 1..4!0.4 2.310.5 2.2X0.5 2.1r0.3 0.5410.11 Phosphorus(mg/L) 0.006:t0.0010.002:10.0020.1,7!0.070.41r0.16 0.6810.25 0.0161.002 0.005r0.002 Ctrlorophyll (mgi L) 0.017r0.0080.00910.0030.0510.02 0.0310.02 0.008:!0.0050.00810.0030.01110.009 NS Latitudeb 25.9X0.2 24.9!0.5 25.010.3 25.110.3 24.8!0.1, 26.3!0.2 23.9!0.04 115.0r0.1 71,4.7X0.2 114.2!0.2 114.510.1 1t4.2X0.2 113.5r0.1 113.710.03 Flow' 3.2 1.1 1.0 1.0 1.0 1.0 ' ratio o{ ionic compositions expressed as milliequivalents b Decimal degrees 'Flow category(see text) 224 S.A. Halse, R.J.Shiel, A.W. Storet D.H.D. Edward, I. Lansbury, DJ. Cale, M.S. Harvey

0.3040 a.1892 L-2',744 2,2444 2.'.7344 I I CBO4 1)_ cB05 2')_l- cB30 r2J_ | 1?)_l_l cB34 13)_ cB42 18)_ | | 1Wl, 893d 50) | | 893f 52)_l_l_ cB44 20)_ |

cB62 31)_ _l_ CB93 4',7)_ 48)_l_J,

cB20 ( 9)_ t32)_ll IW2 B67b I rtr,_l_ | 86'7a ( 34)_L _ l B62c | 1-1r_

B05a 3)_ B54a 24)_l_ 82'.7b 10)_ I cB68 t- Ih]3 cB82 46)_1, cB49 2r)_ cB54 23)_l_ CB51 L CB73 38)_l_ll cB56 25)_l ll I B06b 4l cB38 16) I B58b 2'7)_ |

r!t4 B75c 41)_l_ l_ | 14) _ | B58c 28)_ I I B70b I 40)_ l_l_l_ _ I ( 1s)_ | ( 1e)_l_ | rw5 { 26)_ | ( 3o)_l l_ l B58d t 291

cB09 ( 5)_ 829a I Trr I IW6 B09a

cB16 8)l_l_l

CB75 42J, rvtT cB7'l cB?9 cB78 AA\ I

0.3040 4.1892 L-2744 1.'t596 2.2448 .2.7t00

Figure 7 Classification oI wetland sites in the southern Carnarvon Basin into seven groups (IW1-7) according to their invertebrateIauna in winter (site CB25was excluded).See Appendix 1 for site codes.

Based on their pattern of occunence in winter, depauperate drying-phase claypans) although some sixteen groups of invertebrate species were species showed a preference for the two turbid identified. These included groups that were more claypangroups (IW4 arrd5). or lessrestricted to eachof site groups IW1-4 and 6- 7. There were two groups of generalistspecies that Summet occurred commonly in all but the saline sites. There Six groups of wetland sites were identified was no speciesgroup restricted to IWs sites (the according to their environmental characteristics Aquatic fauna of the southen Carnarvon Basin t.ro

Table 6 Sisnificant correlations between environmental vaiiables and the distribution of 51 wetland sites in ordinations based on the winter and surrmer inve*ebrate faunas of the FLOW ***, *., ., sites. P<0.001; P<0.01; P<0.05. OQ . LoNG 0.5 Variable Winter Summer ATDS ctlA 1f.^\Hfu ordination ordination l^ x0 A . o NadAaO-o Calcium/alkalinity 0.9401 0.8347 -0.5 TDS 0.8913 0.7796 . .l o \o ,u"" 0.8647 0.8724 P/COL Colour 0.8518 0.6198 N-tr Flow 0.7544 0.7624 Longitude 0.6745 0.7272 -1.5 -1 {.5 0 Phosphorus 0.6133 0.6797 Axis 1 Nitrogen 0.5129 0.6070 Latitude 0.4286 0.5405

FLOW o flq LONG in summer. The classification showed some tr o differences from that based on winter .v col environmental data, perhaps because it was a r)0 P/TURB .9 ' smaller dataset, but rivers, marine-influenced AA Aoo sites and claypans still constituted the major o a Tos groupings (Figure 9). Concordance of the classifications based on summer environmental /l and invertebrate data (Figure 10) was equivocal, .3 with 25 of 37 sites being placed in the same a groups in both classifications and a Hubert/ -o.5 0 = Arabie Rand statistic 0.5255. Axls 1 Six groups of wetland siteswere identified on the basis of their invertebrate fauna ir summer (Figure 1.5 10). Group 1S1sites were mostly claypans,wiih the '1 exceptlonof two siteson the lower Murchison River LONG (Hardabut Pool CB04 and Bullock Pool CB05),and FLOW had moderate turbidity and colour, and high tr species richness (38.4a2.8).Group IS2 sites were OO claypans in the drying phase with high turbidity, iAo high nutrient levels and moderate colour (Table4). They had lower speciesrichness (19.3t4.8). Group -1 IS3 contained the bore swamp at Hamelin Station homestead (CB20), Birdrong Spring (CB57a) and - 1.5 two small flowing stream sites,which had slightly elevatedTDS and averaged30.2t3.2 species.Group -2 IS4 contained river pools, rock pools and larger flowing stream sites and had high numbers of species(40.0t3.9). Group IS5 sites were saline birridas with very high ratios of calcium/alkalinity Figure 8 Ordination of wetland sites in the southem and few species(8.3t0.7); 156 comprised the Lake Camarvon Basin based on their invertebrate Macleod sites,which were also salinebut had more fauna in winter, showing environmental gradients species(15.3t2.9). in the ordination space (three dimensions,stress=o.17). Fourteen groups of invertebrate species were identified, based on their pattem of occurrenceat sites in summer. There were speciesgroups Wetland types in the southern Carnarpon Basin and restricted to all site groups except IS3 (seepsand important enaironmaltal pErameterc streams),and one group of infrequently occurring Wetland classiJications derived from winter and speciesshowed preferencefor this habitat. Several summer invertebrate data for a comnon set of 34 groups of generalistspecies occurred comnonly at sites showed a high degree of concordance at the all but the salinesites. five group level (Hubert,/Arabie Rand 230 S.A. Halse, R.J.Shiel, A.W. Storey,D.H.D. Edward, I. Lansbury, D.J. Cale, M.S. Hawey

0.0568 0.1930 0.1293 0.4555 0- 6018 0.7380 I tlI I cB04 ( 1)_ I cB0s ( 2) l_ ESl" B2'7a ( 8)_ B27b ( 9r_'_l_ cB82 { 32).-'-=-_- l_ I cBlo ( 11)_ I ( cB34 12)_l___ i cB42(15)tl B9ld ( J6r lr I ErO1^ | rl| ES2 BlSa ( .t4)_--i cBe3cB93 i( 33)_ I I cB62 ( 20)_ B62a | 271 | B67b I 24 )-. l--=-=- l_

ES3 B67a ( 23l_, l

B06b ( 3 )-.- B75a | 2t) l B62b | 22) I cB54 { 1818)_)--'-- .q?1 | ES4 cB38 ( ttl_ | cB49 ( 16)_t6) | | cB5l ( 1?) | | | aEq< B 75b | 2At I qTnhr)qltll I cB09 ( 4)_ cE16 ( 5) I I ( I ES5 cB15 5) | I cB2o { 7)-' _-_-l_ B2'tc ( 10)-.=---.. ----l_ I cB76 l l 29)_ | I ES6 cBttcB'7'7 { 30) I cB?9 ( I 31) -l trtl I 0_ 0558 0.1930 A.3293 0.4655 0.6018 0.?380 Figure 9 Classification of wetland sites in the southern Camarvon Basin into six groups (ES1-6) based on their environmentalcharacteristics in summer.See Appendix 1 for site codes.

