Records of the Western AlIstralian MlIsfllm Supplement No. 61: 1-12 (2000).

A biogeographic survey of the southern Carnarvon Basin,Western Australia: background and methods

Allan H. Burbidget, N.L. McKenzie1 and Mark S. Harvey2

1 Department of Conservation and Land Management, PO Box 51,Wanneroo, Western Australia 6065, Australia

2 Department of Terrestrial Invertebrates, Western Australian Museum,Francis Street, Perth, Western Australia 6000, Australia

Abstract - This paper describes the setting and aims of a multi-disciplinarystudy to sample the biodiversity of a 75000 km2 region of Western Australia(the southern Carnarvon Basin), and identify biophysical factors related tothe. observed patterns of plant and animal distributions. The study wasdesigned to provide a quantitative biological basis for conservation planningand monitoring.

INTRODUCTION

Decisions on the management of Australianrangelands (e.g. Harrington et al., 1984; StaffordSmith et al., 1997) have been constrained by the lackof data on patterns in biodiversity over most of theAustralian arid zone. Even the species inventoriesof most phylogenetic groups in our arid bio-regionsare fragmentary. While quantitative surveys ofrange condition have been carried out over largeareas of the pastoral zone in Western Australia inrecent years (e.g. Payne et al., 1987; Pringle, 1991),most have focussed on a sub-set of plants and allhave ignored indigenous animal taxa entirely,thereby providing only a biased first approximationto the patterns in biodiversity. The need to acquiresuch information for long term planning ofrangeland management has led a number ofauthors to recommend that the conservationresources of the Australian arid zone be identifiedand measures taken to ensure that these values areconserved (e.g. Foran et al., 1990; James et al., 1995;Landsberg et al., 1997).

James et al. (1995) identified four key researchthemes relevant to the Australian arid zone: "(1)identification of spatial and temporal patterns ofdistribution of native biota; (2) quantification of theimpacts of pastoralism on native biota; (3)identification of, and control of, potential non­native pest species; and (4) development ofmethods and technologies to allow regionalconservation planning". We contribute data underthe first (inventory) theme and analyse these data toprovide a basis for improved regional conservationplanning. The existing conservation reservenetwork is highly biased in many parts of theAustralian arid zone (Thackway and Cresswell,1995). In particular, the major conservation reserves

in the Carnarvon Basin are confined to its peripheryand do not include examples of some prominentland surfaces (Burbidge and McKenzie, 1995; seealso Figure 1). Although there was significantinformation available in the form of opportunisticdata and surveys of restricted areas or relativelynarrow taxonomic groups, there has been nosatisfactory basis for planning a representative, yetcost-effective reserve system. We therefore set outto design and conduct a survey that would samplethe main environments of the region, allow aninterpretation of factors determining the patterns ofplant and animal distributions in the study area,provide an explicit basis for designing aconservation reserve system according to CARcriteria (comprehensiveness, adequacy andrepresentativeness; see, for example, Woinarski andNorton, 1993), and to commence the design of sucha system.

Stafford Smith et al. (1997) have argued that whileecologists need to provide ecological understandingof arid lands, it is also important that they activelyseek the integration of this understanding in thewider policy debate. We attempt to provide a muchbetter understanding of the biogeography of one partof the Australian arid zone, and a beginning to theintegration of such knowledge into the wider debate.

THE STUDY AREA

The Carnarvon Basin extends from near CapeRange in the Exmouth area, south to the MurchisonRiver and inland to the Kennedy Range. It is one ofthe major sedimentary basins of Western Australiaand has a geological history spanning much of thelast 450 million years (Wyrwoll, Stoneman, Elliottand Sandercock, 2000).

2 A.H. Burbidge, N.L. McKenzie, M.S. Harvey

CALM Leasehold

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World Heritage AreaBoundary

Bemier IslandNllIure Reserve

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Figure 1 The extent of present day pastoral leases and major conservation reserves in the southern Carnarvon Basin.

Carnarvon Basin survey - background

IBRA RegionsThackway and Cresswell (1995) provide an

Interim Biogeographic Regionalisation of Australia(IBRA) in which the country is divided into 80regions based on various physical and biologicalattributes. Western Australia includes 26 of theseregions, most of which are restricted to the state.The Carnarvon Region, located on the mid-westcoast of Western Australia (Figure 2), extends fromnorth of Exmouth Gulf south to Shark Bay. To thesouth lies the Geraldton Sandplains Region. Theboundary between these two regions is also part ofthe boundary between the South-West Province andEremaean Province of Beard (1990), i.e. theboundary between the arid and southern mesicparts of Western Australia. The present study wascentred on this area.