statistic=0.7963), with only Hardabut and Bullock alkalinity, TDS, turbidity, colour, flow, longitude, Pools (CB04 and CB05) on the lower Murchison phosphorus and nitrogen were significant both River changing from river pool to claypan groups seasons (Table 6.). Several of these variables were (see Figures 7 and 10). The Murchison River was in rnrer-correlated { t able /). spate in summer and the main river charurel could not be sampled. Backwaters and small pools associdted with the river were sampled initead: DISCUSSION these were likely lo contain faunas with stronqer Clirnatic variation has considerable implications claypan elements than the river channel ald m1y for any attempt to document the fauna oi a region have causedthe changesin classification. such as the southem Camarvon Basin. Apart fiom All classifications based on invertebrate data obvious samp.ling dilliculties if wetlands remain showed the existence of five major wetland dry in low rairfall years, the fauna of many sites groupings: (1) river pools, rock pools and larger differs according to whether they are full or flowing streams, (2) seeps, springs and small partially flooded. A Westem Australian example of flowing streams, (3) claypans, (4) birridas, and (5) aLmostcomplete tumover of invertebrate fauna was Lake Macleod. There were slight differences provided by Lake Gregory, southern Kimberley, between seasons in terms of environmental between 1989 and 1997, as a result of the lake variables that were best related to invertebrate flooding and salinity being dramatically reduced community composition but ratio of calcium/ (HaJseel nl.. 1qq8b). fhe total aquatic invertebrate Aouatic fauna of the southern Camarvon Basin

Q -2110 0 .44'.76 0 .6782 0.9088 1.1394 1.3?00 I I I I I I cB04 ( 1) cB05 | 2) B06b ( 3) cB82 | 32) t_ B2'7a ( 8) -l B70b | 25) IS1 BTsb ( 28) t_t_l B27b ( 9) -l B27c ( 10) L- cB38 { 13) cB51 ( 17) -L- i cB49 ( 16) ,ii cB56 ( 19) ttl B62b \ 22'1 t,l_t_t_

cB54 ( 18) IS2 cB73 ( 26) t_ ( 21)

cB20 ( 7t B62a ( 2r) B6?b ( 24) B67a (

cB30 { 11) B93c { t- B93f t 37) t_ cB34 ( 12) IS4 B93d ( 36) cB42 ( 15) B38a ( 14) CB93 ( l3) _tl B93a ( 34) t_t_t_ cB62 I 20) _l

cB09 4) rs5 cB15 s)_ | cB16 5)_l_l

cB76 29t_ cB79 1r| | 30)_l I I l a.2110 0 .4476 0-6782 0.9088 1-1t94 1-3700

Figure 10 Classification of wetland sites in the southem Carnarvon Basin into six groups (151-6) according to their invertebrate fauna in summer. See Appendix 1 for site codes

fauna of an area such as the southem Camarvon Basin is unlikely to be documented in one year (especially a dry year), nor can the conservation value of individual wetlands be assessedfully. Table 7 Highly significant correlations (P<0.001) Depending on rainJall pattems, the full value of a between environmental variables in winter wetland may be expressed at intervals of many and summer in southern Carnarvon Basin yearsor even decades(see Halse el a1.,1.998a). wetlands.

Variables Waterbirds Winter Summer Waterbird data collected during this survey reflected TDS and calcium/alkalinity 0.795 0.611 what was already known about broad Colour and phosphorus 0.666 0.817 distributional pattems in the southern Camarvon Colour and turbidity 0.561 0.758 Basin (Johnstone et al., 2000). Data from individual Flow and longitude 0.534 sites highlighted the importance of pools in larger Phosphorusand turbidity 0.477 0.624 rivers, rather than freshwater claypans, as TDS and longitude 4.462 -0.497 waterbird habitat. Timms (1997) found turbid Nitrogen and colour 0.452 0.640 Ireshwater claypans in northem New South Wales Nitrogen and turbidity 0.575 were also little used by waierbirds. S.A. Halse, R.J.Shiel, A.W. Storey,D.H.D. Edward, I. Lansbury D.J. Cale, M.S. Harvey

Two groups in the wetland classification waterbtds moving inland from coastalregions after contained most sites with high waterbird rain (Bekle, 1983;Halse et a1.,7992)applies in the conservation value (81, river pools and vegetated arid as well as the temperatezone. swamps; 82, Lake Macleod), although 83 contained Lake Julia (CB62b) and Minilya Pool (CB82), which Aquatic inveltebrates were among the richer sites in the Basin for waterbirds, and 87 contained the pan north of Big Wetlandclassifications and enuironmental aariables Lagoon (CB29a), which clearly had potential to As for waterbirds, some aspects of the aquatic support large numbers of migratory shorebirds. The invertebrate analysis should be used cautiously. survey reinforced the pre-eminent status of Lake The invertebrate surveys were not conducted Macleod as waterbird habitai in the southern during particularly wet years, although Camarvon Camarvon Basin ( Smith and Johnstone, 1985; Lane and eastern parts of the Basin received above- et aL, 1996) but also showed that some species do averagerainfall in summer 1995(Table 1, Figure 2), not utilise the lake and rely on other habitais, and some of the wetlands chosen during especial.ly river pools (Figure 5). Twenty-one sites reconnaissance were dry during all three sampling were excluded from analyses because either no periods. Theseincluded the drier type of 'crabhole' waterbird or only one species was recorded. There swamps/ representativesof which were identified are many possible reasons for waterbirds not being on Carbla and Minilya Stations. Site CB25 on recorded at a site and depauperate sites should not Yaringa Station, the only saline claypan identified be treated as a natural grouping. in the study area that did not have existing marine Although we analysed the waterbird data, results connections,was sampledbut, becauseit was in the should be used cautiously. Sampling effort was low final stages of drying, contained a depauperate, and variable, with a maxirnum of three surveys per unrepresentative fauna and was excluded from site. Extra surveys would have increased the analyses. waterbird list at most sites: for examnle, two Wetland classilicationsderived from invertebrate additional surveys at three sites produced an data gave consistentpattems across seasonswith average of 2.7 exfta species (Table 8). A further five major types of wetland site being recognized complication is that waterbird populations may (1) river pools, rock pools and larger flowing have been unusually low, and thus waterbird use of streams, (2) seeps, springs and small flowing sites unrepresentative, during the summer 1995 streams, (3) cla)?ans, (4) birridas, and (5) Lake sampling period because of widespread, above- Macleod. The claypan group could be further average inland rain (Figure 2). Rainfall in the divided on the basis of turbidity and stage in the Goldfields between Jaauary and March 1995 was drying cycle (Figures7 and 8). Lessturbid claypans ihe highest on record (Bureau of Meteorology, and those where water levels had not receded 1995). Thirieen sites, at which waterbirds were noticeably had higher species richness, although surveyed during summer both in the 1994 some specieswere more or less restricted to highly reconnaissance trip and in 1995, had almost three turbid and drying-phase claypans. Saline pans times more species in 1994 than 1995 (3.8t1.0 z.rs without current marine influence, such as CB25, 1.3t0.4), when most of the summer surveying was may comprisea sixth, uncommon type oI wetland. done. This suggests that the phenomenon of Preyious sfudies of communitv composition in wetlandsof Western Australia have examined much smaller geographic areas and a restricted Table 8 Numbers of waterbird species recorded at range of wetland types. Growns ef aL (1992) arrd three sites in three sampling periods during Davis et sL (1993)classi{ied 40 shallow, permanent the survey (summer 1994to summer 1995)and or seasonal lakes on the Swan Coastal Plain in two subsequentsurveys (winter 1995 and according to their invertebrate communities and summer 1996).Numbers of speciesfirst seen in winter 1995 or summer 1996 are also found two small outlying groups, one of which was shown. SeeAppendix 1 for explanationof site related to high salinity and the other to low pH. codes.ns, not surveyed. Groupings among the remaining wetlands appearedto be related to colour and nutrients. In a similar analysis of 23 shallow, permanent lakes on the south-westemcoast, Edward et a/. (1994)found Summer 1994 62NS Winter 1994 712 10 groupirrgs based on invertebrate communities Summer 1995 102 appeared to be related to saliniiy (although all Winter 1995 566 salinities were <3000 mg Li) and nutrients. In a Summer 1996 11 10 1 study of the macroinvertebrate communities of Extra specieswinter 1995 201 rivers across north-western Australia, based on Extra speciessummer 1996 0 familyJevel identifications, Kay et aI. (1999)found Total number of species 15 14 11 variables measuring geographic positiory salinity Aquaticfauna of the southemCamarvon Basin 233 and river discharge were more important than (Table 4). Lack of survey in Western Australia turbidity, alkalinity and nutrients. means that rnany widespread species have been In this survey, wetland groups based on recorded rarely. For example,for 50 years the only invertebratecommunities were best correlatedwith known Australian locality of the harpacticoid ratio of calcium,/alkalinity (not measuredin other Cletocamptusconfluens was Shark Bay (Lang 1948), studies), salinity, turbidity, colour, flow (which until it was found throughout Westem Australia in separatedrivers and springs from lentic wetlands), the 1990s (Halse ef aI., 1996; this study; Halse, geographic position, phosphorus and nitrogen. unpubl. data).However, some speciesbelonging to Storey ef aL (1993) suggestedthat environmental groups for which therehas been comparativelyhigh variables best related to wetland groupings vary collecting effort should probably be treated as according to the range of wetland types being endemic to the Basin until there is evidence to the studied, scale of the study and landscape setting. contrary. These include the calanoid copepod For example, environmental variablessuch as ratio Calamoecinhalsei (Bayly, 1998),several ostracods and of calcium/alkalinity will show far more variation the anostracanBranchinella sp. nov. (aIf.Iyrifera). if birridas, rivers and freshwater claypans are Gibson et al. (2000)found that, while most of the sampled than if only one wetland type is studied. wetland flora in the southem Carrrarvon Basin was In surveys of small areas,geographical coordinates typical of arid areas,significant numbers of south- are unlikely to be important. Similarly, turbidity western and tropical species occurred at the will probably be more variable in lentic waterbodies northem and southem limits, respectively,of their than rivers, and show greater variation in north- ranges. The same is true for aquatic invertebrates western Australia, where soils often contain a high and the southern Carnarvon Basin aDDears to proportion of clay or loam, than on the Swan represenla zonewhere Bassianand Torreiian biotic CoasialPlain, where they are sandy. elements meet (see Serventy and Whittell, 1967). bxter-couelation among environmental variables Examples of Torresian speciesthat have extended in this survey creates doubt about the vaudity of their range southwards are the copepod some correlationsbetween wetland groupings and Eudiaptomuslumholtzi (see Timms and Morton, environmental variables. The observed corelation 1988)and beetlesBerosus dallnsae and Hydroglyphus between longitude and wetland groupings may leal (see Watts, 1978, 1987). Examples of Bassian have been at least partially the result of strong speciesextending north are the copepod Calamoecia correlations with TDS, flow regime or distance from tasmanicasubattsnuata (see Maly et al., 1997) a\d the coast (which was not measured),rather than ostracodsMytilocypris mytiloides and Australocypris reflecting large-scalezoogeographic pattem. Strong insulais (seeDe Deckker,1978). Nevertheless, most inter-corelations between nutrients, tu$idity and aquatic invertebrates in the Basin either have colour may obscure the relative influence of these Eyrean affinities or occur throughout Australia. variableson comnunity composition, Groups such as anostracansand notostracansare typical of the former (Ceddes,1981; WilLiams, 1968); Biogeography many corixid hernipterars are examples of the latter Analysis of biogeographical patterns in the (Wroblewski, 1972;Knowles, 1.974). southem Carnarvon Basin data is made difficuli by Until now, only the chironomid Archaeochlus, the fact that distributions of most aquatic which occurs in small temporary streams on granite invertebratesare poorly known. About 50 species outcrops in south-western Australia and is also previously thought to be restricted to the eastem known from the Drakensberg Escarpment ald half of Australia were recorded in this survey and Namibia in southem Africa, was recognized as a speciesthat currentV appear endemic to the Basin Gondwanan relic in arid parts of Westem Australia 'fhe may be recorded elsewhere when surveys are (Edward, 1986; Cranston et al., 7987). conducted in other little studied regi.ons of occurrenceof aquaticinvertebrates, hitherto known Australia. Inability to identify some faunal groups only from South America, in the southern to species level further obscuresbiogeographical Carnarvon Basin raises the Dossibilitv that pattems, as well as preventing assessmentof the Gondwanan relics may o.c,.,t q..,ite commonly il conservation status of the unidentified animals. arid areas. It seems unlikely that the copepod Despite being a major component of the fauna of Robertsoniamourei, whic}l occurs in Lake Macleod, Australian wetlands (e.9.Geddes et a1.,1.981.;Davis could have been transported by migratory et aI., 7993),about 40% of ostracodscollected were shorebirds (Procteret aI., 1967)because there is no either undescribed or could not be confidently established flyway between South America and identified to species level. None of the Westem Austra-lia. The rotifer Hexsrthra brandor{fi ceratopogonid dipteran larvae could be identified seemseven more unLikelyto have been translocated to species. and almost certainly representsa relictual species. Most of the undescribed speciescollected in the Similarly, the undescribed Daphnia sp. nov. (aff. southem Carnarvon Basinare probably widespread barbata)artd. D. sp. nov. (aff. gibba) have strong S.A. Halse, R.J.Shiel, A.W. Storey,D.H.D. Edward, I. Lansbury, D.J. Cale, M.S. Hawey

African affinities and may be Gondwanan (C. wetland (Gibson et al., 2000). The most obvious Wilson, pers. comm.). Past studies of Gondwanarr implication for both invertebratesand plants is that relics in Westem Australia have focussed on the the number of speciesrecorded in the Basinwill wetter south-west (Edward, 1989; Cranston and grow as survey effort is increased. Edward, L992; Horwitz, 7997) and the signilicance Inlrequent occurence may be the result of aquatic of arid areas as habitat may have been invertebrates occurring in a narrow range of underestimated, particularly for species with a wetland types or conditions, being present for a resistant stage in their life history. short time in the flooding cycle, or having poor colonizing ability and locally restricted Wetland community pnt terns distributions. Maly et aI. (1997) argued that poor Waterbirds and aquatic invertebrates in south- dispersal,rather than narrow ecologicaltolerances, western Austra.lia appear to respond to the same shaped the distribution of most calanoid copepods environmental variables, such as salinity and in Australia. However, many of the species with a nutrient levels (seeDavis et. al., 1993;Halse et aI., single occurrence in the Basin were insects (see 1993; Storey et aI., 7993), yet wetlands supporting Appendix 4) with strong powers of flight. the largest numbers of waterbird species frequently Differences between communities of drying-phase differ from those with most inyertebrates. For (IW5 in winter and IS2 in summer) and other example, in a recent survey on the Swan Coastal claypans suggestedthat temporal successionmay Plain none of the five wetlands with hishest have contributed to sporadic collection of some invertebrate richness (Davis et. al., 1993)-were speciesbut it seemsunlikely to have been the factor among the 20 wetlands with highest waterbird underlying all rare occurrences.Gibson ef al. (2000) richnessor most breeding waterbfudspecies (Storey suggestedthe southem Camarvon Basin naturally et aI.,7993). contains many rare plant speciesand the same The same pattern applied in the southern seems true for aquatic invertebrate species, Carnarvon Basin. Although there was some although the reasonsfor rarity are not understood. superficial agreement between classifications of Rarity or infrequent occurrence of aquatic sitesbased on waterbirds and aquaticinvertebrates, invertebratesand other speciesposes problems for site-by-site comparisons suggested that the their conservatiory particularly when it is unclear processesunderlying fomation of waterbird and whether they are restricted to very few wetlands or invertebrate assemblages were different. occur very infrequently at many sites. Given the Information about community composition of high proportion of'rare'species with poorly waterbirds at a wetland could not be inferred understood and unpredictable distributions, it is reliably from data on invertebralesor aiceztersa (cl. udikely that all species can be conserved in a Figures 3 and 7). Plant communitiesprovided even regional nature reserve system. The realistic less information about the biota: the same plant objective for the reserve system should be community occurred at marine sites CB09 and protection of examples of all wetland types in the C877, brackish river site CB38a and freshwater southern Carnarvon Basin and their tvpical coastalpan CB36 (Figure 2 in Gibson et al., 2000), invertebratecomm u n ities. Other mechanisms, despite these sites supporting three waterbird and designed to ensure careful management of wetlands four invertebrate communities (Figures 3 and 7). on different land tenures, will be a necessary The Camarvon Basin results mirror those of yen adjunctto nafurereserves for conservationof some (1987),who found terrestrialbeetles, mammals and rarer, or infrequently occurring,species. vegetation each provided only limited information about the composition of the other two biotic elementsat sitesin Victoria. ACKNOWLEDGEMENTS Lack of concordancebetween classifications based Funding was provided by the Commonwealth on different taxonomic elements means that through ihe National ReservesSystem Co-operative inventory of each element is required to identify Program of the Australian Nature Conservation distribution pattems artd important habitats before Agency (now Environment Austraha), together shategiescan be prepared for the conservationand with funds provided by the Western Australian protection of that element. Wetlands wiih 1ow Department of Conservation and Land conservation value for plants may be irnportani for Management. We thank W.R. Kay for help with invertebrates or waterbfuds. stdtisticalanalyses, R. Schulz for chemicalanulyr"r, A. Clarke for sorting invertebrate samplesani fhe Rarespecies and theil conseraation following for assistancewiLh identificitions:I.A.E. A third of aquaticinvedebrate taxa were collected Bayly (somecalanoids), D. Belk (someanostracans), only once during this survey (Table 3). The R. Hamond (harpacticoids), S.A. Harrington pfrenomenon was even more pronounced in the (culicids),J.M. McRae (polychaeres),J.K. Moulton flora, with 55 % of species occurring at only one (simuliids), G.L. Pesce(some cyclopoids),S. Slack- Aquatic fauna of the southern Camarvon Basin 235

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Manuscript receixled29 lune 1998;accepted 4 AtLgtst 1.998.