Table 1 lists the variety of names for thebiogeographic districts referred to in this andfollowing papers. The Carnarvon Region(Thackway and Cresswell, 1995) has a similarnorthern and eastern boundary to the CarnarvonBasin (see Wyrwoll, Stoneman, Elliott andSandercock, 2000). However, the Carnarvon Regionextends south only to Shark Bay where it adjoinsthe Geraldton Sandplains Region [equivalent toBeard's (1980) Irwin Botanical District], whichincludes the southern edge of Shark Bay and thesouthern end of the Carnarvon Basin.

Carnarvon RegionThe Carnarvon Region occupies the northern and

central areas of the non-marine parts of theCarnarvon Basin. Because of its geographicalposition, the region is influenced by both the winterrainfall of the south-west and the summer rainfallof the north. It has an arid to semi-arid climate,with a mean annual rainfall as low as 200millimetres in places. Severe droughts are aprominent element of the climate (see Wyrwoll,Courtney and Sandercock, 2000).

The Region is dominated by extensive, low­gradient, alluvial plains traversed by the Minilya,Gascoyne and Wooramel Rivers (Figure 2; Wyrwoll,Stoneman, Elliott and Sandercock, 2000). Theserivers flow intermittently. Low, open woodlands ofAcacia species, such as Snakewood (A. xiphophylla)and Bowgada (A. linophylla), over shrubs such as

3

Poverty Bush (Eremophila), cassias (Senna) andsaltbush, occur on these plains (Beard 1990 andreferences therein). Shrubs and hummock grassesgrow on the low sand ridges that are scatteredacross the plains. In northern parts, the plains gradeinto red sand dune fields, supporting spinifex andmulga communities that are reminiscent ofAustralia's red centre. In the south, near theboundary with the Geraldton Sandplains Region,the plains support Eucalyptus-Callitris woodlands onyellow sand dunes. Intermittently flooded claypansare common and, close to the coast, birridas(evaporite pans) occur in interdunal depressions.Low-lying areas, such as the birridas, the fringes ofLake MacLeod and the coastal flats, support richsamphire communities. In the east of the study area,erosional uplands such as the Kennedy Rangeprovide a contrast to the surrounding plains.

Land-use in the area is predominantly for pastoralpurposes, with significant areas on the marginsbeing set aside for nature conservation.

Geraldton Sandplains RegionThe Geraldton Sandplains Region extends from

Shark Bay south to the vicinity of Jurien andBadgingarra at about latitude 30° (Beard, 1990). Theclimate is dry warm Mediterranean, with cool wetwinters and hot, dry summers. The region isdominated by heaths and scrub-heaths near thecoast. Extensive areas of coastal limestone, partiallymantled by pale yellow to grey sands, support lowheaths with emergent thickets of Banksia andmallees, such as Illyarrie (Eucalyptus erythrocorys). Aparticularly impressive feature of the area to thesouth of Shark Bay is the Zuytdorp cliffs, which aretopped with windblown, almost prostrateshrubland. Further inland, these are replaced bymallee, Banksia or Actinostrobus scrubs and heaths,with Acacia-Allocasuarina thickets still furtherinland. Small areas of Eucalyptus woodlands occur,mostly in the southern part of the region. The partof the region included in the present survey ismostly covered with sand, with some limestoneoutcropping very near the coast, and extends southto the Murchison River (Figure 2).

Much of the northern Geraldton SandplainsRegion is used for pastoral purposes, with theremainder mostly being in conservation reserves;

Table 1 Comparison of the names used by Beard (1980, 1990) and Thackway and Cresswell (1995) for certainbiogeographic entities referred to in the present study.

Beard,1980

Irwin Botanical DistrictCarnarvon Botanical DistrictAustin Botanical DistrictAshburton Botanical District

Beard,1990

Northern SandplainsCarnarvon RegionMurchison RegionGascoyne Region

Thackway and Cresswell, 1995

Geraldton Sandplains RegionCarnarvon RegionMurchisonGascoyne

4

there is some vacant Crown land north-east ofKalbarri National Park.

Study Area BoundariesThe study area for the current project extends

north to the Minilya River, inland to the easternboundary of the Carnarvon Basin and south toabout the Murchison River (Figure 2). It includesthe floristically and zoologically rich area aroundthe boundary between the Geraldton Sandplainsand Carnarvon Regions [the 'mulga-eucalypt line'of Serventy and Whittell (1967) and others], as wellas Acacia woodlands and shrublands characteristicof the southern two thirds of the Carnarvon Region.It therefore includes the whole of the Shark Bayarea but excludes most of the northern tropical endof the Carnarvon Region, which is dominated byhummock grassland.

Why Southern Camarvon Basin?We positioned our study area to straddle one of

Western Australia's most important biogeographicboundaries, in an area known to have greatfaunistic and floristic complexity. This, it washoped, would help provide a more meaningfulunderstanding of the factors that determine thepatterns of distribution of the plant and animalgroups sampled in our study. We therefore sampledthe woodlands, heaths and shrublands of thenorthern Geraldton Sandplains and southernCarnarvon Regions, as well as the arid zonecommunities of Acacia woodland and hummockgrassland that characterise the Carnarvon Region.The area of the southern Carnarvon Basin asdefined herein was about 7.5 x 106 hectares. We didnot have the resources to sample the northern endof the Carnarvon Region.

An additional reason to extend the study area tothe south was to include all the Shark Bay WorldHeritage Property (Anonymous, 1995).

HISTORY OF EXPLORAnON

Aboriginal OccupationEvidence of Aboriginal occupation at Shark Bay

goes back at least 18000 years (Bowdler, 1990a).Cape Range, north of our study area but still withinthe Camarvon Basin, was occupied by Aboriginalpeople over 30 000 years ago and the range has acontinuous history of occupation since that time(Morse, 1993). Presumably the remainder of theCarnarvon Basin has a similar history, at least alongthe river systems, but there is no archaeologicalevidence available to confirm this.

Seven tribes occurred in the southern CarnarvonBasin, occupying more or less non-overlappingparts of the region (Tindale, 1974). These peoplesubsisted off wild plants and animals, although

A.H. Burbidge, N.t. McKenzie, M.S. Harvey

actual use is mostly not documented; in fact, verylittle is known of the traditional life and customs ofthe pre-European Aboriginal people of Shark Bay(Bowdler, 1990b). However, deposits at Cape Range(Morse, 1993) indicate that coastal inhabitingpeoples gathered a range of food items from themarine environment, including crabs, gastropods,dugongs and turtles. On Peron Peninsula,archaeological evidence shows that macropods andEmu eggs have been used from Pleistocene times.Fish and marine invertebrates have also beenimportant food sources for much of this time, butevidence for the use of dugongs and turtles is onlyavailable from the Holocene (Bowdler 1990a, 1995).Aboriginal peoples in the area probably used fire asa tool of land management, but the impact of theiractivities on the flora and fauna of the CarnarvonBasin are unknown. Clark (1992) provides a seriesof interviews with older members of the Yammatjipeople, who inhabited the Gascoyne region, butthese are primarily a record of life on early pastoralproperties, and little seems to be recorded of theirtraditional way of life.

European Exploration and SettlementThe first recorded landing by a European in

Western Australia was at Cape Inscription, at thenorthern end of Dirk Hartog Island, where CaptainDirk Hartog landed in 1616 (Battye, 1915). He alsomapped Dorre Island and the entrances to SharkBay (named later by William Dampier). Hartog andseveral other early Dutch sailors (including HaevickClaeszoon van Hillegom, J. van Roosenbergh,Wollebrand Geleynzoon de Jongh and AntonieCaen) made only passing reference to any naturalhistory observations they made. Near BernierIsland, Caen did make what appears to be the firstrecorded sighting of Black Swans by a European,although Willem de Vlamingh is known as the onewho brought the bird to the attention of Europeans,following his observations on the Swan River(Serventy and Whittell, 1967).