THIS IS A BLANK PAGE Aouatic fauna of the southem Carnarvon Basin

Appendix 1 * Wetland sites sampled in the southem Carnarvon Basin, 1994-95. sampled August 1994,] October 1994,# July 7995 (winter), i March 1995(summer).

Site no Name Latitude Longitude Tenure TyP" cB04"+ Hardabut Pool 27'53'r1"5 114'34'04"E MtView Billabong cB05'+ Bullock Pool 27.49'04"3 114'46'31"E Riverside River pool CB06a.t Un-named swamp 27'31',26"5 115'04',20"8 Coolcalalaya Ephemeral swamp cB06b*i Un-named claypan n"31,'29"5 115'05'14'E Coolcalalaya Ephemeral pan cB09*i Un-named birrida 26"4513"5 tt3'42',U"8 Tamala Btrida CB09a* Un-named birrida 26'M'31"5 113.42'37"E Tamala Birrida cB15.t Un-named birrida 26'23'05"5 113'20'08"E Carramng Birrida cB16.r Un-named birrida 26't4',20"5 113'23',56"E Birrida cB20*t Hamelin Pool "25'40"5 114"11'33'E Hamelin Permanent swamp c825* Un-named clalpan 26"02',07"5 114"19'50"E Yaringa CB27a+ Nr Wardawarra Pool 26"04'49"5 115'27'rj"E Yalardy River pool cB27W+ Un-named swamp 26'1,5'22"5115"27'49"8 Talisker Ephemeral swamp CB27ct Un-named swamp 26'18'54"5 115'30'50"E Talisker Ephemeral swamp CB29a* Pan N of Big Lagoon 25'36'01"S 113'28'10'E Marine Park Birrida cB30#i Nr Namararra Well 25'39',30'S1t4"25'20"E Wooramel River pool cB34*r Mundilya Pool 25"39'54"S 114"50'54"E Meedo River pool CB35a* Un-named canegrasspan 25"40',52"S114.13',14"E'Wooramel Ephemeral canegrasspan c836* Ephemeral marsh 24'57',51"5 113'42'16"E Brickhouse Ephemeral coastalmarsh pan cB38*+ Chagra Well dalpan 25"11'47'S 11,4"57',02"8 jimba Jimba Ephemeral pan CB38a** Salt Gully 25"04'17"5 115'01'48'E Jimba Jimba River pool cB42*+ Winnemia Pool 25"00'32"S11,4"56'52"E River pool cM3* Un-named clalpan 25'04'77"5 115'03',30"E cB44. Rocky Pool 25"45'23"S 114'08'08"E Brickhouse River pool c849*+ Coollilee Pool 24'42'21"5 113'41'10"E Boolathana River pool cBs1#i Un-named canegrasspan 24'U'21"5 113"43'14"E Boolathana Ephemeral canegrasspan cB54*t Nr Cardabia Swamp 24'33'10'S 113'45'35"E Boolathana Canegrassswamp CBga# Cardabia Swamp 24"34'M"S 113"43'13E Boolathana Crabhole swamp c856.r rfr6rc !@]ya' 24"38'34',9 113'59'29"E Boolathana Ephememl pan C855a* Un-named pan 24"38',19"S113'59'35"E Boolathana c858b* Un-rramed dalpan 24'48'08"S 114'16'15"S Doorawarrah C858c* Un-named bluebush swamp 24"47'44"5 114010'05"E Doorawarrah Ephemeral bluebush swamp cB58d* Un-named canegrasspan 24"47'37',g 114'09'14"E Brickhouse Ephemeral canegrasspan CB58e* Un-named clalpan 24"47'35"5 114"09'14"E Brickhouse Ephemeral pan c 2.+ Mooka Ruin springs 24.53'25"5 rr4'57'29"E Mooka River pool CB62a*t Un-named creek 24.46'28"5 t14"56'17"E Mooka River pool c 2b+ Lake Julia 24'40'46"5 114"56'11"E Mooka Ephemeral swamp CB62c*t Un-named spring 24'46'.31"5 114'56'20"E Mooka Hillside seep cB67*+ Birdrong Spring 24't4',40"5 11,4'52'11"E Mardathuna Hillside seep cB67b'+ ScoopedHole 24"L8'07'S 114'50',28"E Mardathuna River pool c868* Un-named swamp 24"17'46"5 11,4"29'47"8 Hill Snnne Ephemeral swamp cB70b*i Bulgra Pool 24"25'47"5 11.4"32'54"8 Mardathuna River pool cB73*+ BoolanPool 24"28'38"5 113'40'36"E Boolathana River pool CB75a.* Cattle Camp Pan 24.28'25"5 11,4"13'27'E Cooralya Ephemeral pan cB75b*i Bluebush Bore Swamp 24'28'79"5 114"18'08"E Cooralya Bluebush swamp CB75c. Dwyers Pan 24.26'O0"5 rr4'27'18"E Mardathuna Ephemeral pan cB76*+ Lake McLeod 23'57'39"5 113"36'40"E Gnaraloo/Dampier Salt Birrida CB77+ Lake Mcl,eod 23"54',53"S113'39'23"E Gnaraloo/Dampier Salt Birrida cB78+ Lake Mcl,eod 23'52'19"5 113'42'43"E Gnaraloo,/Dampier Salt Marine pan cB79fr Lake Mcleod (Blue Holes) 23'47'16"5 r73'44'44"E Gnaraloo/Dampier salt Marine pan cB82*+ Minilya Pool 23'52'04"5 113'58'44"E Minilya River pool cB93*+ Cardilya Pool 25'37'34"5 115'31'28"E CareyDowns River pool CB93a.+Bidgelang Pool 2534'32"5 115'36'01"E Carey Downs River pool CB93c*+Callltharm Spring 25"52.59"5 115'30.14"E Carey Downs River pool cB93d*t Nwnery Pool 25'51'45"S 115'31',53"E Carey Downs River pool CB93FTBoothawalla Pool 25'47'37"5 775"17'26"E Carey Downs River pool CB93g. Meedo Pool 25M',29"5 11 Gilroyd River pool S.A. Halse, R.J.Shiel, A.W. Storey,D.H.D. Edward, I. Lansbury, D.J. Cale, M.S. Hawey