On the other hand, William Dampier, whoexplored the Shark Bay islands in 1699 (Marchant,1988; George, 1999), compiled some of the earliestbotanical collections by Europeans in Australia(Finney, 1984). These are still preserved in theUniversity Herbarium, Oxford. His illustrations offour birds he saw in the Shark Bay area (Red­necked Avocet, Pied Oystercatcher, Bridled Ternand Common Noddy) in his book Voyage to NewHolland, are the earliest known illustrations ofAustralian birds (Whittell 1954). Dampier alsodescribed a macropod, presumably a Banded Hare­Wallaby (Lagostrophus fasciatus) (Troughton, 1973).This was not the first reference to a macropod, theDutch having made earlier references fromHoutman's Abrolhos and Rottnest Island, but itdoes appear to be the first from north-western

Carnarvon Basin survey - background 5

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Figure 2 Location of sampling sites, place names mentioned in the text and major biogeographic boundaries(Thackway and Cresswell, 1995) in the southern Carnarvon Basin. Terrestrial sampling sites are marked assolid circles and aquatic sites as solid triangles.

6

Australia and the Carnarvon Basin. Dampier alsoreferred to other animals at Shark Bay, includinglizards, "manatees" (Dugongs, Dugong dugon), fishand various marine invertebrates.

Important plant and animal collections (includingterrestrial and marine collections) were also madeby the French, during voyages in the early 1800s[by Peron and Leschenault de La Tour withBaudin's expedition in 1801 and Quoy and Gaimardwith Freycinet's expedition in 1818 (Marchant,1998)), and by several English and Germanexpeditions (sea and land) in the mid to late 1800s.The German expedition led by Michaelsen andHartrneyer in 1905 produced significant collectionsof terrestrial and marine invertebrates (Michaelsenand Hartmeyer, 1907-1930). Numerous studieshave been made since the mid-1900s, addingconsiderably to our knowledge. Nevertheless, thesehave all been opportunistic collections orobservations, or have been confined to a relativelynarrow range of animals or plants.

Exploration of the interior for agricultural anddevelopment reasons began in 1839, when CaptainGeorge Grey discovered the Gascoyne River andexplored the coast in the vicinity of the river and thepresent day Carnarvon, where he reported thepresence of fertile soils (Battye, 1915). FrancisThomas Gregory explored the length of the GascoyneRiver in 1858 and, on his return to Perth, reportedthat there were several tracts of well-watered landalong the river that were admirably suited forpastoral purposes. Pastoral settlement followed soonafter. C. von Bibra took up land near the Gascoyne in1863, quickly followed by others nearby and at DirkHartog Island and near Freycinet Harbour. Theseearly attempts apparently failed; the first successfulattempt in the area appears to have occurred whenAubrey Brown and John Monger took up land at themouth of the Gascoyne River, Brown havingoverlanded 4000 sheep from York. In 1876 Brownestablished the pastoral property that later becameknown as 'Brickhouse' (Burnside, 1979).'Boolathana', just north of the mouth of the GascoyneRiver, was established by Charles Brockrnan in 1877.'Lyons River', east of the Kennedy Range, was takenup before 1877 and 'Jimba Jimba', at the junction ofthe Gascoyne and Lyons Rivers, in 1878. Stationsaway from the coast or river frontage wereestablished later; for example, 'Mardathuna' wasestablished in 1902 (Battye, 1915; McDonald, 1991).An interesting account of station life in the 1880s isprovided by Carter (1987); a different perspective isprovided by Clark (1992). The extent of present daypastoral leases is shown in Figure 2. Sheep numbersin the Carnarvon Basin peaked at over 1 million inthe mid-1920s but dropped sharply to less than halfthis number by about 1940. In the following 40 yearsnumbers did not exceed about 730 000 animals(Payne et al., 1987).

A.H. Burbidge, N.L. McKenzie, M.S. Harvey

Other exploitative activities began in the 1840s(Battye, 1915; Cooper, 1997). Guano mining startedin Shark Bay in 1847 and a small pearling industrywas commenced in the 1860s. Sandalwood cuttingcommenced on Peron c. 1860 and was being loadedonto sailing ships at Gladstone from the early 1870s.American and French whalers probably operated inthe area from about 1803, and in the 1950s and1960s whaling stations were operating first fromPoint Cloates and later from Carnarvon. Fishing (fora variety of sea foods) has also been carried outover a long period (Cooper, 1997; Fry, 1988). Apartfrom fishing, none of these industries has persisted,but salt extraction now occurs at Useless Loop,Shark Bay.

EXISTING KNOWLEDGE

Existing knowledge of the physical and climaticenvironment and the forces and processes that haveshaped the landscapes of the Carnarvon Basin issummarised in Wyrwoll, Courtney and Sandercock(2000) and Wyrwoll, Stoneman, Elliott andSandercock (2000).