Appendix 2 Values of environmental variables collected in the southem Carnarvon Basin in winter and suruner. See text for descriDtionof variables. 3 5']P a 2 - .? ;*: ;Y;:!AEHa,'boq.: ^*:,! E".!€E=!odzr[Ii s F - J * € i' S i _{ 6 3 t E i; H J 0 J EE€€ E- ;.:o; _sf da CB04 sururer 7.7 85 63 310 300 0.507 0.891 1.833 0.68 0.08 0.005 9.6 n.886 It4.S68 z CB04 winter 8.2 98 14 0.24 3600 1.035 0.265 0.582 0.39 0.01 0.002 27.886 714.568 2 CB05 surntr€r 6.2 83 77 280 260 0.468 0.926 7.917 0.75 0.06 0.007 7t 27.818 :fr4.n5 2 CB05 winter 8 110 8 2 1100 1.097 0.286 0.675 0.41 0 '14 0.007 27.818 1I4.nS z CB06a winter 8.3 120 0.76 23 0.196 0.669 4.793 0.44 0.01 0.006 27.524 LIS.OZZ I CB06b summer 7.5 68 1200 62000 1100 0.060 0.377 0.872 Z6 27.525 115.087 1 CB06b winter 7.4 108 1100 17000 84 0.217 0.203 1..917 3.4 0.01 0.005 2Z52S LIS.O87 I CB09 summer 7.3 52 50 15 52000 13.740 0.228 0.415 0.53 0.04 0.024 6.2 26]5g t'I3.7Og 1 CB09 winter 7.8 106 3 0.91 180000 25.618 0.186 0.370 2.1 0.01 0.004 26.753 rr3.709 | -1 CB09a whter 8.6 126 23 0.85 85000 34.043 0.269 0.543 3.2 0.02 0.002 26.742 113.710 CB15 suruner 7.8 77 39 5.1 140000 18.994 0197 0.471 1.6 0.02 0.147 3 26.385 113.336 1 CB15 winter 8 85 3 0.45 57000 U.043 0.286 0.657 2.1 0.01 0.015 26.385 113.336 1 CB16 sluruner 7.7 77 200 20 270000 6.596 0.195 0.186 1.2 0.005 O 2.4 .239 179399 1 CB16 winter 8.1 119 5 0.5 55000 44.907 0.282 0.719 1 0.02 0.002 26.239 113.399 1) CB20 surnmer 7.6 40 36 3.6 6nO 2.537 0.258 0.513 0.24 0.005 0.004 9.5 .428 II4.t9g 2 CB20 winter 8.2 167 3 0.35 6000 2.585 0.264 0.568 0.35 0.01 0.001 26.428 114.793 2 CB25 winter 83 290000 0.159 26.035 114331 1, CB27a summer 8 77 56 250 310 0.351 1.266 3.082 0.87 0.09 0.014 6.9 26.08 IIS.4S3 2 CB27b summer 6-7 75 60 300 150 0.303 1..171 2.397 0.67 O.O7 0.015 1.9 26.256 1-15.4641. CB27b winter 7.38 92 34 440 45 0.127 0.323 0.799 1.7 0.71, O.'25 26.256 115_464 1 CB27c sumrrer 7.5 78 51 26 1700 4.154 O3U 1..046 4.2 O.O4 0.092 I.3 26.315 115.514 1 '25.600 -t CB29a winter 8.6 139 7 0.5 94000 38.521 0.267 0.530 2.2 O.O2 0.017 773.470 CB30 suruner 8.6 110 17 3.5 150 0.780 2.325 2.820 0.21 0.01 0.003 I .658 II4.422 4 CB30 winter 8.91 702 24 1.1 140 0.811 2.611 4.622 0.54 0.01 0.11 .658 114'422 2 CB34 surrttlrer 8.1 95 15 1.5 260 0.939 7.604 2.242 0.31 0.01 0.004 g 25.665 II4.U8 4 CB34 winter 9.4 137 7 0.38 230 0.656 0.452 1.551 0.69 0.01 0.014 25.665 1I4.UB 2 CB35a winter 7.a 58 55 37000 480 0.101 0.723 2.397 0.84 0.24 25.68I II4.22I I CB36 winter 8 88 400 9100 630 0.075 0.040 0.399 7.1 0.67 O.O2Z 24.964 113.705 7 CB38 surunei 7.5 81 120 6700 370 0.076 0.426 0.240 1.2 0.43 9.2 25.196 tI4.gSI 1 CB38 winter 8.2 105 400 10000 190 0.060 0.177 0.147 1.6 0.58 'l,4OO 25.196 114.951 1 CB38a summer 8.6 88 8 5.8 7.293 0.982 0923 0.41 0.01 O.OZ 4.3 25.071 115.030 2 CB38a winter 9.1 115 6 1.8 690 5.324 0.802 0.971, 0.28 0.01 0.001 25.071 115.030 2 CM2 suruner 8.1 96 12 5.1 660 1.153 0j30 7.289 0.17 0.005 0.003 7.8 2S.OO9114948 z CBAZ winter 8.6 74 5 0.35 1700 2.387 D.4U 0.767 0.45 0 0.002 E.N9 114.948 Z CB43 winter 8.5 95 180 36000 550 0.021 0.061 0.799 1.5 0.39 O .072 115.058 1 CB44 winter 8.6 130 7 0.27 360 0.622 0.804 2.895 0.58 0 0.001 25.756 114.136 2 cB49 suErner 6.9 87 11.0 1200 460 0.190 0.776 1.525 0.82 0.21 0.036 16 24.706 113.586 1 CB49 winter 8.2 98 88 4100 230 0.119 0.233 0.737 1.3 0.48 0.008 24.706 113.686 1 CB51 summer 6.5 91 110 Z4O0 770 0.086 0.426 0.599 0.62 0.35 2.4 24.739 113.221 1, CB51 winter 8.M 704 70 5300 410 0.055 0.263 0.533 1.5 0.33 0.028 24.739 I7372I I CB54 sururer 8.2 40 91 50000 930 0.076 0.079 0.409 5 0.96 2.8 24.553 1.1.3.260I cB54 winter 8-2 95 1400 12000 260 0.046 0.138 0.369 1.8 0.05 0.008 24.553 113.760 7 CB54a winter 6.59 53 35 410 110 0.254 2.228 14.380 0.81 0.03 0.01s 24.s79 11.3.7201 cB56 suruner 7.8 85 91 4500 550 0.300 2.672 5.592 7.7 o.r7 0.05 20 24.649 1r3.gg2 1, CB56 winter 8.5 727 260 820 56 0.380 0.436 2.996 0.43 0.06 O.O2 24.643 113.992 1 CB56a winter 7.4 100 12000 38000 500 0.299 1.338 8.783 3.9 0.84 0 24.639 113.993 7 CB58b winter 8 101 1900 43000 250 0.069 0.276 2.397 B.I O.S2 0 24.802 174.271 7 CB58c winter 8.1 95 48 380 68 0.372 2.869 9.587 O.n O.O3 O.OO3 24.796 714.168 7 CB58d winter 9.3 156 780 880 91 0.053 0.339 1.198 1.9 0.01 0.015 24.794 114.154 1 CB58e winter 8.2 104 8800 22000 680 0.056 0304 1,.79A 2.4 1.5 0 24.793 114.754 1 cB62 sumrer 8.3 150 120 17 11000 2.776 0.477 0.472 2.8 0.01 0.006 2s 24.89L 1149s8 4 CB62 winter 7.6 74 6 0.45 2000 1.150 0.573 0.422 0.51 0.01 0 24.891 114.958 4 CB62a summer 8.2 124 76 2.9 11000 0.733 0.396 0.257 o9g 0.01 0.007 64 24.774 r1,4.gg} 4 CB62a winter 8.8 158 23 0.34 10000 1.120 0.387 0.194 0.68 0 0.006 24.n4 ]14938 4 Aquatic fauna of the soulhern Carnarvon Basin 247