At the beginning of the survey, knowledge of thestudy area's plant and animal species was based onopportunistic collections made for variousmuseums and herbaria (see above), particularly forthe Western Australian Museum and WesternAustralian Herbarium. Some of these (e.g. Burbidgeand George, 1978) have provided a good overviewof particular areas, but there has been no previoussystematic survey of the entire area.

A number of vertebrate pests occur in the studyarea, with the main ones being rabbits, goats, foxesand cats; their occurrence in this area is summarisedby Payne et al. (1987). Amongst plant pests in theregion, Buffel Grass (Cenchrus ciliaris) is known tobe a serious environmental weed (Humphries et al.,1991; Griffin, 1993) although there seems to havebeen little done concerning documentation of itsdistribution or environmental effects in the studyarea or development of management options.

The Western Australian Conservation ThroughReserves Committee (CTRC) providedrecommendations on a reserve system for theCarnarvon Basin (and all other regions for theState) (e.g. CTRC, 1974). The recommendationswere based on existing knowledge and constituteda valuable advance in reserve system design forthe Carnarvon Basin. However, mostrecommendations concerned areas on theperiphery of the region, with a number of majorcommunities not included in the proposals.

The available data did not provide a basis fromwhich to appraise future trends in biodiversityunder the disturbance regimes (particularly grazingand introduced predators and other pests) that havebeen imposed following European settlement.

Camarvon Basin survey - background

National Reserves System Cooperative ProgramThe commitment of the Commonwealth

Government, in cooperation with the States andTerritories, to develop a comprehensive nationalsystem of parks and reserves, was outlined in thePrime Minister's December 1992 Statement on theEnvironment. Through this initiative the NationalReserves System Cooperative Program (NRSCP)was established and administered by the AustralianNature Conservation Authority. The Program laterbecame the National Reserve System Program ofthe Biodiversity Unit in Environment Australia. Theaim of the NRSCP was to facilitate progress towarda national comprehensive, adequate andrepresentative system of reserves to be establishedprogressively by the year 2000. The southernCarnarvon Basin survey was funded under theNRSCP as a cooperative project between theCommonwealth Government and the Governmentof the State of Western Australia as part of thiscommitment.

THE SURVEY

General aims of the survey have been outlinedabove. The underlying strategy used was a quadratbased survey in which data were collected asbiophysical attributes measured at the same placein the same time frame, to enable us to investigateassociations between species occurrence andenvironmental gradients (Austin et al., 1984;McKenzie et al., 1991; Margules and Austin, 1994;McKenzie, Halse and Gibson, 2000).

General arrangements for the survey are outlinedbelow. Specific details are provided in individualpapers.

The Field Sampling ProgramIt was agreed between the State and

Commonwealth Governments that the surveywould include sampling of vascular plants, aquaticinvertebrates, at least one major terrestrialinvertebrate group, reptiles, amphibians, birds andsmall mammals, together with an analysis of soilattributes or water chemistry at each sample site.The sampling program was divided into threemajor phases:

• selection of sampling sites,• first sampling: aquatic in winter 1994, terrestrial

in spring 1994,

• second sampling: aquatic in summer 1994/95,terrestrial in autumn 1995.

Some additional sampling, to extend thegeographic and temporal coverage, was doneoutside these periods (see below and relevantpapers following).

Specifically, we aimed to:• carry out the first systematic survey of vascular

7

plants, aquatic invertebrates, terrestrialinvertebrates, reptiles, amphibians, birds andsmall mammals in the Carnarvon Basin,

• collect voucher material for relevant museumsand herbaria (particularly the WesternAustralian Museum and Western AustralianHerbarium) for detailed taxonomic appraisal,

• investigate relationships between speciesoccurrences and attributes of the study area'sphysical environment, to identify predictors ofgradients in species composition across thestudy area,

• provide an overview of the status anddistribution of species (both native andnaturalised) and species assemblages,

• provide a quadrat-based benchmark, againstwhich future changes in the status of the biotacan be measured,

• provide a regional context for conserving thestudy area's biodiversity,

• provide recommendations for improving thedesign of the present conservation reservesystem and

• provide recommendations for the conservationof priority species and species assemblagesoutside the existing conservation reservesystem.