,! bD D -F-b. \- ^=Z'- itr=ut6oq qa;;:<:€E=;r olc-q orft-.\;ER\+ -oX .6.O.o:!Y .i''< ; 3 d E i 0 J \JFFtJ(A CB52b sumrner 7.6 70 490 3100 400 0.145 1.013 1.198 3.7 0.01 22 24.68 114.936 1 CB67a summer 6.4 10 13 21 1200 1.796 7.047 0.474 0.47 0.21 0.034 15 24.244 \74.870 -1.029 3 CB67a winter 6.9 68 8 0.21 7300 \.921 0.499 0.91 0 0.012 24.244 774.870 3 CB67b summer 8.6 138 24 0.7 7200 1.217 0.534 0.234 0.43 0.01 0.005 M 24.n2 114.841 4 CB67b winter 8.2 145 12 0.34 6500 1.473 0.607 0.288 0.64 0 0.016 24.n2 114.U1, 4 -1.332 CB58 winter 8.1 110 140 400 43 0.435 2.933 0.53 0.02 0.009 24.296 11,4.497 1 CB70b summer 9.4 120 80 250 300 1.217 0.894 7.667 2.2 0.06 0.14 24 24.428 114.548 1 CB70b winter 7.8 101 1100 1400 74 0.294 1.278 2.397 5.5 0.64 0.134 24.428 1\4.548 1 CB73 suruner 8.8 94 160 960 3000 0.238 0.-145 0.723 7 L.1. 0.042 1,5 24.477 713.677 1 CB73 winter 8.5 97 7m0 2300 240 0.077 0.507 1..370 4.5 0.56 0.113 24.477 113.6n 7 CB75a summer 7.5 72 810 30000 1200 0.069 0.857 0.799 6 5.1 20 24.474 114.224 1 CB75a winter 8.2 110 2300 2700 91 0.124 0.954 1.198 2.1 1.5 0.047 24.474 114.224 7 CB75b summer 8 1.21. 160 1500 730 0.254 4.686 9.587 1 0.15 0.076 39 24.472 114.302 7 CB75b winter 7.5 84 550 1400 76 0.375 1.886 3.595 1.5 0.06 0.012 24.472 114.302 7 CB75c wirrter 7.2 104 460 700 53 0.434 0.s51 0.599 1.5 0.07 0.011 24.433 774.455 1 CB76 sufiuner 7.9 r52 13 3.6 53000 9.465 0.223 0.382 0.55 0.01 0.005 20 23.961 113.611 1 CB76 winter 8.2 1.47 9 0.26 46000 9.032 0.219 0.404 0.51 0.01 0.002 23.961, 113.611 1 CB77 suruner 7.8 164 1.4 4.8 41000 7.811 0.242 0.386 0.24 0.005 0.01 3.9 23.915 113.656 1 CB77 winter 8.9 145 5 0.21 40000 8.318 0.237 0.402 0.31 0 0.02 23.915 173.656 1 CB78 winter 7.4 133 8 0.05 32460 7.402 0.268 0.608 0.85 0.005 23.872 113.712 1, CB79 sufuner 7.2 84 1.7 0.6 40000 5.479 0.242 0.416 0.28 0.005 0 3.2 23.788 113.746 7 CB79 winter 7.3 105 72 0.05 41220 7.088 0.267 0.37s 0.48 0.005 23.788 113.746 7 CB82 sulY[rer 6.9 74 21, 540 140 0.741, 5.356 2.846 0.41 0.06 0.017 6.3 23.868 113.979 2 CB82 winter 8.8 U 110 100 340 0.676 0.9n 2.538 0.56 0.02 0.026 23.868 1,1,3.9792 CB93 summer 8 109 28 0.7 80 0.455 4.712 7.190 0.41 0.01 0.009 6.9 25.626 11,5.524 2 CB93 winter 8.8 1.02 24 1,.7 180 0.692 5.28923.967 2.8 0.01 0.04 25.626 115.524 2 CB93a summer 8.4 108 23 1.4 120 0.609 2.395 6.591 0.44 0.01 0.00€, 1I 25.576 115.600 2 CB93a winter 7.7 106 13 0.28 64 0s37 1.2-10 4.793 0.55 0 0.002 25.576 775.600 2 CB93c surrrrIler 7.6 99 15 6.9 590 1.522 1.023 1.065 0.18 0.005 0.014 6.2 25.877 115.502 4 CB93c winter 8.4 108 7 0.25 1700 1.416 0.590 0.799 0.23 0 0.001 25.877 115502 4 CB93d summer 8.1 99 1,7 16 310 1.41,6 1.414 1.031 0.24 0.005 0.014 6.6 25. 3 11s.531 2 CB93d winter 8.7 1,42 7 0.25 580 1.332 1.103 1.174 0.48 0 0.001 25.863 -1.033 \75531 2 CB93f surrrmer 8.3 89 14 3.4 680 1.712 1.027 0.28 0.005 0.009 6 25.792 175.291 4 CB93f winter 8.4 108 17 0.25 420 0.832 1.-179 3.414 0.78 0.01 0.026 25.792 71,5.291 2 CB93g winter 9.5 161 10 0.31 480 0.949 1.022 1.580 0.72 0.07 0.03 .747 115.116 2

' decimal degrees b flow category 242 S.A.Hals€, R.J. Shiel, A.W. Storey, D.H.D. Edward, I. Lansbury'D.J. Cale, M.S. Hawey Appendix 3 waterbirdsrecorded at wetlandsin the southerncarnarvon Basin 199495. See Appendix 1 for wetland sitenames,

* - €€ + A - € o-'!€ * 3o = 6 q"s4 rr rq-- s S Speci€j SEEEBEssBEEBS tEsESEEEEBEESE BlackSwan ll 2l AustralianSh€lduck 4 4 | t7 AustralianWood Dock 4 | ZO PacificBlackDu€k1863 56 I I GreyTcal l7 l0 347 185ll 2 49 PiDl-earedDuck z 33 Haftlhead Udd.ntified duck Au$ralasiu Grebe I 3 6 3 Hoary-hcad€dGrcbe I 3 2 GreatCrested C,rcb€ Unidcntificdtrtb€ 8 Dad€r I Linle Pied Coso6t 2 I 2 Pied Cormorant Litrte Blekconnorant 2 l8 Unidentifi€dcorlnoranl

Whit€-facedH€ron z9 2l 1632 Linle Egrct I 3l Gr€atEgEt I I 2\ Unidentifr€degrct Suiat€dHemn Ndkc€n Night Hcroo

Royal Spoonbill Y€llow-billedSpoonbil Whir€-b.uicdS.a

Mdsh Sandpipcr CornmonGrc€nshank Terek Saldpip€r CommonSadpiper I Grcy-tarl€dTattld

Gr@t Knot Red-neckcdStinl 7 SharFtailcdSandpip€r CurtewSandpiper 15 Pien Oystercar.her Black-wingedStilt Bandci Slilt 808 I PacificGolden Plover Grey Ptov€r 3 13 I 4

Black-front d Dottcrel 7 1322 Red-kne€dDotterel BandedLapwing

Silver Gull Gull-bilhn Tem CaspianTem Unidentifi€dTem

No. oI birds t3 a7 4a 44 a3 023 2 235 4 023 0lZ85 0122 r 36 844 029 Z 415 0 No. of sD€cies 7lot26 2 0 4 217l0 5 0 4 0l 416 2130 9 214 0 Aquatic fauna of the southem Carnarvon Basin 243

r E^nAaEHff€ cceHfifi*fificfe Be EesnsdBEEEuD if $€cf r{E{ EB EBF,. B dEBE838Et 163 l2 853 I 903 26 t7 '13 t212 92 I 22 1605 63 228 lM4 42 2 2 3135 2 tl 3l 3 0 2 l5 20 6 t2 I2 3 lo 2 29 1 82zo 3 ll 416 39 444 2 257 3A &11 4 ZO7 153 22 22 702 3682 lll 445 lll 6 I 2t67 I 16 16 l0 11123 ll I l1223 5 t0 34 z z 0 I l6 I I 6 E I I I 6 I t2 20 163 6 6 l0 l0 5 5 40553 tu 2 2 I 5 4 24 24 I I 85 92 4 27 42 3 3 53 3E l5l UE 70 8084 9492 rs291 305 I I 3l 3l 8 150 r92 t0 l0

3 l4

17781 1778I 42243 88 IO 2t47 50 t0 l0 12214222094000,1 0 4 7ll0 l89o2t9 50 787 292q2 80 6 2 I 16 3 r 34504 3 212 2 2 | O t2 0 0 0 3 O 3 4 | 4 tS t2 3 28 25tO Z | | I2 | 244 S.A. Halse, RJ. Shiel, A.W. Storey,D.H.D. Edward, I. Lansbury, D.J. Cale, M.S- Harvey

Appendix 4 Aquaticinvertebrates collected durin8 winter 7994or 1995and summet1995 in the southernCamarvon Basin. See Aooendix1 for wetlandsite names.

PROTISTA LOBOSEA ARCELLINIDA Arcellidae Arcella sp. Centropyxidae Centropyris sp. Difflugiidae D iJJlu g i a c o r o na W allich Diflugia ^ff. Bramen Penard DifrIugia sp. A Lesquercusidae Netzelia sp. NEMATODA Nematodasp. ROTIFERA DIGONONTA Digononta sp. BDELLOIDEA Philodinidae Philodina sp. MONOGONONTA Conochilida€ Cono chi lus do ss uarius H'udson Cono c hi lus hip poc repis (Schr?l*) Co no c hi I us natans (Seligo) Conochilus sp. Trochosphaeridae Horcella brehmi Donner Filiniidae F ilinia aus traliens is Kosre .t F il inia longis eta (Ehrenberg) F ili nia longis e ta LiM. tica (Zacharias) Hexarthridae H exart hra b rando4f Koste H ercrt htu fennic a (Lev afiet) Hercrthra mira (Hudson) Testudinellidae Tes tudine I Lr patina (Hermann) Flosclariidae Iacinularia sp. Sinanthe rina p rcceru (Thorye) PLOIMIDA Brachionidae B rachionus anSularis Gosse B rachionus bidetttatus Anderson B rac hionu s caly c iJlorus P allas I B rac hionus dicho tomu s Sheph^rd B rac hionus dinidiatus (B ry ce) B rachionus falcatus Zachadias Bruchionus keikoa Kosle Bruchionus lJratus Shephai Brachio us nihoni Ahlstrom B rachio nus aff . nor ae.ealandiae (Morris) Brachionus ^ff. pinneendr.r Koste & Shiel B rachionus plicatilis (Mluller, Brcchionus quad dentalrr Hermann Aouatic fauna of the southem Camarvon Basin 245

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U(J ()UUOOOO Bruchiona[ quadridentalus rhenan s (Lauterbom) B rac hio nus rubens Ehrenberg Brachi onu s urceo lari s (M\eller\ Brachionus sp. K erate I Ia austrulis (Berzins, lll Keratella sp. no\. (aff. australis grp) Ke rate lla proc ufl a (Thorw) Kerut e IIa q uadrata (Mttller) K erute I Ia s hc ki (Benins) Ke rute lkt tropica (Apsteln\ Keratella sp. A P lationus patulus (Mnller) Platyias quadico m is Ehrcnberg Asplanchnidae A sp lanc hna bri I htw e lli (Gosse) Aq lanc hna siebo ldi (L.y dig) Asplanchna sp. nov. (afi sierrldr ) Asplanchna sp. B Notorunatidae Cep halo de I Ia gibba (Ehenberg) Eosphora aff. anthadis Hafiing & Mye$ Eo sphora naj as Ehrcnberg Notonamta aff. copeus (Ekenberg) Notomamta aff. tipus Ehrenberg Notommatidaesp. A Synchaetidae P olyart hra dolic hop te ra Idelson Proalidae PDal€r ri8/roidea (Skorikov) Trichocercidae Tric hocerc a pus i IIa lennings Trichoce rc a simi lis (Wierzejski) ll Trichocerca similis grandis (Hauer) Trichocerca sp- Dicranophoridae Dicranophnrus epichais Halling & Myers Lecanidae kcane Brcndis (M\nay) Iacane ho enanni (Etldenberg) kcane luna (Mueller) kcane bulla Gosse Lecanepapuana (Mvr^y) Iacane thalera (Harring & Myers) Iacane ungulatu (Gosse) Ircane sp. B I*cane sp. C Euchlanidae Eachlani s di latata Ehrenberg Euchluis sp. nov- Euchlanis sp. B Mytilinidae My tilina I entralis nlncrucant ha (Gosse) Trichotdidae M acroc hae tus col Linsi (Gosse) MOLLUSCA BIVALI'IA Biva.lviasp. VENEROIDA Laternulida€ LatzmuLa aff. anatina (Linnafl)s, Aquatic fauna of the southem Carnarvon Basin

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OOOUOO(JU GASTROPODA NEOTAENIOGLOSSA Truncattida€ Truncatelli aff. guerinii Vill^ & Villa Bithyniida€ Gabbiasp. A Gabbia sp.B Marginellidae Marginellidaesp. OPISTHOBRANCHIA Scaphandridae

Diaphanidae ?Diaphanidaesp. BASOMMATOPHORA Planorbidae '!GlJtophysa sp. Gyfttulus sp. Isidorclla newcombi (Aduns & Aneas) Isidorellasp.B ?Leichardtia sp. Lymnaeidre A ustropep le a les soni (Deshayes) ANNELIDA POLYCIIAETA PHYLLODOCIDA Nereidae Nereidaesp. Polynoidae Polynoidaesp. CAPITELLIDA Capitellidae Capitellidaesp. ORBINIIDA Orbiniidae Orbiniidaesp. OLIGOCHAETA TTIBIFICIDA Enchytraeida€ Enchytm€idaesp. Tubiffcidae Tubificidaesp. Naidida€ Allonais pectinata (Stepenson) Dercfurcata (Mnller)

Dero sp. rvdir sp.CB1 Nair sp. Pristinellajenkina. (Stephenson) Pi stina longiseta Ehrenberg Pistina sp. OPISTHOPORA Opisthoporasp. HIRIJDINAE Hirudinaesp. Hl ARTHROPODA ARACHNIDA HYDRACARINA Hydracarinasp. Hydrachnidae Hydrachnaapp roxim.Lta Hal\k Hydrachna sp. Aquatic fauna of the southem Camarvon Basin

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UQ(.)O(.)UOOOOUOU Eylaidae Ellais sp. Hydrodromidae HJdrodrona sp. Limn€siidae Limnesia aff. aurtralica L\ndblad Hygrobatidae Coau s tralio b areI Ion I ipalp is (Lundblad) Corticacarus sp. Unionicolida€ Recifella sp. Unionicola aff . Iundbladi K.O. thets Pionidae P iona austraLica K.O.'l iets P iona cumbe r lande nsi s (Rainbow) Aturidae Albid ^ff. rectifrons K.Yiets Arrenuridae Arrenurus balLadonie n sis Halik

Arrenurus sp.B CRUSTACEA ANOSTRACA Branchipodidae Paraftemia informis L)nder

Thannocephalidae B ranc hineUd rlf nis won gane ns i s Linder llt B ranc hine lla dent ic ulata Linder BranchinelLa sp. ^o\. (^ff. llrifera) B ranc hine I La oc cidentulis (Dakir) .l tl B runc hine I La v,e Uardi Mllner I B ranchi ne I Ia probis cida Henry Brachinella sp. A NOTOSTRACA Triopsidae T ri ops aus trali ensi s aus| ruliens i s Spencer& Hall CLADOCERA Sidiidae D iap h@tos otna aff . austrcl iensi s Korovchinsky D iap halnos ona excis um Sars

D iap hano soma ung uic ulat um Grmey Diaphanosoma sp. .t Ialonopsk alstrul^ SaIs Lat a nop si s b rc hmi Petkov skj SarsiLatonaaff. papuana (Daday) Chydoridae Alonine gen. nov. Alona cdnbouei Gueme & Richard Alona diaphana King Alona aff. inrcticulata Shenet al. Alona aff. rectangula SaIs Alo na re c tan gu h noweze o Landiae Sars Alona rectanguh pulchra Hellich Alona setuloides Smimov & Timms Alona sp. nov. (aff. A. diaphana Aquatic fauna of the southern Camarvon Basin

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-iqi-i7 hh\o@ooh oooooo(J Alora sp. nov. B Alona sp. no'r'. C Alona sp. nov. D Alona sp. eo'r'. E Alona/B iapertura sp. A Biapertura afrnis s.l. (l,eydig) Biapertura aIf.longinE d Smimov Biapeftura aff. nacrccopa (SNs) .l tl B iape rturo ri Sidicaudis Sn..ffio'r' ll Biapertwa afI. riSidicaadis Smimov Biapertura sp. nov. l Biaperturu sp. B Biapertura sp. C Ce Isinotum hJp$ilop hum Frcy tl Chydorus eurynotus Sars ll - Dunhevedia cru$a King Ep heme roporut barrois s i Richard Leydi gia ^ff . acanthoc ercoides (Fischer) Monospilus' diporus S'mfiov & Timms Monospilus' aft. dipontr Smimov & Timms Monospilus' elongatus Smimo\ & Timms PLanicirclut alticainatus Frey Rdt sp- nov. Macrothricidae M ac rohrit brer k eta Smtnov M ac ro thrix aff . hi rs uticomis Norman & Brady .l Macrothrix ^ff. spinosa King Macrothrix sp. nov. Macrothrix sp. A. I Neothit anlata Gtrmey Ne ot hr ix p aucis e t osa Smimo!r' Neolin, sp. nov. (afi ruperamlata)