Terrestrial SitesPreliminary selection of the general areas in

which sampling sites would be established wasdone by N.L. McKenzie, A.H. Burbidge, G.].Keighery, K.P. Aplin and M.S. Harvey and, basedon an appraisal of existing knowledge, includingvegetation mapping (Beard, 1975, 1976) andgeological mapping (Denman and van de Graaff,1982; Denman et al., 1985; Butcher et al., 1984;Hocking et al., 1982, 1985, 1987; van de Graaff et al.,1982). The preliminary selection was then refinedfollowing liaison with CALM regional staff (G.Leaman and R. Shephard). Actual sites wereselected in the field by N.L. McKenzie, MS. Harvey,G.]. Keighery and A.H. Burbidge between 7 Apriland 15 April 1994.

Sample sites were positioned throughout thegeographical extent of the study area in a stratifiedrandom array, with quadrats in typical examples ofeach of the main surface stratigraphic units thatcharacterise the study area. From two to six siteswere clustered around each of 13 survey areas,herein referred to as 'campsites'. A total of 63terrestrial sites were sampled. Details are describedin the relevant chapters following, but in generalterms, for the purposes of vertebrate sampling, eachquadrat was 400 x 400 m, centred on the pit linesand a smaller (30 x 30 m) quadrat used for samplingvascular plants. If there was obvious internalheterogeneity at a site, a second plant quadrat was

8

established. Invertebrate pit lines were placed nearthe vertebrate lines.

Campsite codes (Figure 1 and other papers inthis volume) are as follows: BB - Bush Bay;BO - Boolathana; CV - Cuvier; EL - Edel Land;GJ - Gascoyne Junction; KE - Kennedy Range;MD - Meedo; MR - Mardathuna; NA - Nanga;NE - Nerren Nerren; PE - Peron Peninsula;WO - Woodleigh; ZV - Zuytdorp.

All relevant pastoralists (who previously hadbeen written to) were visited at the time of siteselection. Site establishment (digging pitfall traps)was conducted from 27 April to late May 1994.Some opportunistic data collection (e.g. occurrenceof birds at sites) was conducted during the siteselection and establishment periods. Initially, 61sites were selected, but two additional sites wereestablished on Edel Land (EL) when it becameapparent from preliminary analyses that this areawas under-sampled.

Invertebrate pits were installed in August 1994and left open until August 1995.

Spring zoological sampling was conducted from26 September to 19 October 1994. Personnelinvolved, and sites visited, are outlined in Table 2.Sites listed in Tables 2 and 3 were sampled for fivedays and five nights in each period. Sites at EDwere sampled for birds by P. Stone in August 1995.These sites and 21 others were sampled for ground­dwelling vertebrates by N.L. McKenzie, J.K. Rolfeand W. Muir in November 1995. Further samplingfor ground-dwelling vertebrates was carried out onall but the Edel Land (EL) quadrats in March 1996[see McKenzie, Rolfe, et al. (2000) for details].

A.H. Burbidge, N.L. McKenzie, M.S. Harvey

Autumn zoological sampling was undertakenfrom 8 May to 3 June 1995 (Table 3).

Physical attributes were sampled by K-H.Wyrwoll and T. Stoneman (Wyrwoll, Stoneman,Elliott and Sandercock, 2000).

Aquatic SitesPossible sampling sites were identified by Halse

et al. (2000) from existing biological and geomorphicknowledge and field inspection during areconnaissance survey early in 1994. Relevantpastoralists were also contacted at this time. Siteselection was finalised at the time of first (winter)sampling (August 1994). The selected set of sitesincludes representatives of all common wetlandtypes in the region. Where possible, sites were alsosampled in summer (March 1995), but some did notcontain water at this time. A total of 56 sites weresampled, representing 53 wetlands. Samplingfocussed on aquatic invertebrates, waterbirds andvascular flora.

Conditions at the Time of SamplingThe study area has a rainfall regime showing high

variability through time and high patchiness(Wyrwoll, Courtney and Sandercock, 2000). Duringthe years 1994 and 1995, when most of oursampling was done, Camarvon and Meedo hadbelow average rainfall in 1994 and above average in1995, Gascoyne Junction had above average rainfallboth years, Hamelin Pool had below averagerainfall both years and Gnaraloo (north of Quobba)had close to its average rainfall in both years (Table4). At the terrestrial sites, the differences between

Table 2 Personnel, attributes sampled and sites visited during the spring zoological sampling in the Camarvon-Irwinstudy area during 1994. M = mammals, B = birds, R = reptiles and amphibians, 1 = invertebrates.