Moinidae Moina ^ff. australiensis Sals lt Moina ^ff. flexuoso SaIs Moina micruru K\rn Moina monqolica Dad^y Moina aft. weistanni lshikawa Moina sp.D Bosminidae Bosmina meridionalis S arc Daphniidae Ce riodap hnia c omuta S^rs I Ceiodaphnia aff. darta Richard C eriodap hnia af f . laticaudata Mueller Daphnia angulata Hebef Daphnia carinata KinE Daphnia cephalata Kjng Daphnia prcjecta complex Hebert Daphnia sp. rov. (^ff. barbab) Daphnia sp. nov. (aff. Bibba \ Daphnia sp. Dap hniop s is p usilla Sewenty Scap ho kbe ris kinqi Sars Simocephalus aff . heilongjianqensis shi & shi Sinocephalas w tuluselisabet hac (Kins) Aquatic fauna of the southem Carnarvon Basin

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QUUU UO OUUUOQ Simocepholus ))ictoriensis Smimov & TiInms Simocephalus sp. Ilyocryptidse IlJocryptus sp. r,ov. .ll OSTRACODA Trachyleberidida€ ActinocJthereis scutigera (Brady) Cytherideida€ Cypridei s aust ralie ns k Harlmar|.]f. Paradoxostomatidae Paradoxostonu sp. Pectocytherida€ M ckenzieartia po j ac ksone ne sis (McKenzie) Darwindidae Daryinula sp. Candonidae P hly ctenop ho ra ^ff . zealandio (Brady) sensu Hartmam .t Limnocytheridae Linnocythere dorsicula De Deckler Limnocyt he rc po rp hy re tica De Deckker ,l Limnocythere ^ff. porphJretica De D€ckter Parclimnocythere sp. nov. 2?5 Paralimnocytheridgen. nov- Cypridopsidae CJpridop s is funeb ri s Blady I Safi c, p rido p sis ac ule ata lcosta) Ilyocyprididae I lyoq p ris australi€ ns is SNs l Iboqpris perigundi De Deckker Cypddidae Alboa worooa De Deckler AnpullacJpris sp. nov. 469 ?Ampullacypris sp. nov . 498 Austrclocypris insularis (Chapman, Bennelongia sp. nov. 414# B enne I o ngia aust ralis (Bftdy) I

Bennelongiabarangaroo De Deckker tt tlll .ll. 11 ,III Bennelongia sp. Candono cyp r i s nov aezelandiae (Baird) I Cypericercussalinus De Deckker Cypericercus sp. nov. 415 ll Cf peric erc us sp. nov. 422 Cyperircrcus ?sp.nov. 444 ,t,, Cyp rct ta b ay IJ i McKenzre I .llll I Cyprettu sp.488 I Cyp rinotus kimbe rley ens is McKenzie llt Cyp rinot us edw ardi McKenzie Diaclp ri s die tzi (Herbsr\ ll Hemicyprk megalops Sals sens\l McKenzie ?Heryetoclpris sp. H e t erocJ p ri s tatei (Brady) HeterccJpris afi vdlia De Deckker Heterocypris sp. nov. 489 tl

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OQ ('J() () Ilyodromus candonites De Deckker Ilyodromus dikrus D€ Deckker yodromus aff. ,iridulus (Bt^dy) Ilyodrom8 sp. nov.255 Myilnqpris nJtiloides (Brady, .l Mttilocypris sp. nov.426 Tngonocypris aff. globosa De Deckl(er Newnhamia fenestra King ReticJpris pinguis D€ Deckker Strandesia aff. phoenix De Deckker Zonocypris sp. nov.466 Zonocypretta kalimna De D€ckker CONCHOSTRACA Lync€idae LJnceus sp. A Limnadiidae Limnadopsis sp. Limnadia sp. A Cyzicida€ C)zicrr sp. A

Clziclr sp. B .l Clzicrr sp. C Cyzicus sp.D

C)zi.,rr sp. E I Clzicrs sp. F I .t C)zic,ir sp. H CJzicus sp.I Cyzicus sp. CALANOIDA Acartiidae Acatlia sp.351 Centropagidae

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Thermoctc lop s decip iens (Kiefer) HARPACTICOIDA Canthocamptidae Mesochra Jlaya lang Aquatic fauna of the southem Camarvon Basin

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->i-> 888:e (JQU(J() OOO UOOUO('J Cletocamptus c on|lue ns (Schmeil\ ll Clctocanptus die te rs i (Rlch^rd) .l Nannonesochra arupinerrir (Brian) Laophontidae H ete ro kup honte oculata (Gnrney\ Onychocamptus bengalerrtr (Sewell) .l .ll Quinq ue laop honte w ellsi (Hamond) Ameirida€ AnEira sp.308 {ff. Anvira sp. Nitocra aJfinis Gumey Diosaccidae Amonardia sp- I],ov- Amphiascoides subdebilis (willey\ Anp hiascops is cinctus (C\ans) B u lbamp hiasc us i mus (Bndy) Robe rtso nia mo ur ei (Nogrera) Rob erts onia p rop inq ua (Scott) Robertsonia sp. A Diossacidaesp. Tisbida€ Tisbella timsae Gvney Thalestfidae Eudactylopus sp. Louriniidae Lourinia armata (Clat)s) ISOPODA Sphaeromatidae Sphaeromatidaesp AMPHIPODA Paracalliopiidae Paracalliopiidaesp. A1 Paramelitrdae Paramelitidae sp- ,A4 Ceinidae Ceinidae sp. A2 .I C€inidae sp. A3 DECAPODA Decapodasp. M I INSECTA EPHEMEROPTERA Caenidae Tasnanocoenis sp. ARR J or M lltl Baetidae Cloeon sp. llI Baetidaegenus I WA sp. 2 ; ZGYOPTERA Coenagriidae Austroagrion coeruleum (T1llyard) Is c hnu ra auro ra (B r a!€r, I k chnu rc hete rc sticta (Burmeister) .l I I Xanthag ion ery t hrcne u rum Selys tl lttl .l Megapodagrionidse Argiolestes pusillus Tillyai I Lestidae Aust ro les te s annulos us (Selys) Aust ro les tes aridus (T illy ard) lt Austrolestes io (Selys) ANISOPIERA Gomphidae Austrogonphus go oni Walson Aeshnidae Aeschna brev isrl la Rarnbw H eniana.x papuens is (Burmeister) Aquatic fauna of the southem Camarvon Basin

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U(JUQ()U U Corduliidae Hemicor.lulia tau Selys Prccordu lia afrini s (Sely s) Libelhlidae Aust rothemis nigris cens (Martif,) Dip lacode s bipunc tata (Bft\tet\ Dip hcode s tuunatode s (Burmeister) ll O rthe I rum cale do nicum (BraDer) .t lt lll P antala flav esc ens (F ^bficins) ll Trapezos tigmata stenoloba Walson Libellulidae sp. HEMIPTERA Saldida€

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Lhnogonus sp. Veliidae Micr elia oceanica Dist^nt M icrov elia p eramoe na Hale Micror,elia sp. Mesoveliidae Mesovelio I ittigera Hor"',ald]. Mesovelia sp. Corixidae A I rap toco rixa e urynomz (Kirkaldy) A I rcp toco r Lra hi rt ifrons (Hale) A g rup to c o ri:@ pan ipunc lata (Hale\ Agraptocori\a sp. Micrcnecta annae s.1.(Kirkaldy) I .l M icronectu gracilis H^le I M ic ronecta robusta Hale ,l Micrcnecta ? halei Wroblewski

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Tabanidaesp. B Dolichopodidae Dolichopodidaesp. Empididae Empididae sp. Sciomyzidae Sciomyzidaesp- Ephydridae Ephydridxe sp. A I Ephydridaesp. B I Ephydridae.p.C Ephldridae sp. D Ephydridaesp. E Muscidae Muscidae sp. LEPIDOPIERA Pyralidae Pyralidaespp. I TRICHOPTERA Hydroptilidae A ff i ! op i La g l obo sa \ryells HelLlethira sp. Ecnomidae Ecnomus sp. Leptoceridae Notalinafulva Ktmmin, I O?c€rrssp. AV6 I Oecer,r sp. AV28 l llll Tnpl?rndc\ autlral^ Na\as I .t tll lll COLEOPTERA Coleopterasp- C l Haliplidae Haliplus fuscatus Clalk l Holtplus kstudo Clatk I Glrinidae Dneu rus a ust ral is lFahnciust Dyliscidae Alldd€rrrr rrrrliSdrir 'Clarl) | | ltllll

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