Team Personnel

1. N. Hall and P. Boglio (M), AH. Burbidge (B), ].K. Rolfe (R),A Sampey (I), I. Cresswell (M,B,R,I)

2. N.L. McKenzie (M, R), R.E. ]ohnstone (B), W. Muir (R),P.].L. West (I), P. Stone (B)

3. AA Burbidge (M), P.]. Fuller (B), A Desmond (R), M.5. Harvey and].M. Waldock (I)

Sites visited

Ba, G], PE, ZU

CU, MR, MD, NA

BB, KE, wo, NE

Table 3 Personnel, attributes sampled and sites visited during the autumn zoological sampling in the Camarvon­Irwin study area during 1994. M = mammals, B = birds, R = reptiles and amphibians, 1= invertebrates.

Team Personnel Sites visited

1. N. Hall and P. Boglio (M), AH. Burbidge (B), ].K. Rolfe and M. Cowan (R), EL, NA, MD, MR, CUA Sampey (I)

2. N.L. McKenzie (M, R), R.E. ]ohnstone and P. Stone (B), W. Muir and NE, WO, BB, KEL. Smith (R), P.].L. West (I)

3. AA Burbidge/P. Boglio (M), P.]. Fuller (B), A Desmond and ZU, PE, Ba, G]B. Maryan (R), M.5. Harvey and ].M. Waldock (I)

Camarvon Basin survey - background

Table 4 Annual rainfall for 1994 and 1995, and long­term average annual rainfall for selectedrainfall stations in the study area (data fromAustralian Bureau of Meteorology).

Station Rainfall (mm)1994 1995 average

Carnarvon 135 269 228Hamelin Pool 134 109 207Gnaraloo 200 233 230Meedo 92 322 216Gascoyne Junction 259 273 206

the seasons were most marked at Meedo. At thissite, sampling was difficult in spring 1994 due tothe dry conditions following a period in which sixof the previous eight months exhibited belowaverage rainfall. In autumn 1995 animals werenoticeably more abundant, and herbaceous growthmuch more obvious, following much above averagerainfall in February, March and April.

As noted above, some wetlands contained waterin both winter and summer, but some did not.

Data storage and analysisData were entered into standard PC databases

from which presence-absence matrices for site byspecies were generated for analysis. We used thecomputer package PATN (Belbin, 1989, 1991), todetect patterns of species composition in the datamatrices. In most papers herein, the followingsequence was used. The association measure 'Two­step' (Belbin, 1980) was used to determine thequantitative relationship between each pair ofspecies, to allow classification of species in terms ofthe sites at which they occurred. The 'Two-step'measure is robust for use in situations, such as thepresent one, where sites vary greatly in their levelsof species richness (Austin and Belbin, 1982). TheCzekanowski measure (Czekanowski, 1932) wasused to compare the quadrats according to theirspecies composition. This procedure is appropriatefor ecological presence-absence data and is robustto variations in species abundance patterns andsampling efficiencies (Faith et al., 1987; Belbin,1991). For both measures of association, a modifiedversion of un weighted pair group arithmeticaveraging (UPGMA - Sneath and Sokal, 1973;Belbin, 1991) heirarchial clustering strategy wasused, with the clustering parameter (Beta) set to-0.1. Two-way tables were constructed from theclassifications of sites and species, in order tofacilitate interpretation of both classifications. Forparticular groups, this analysis path was extendedin various ways, as described in the relevant papers;extensions included the use of group statistics andgeneralised linear modelling to relate biologicalpatterns to attributes of the physical environmentson the sampling sites.

9

All sites were photographed, and scanned imagesare available on CD in selected libraries, includingthe libraries of the Western Australian Departmentof Conservation and Land Management, theWestern Australian Museum and EnvironmentAustralia (Canberra).

FundingCommonwealth funding for this project

($438366) was provided under the NationalReserves System Co-operative Program of theAustralian Nature Conservation Agency (later theNational Reserve System Program of theBiodiversity Unit in Environment Australia). StateGovernment funding for the survey included morethan $100000 provided through the WesternAustralian Museum and more than $300000provided through CALM. The State Governmentcontribution included existing infrastructure andpermanent staff.

ACKNOWLEDGEMENTS

First, we would like to thank all those landmanagers who gave us permission to work on thelands under their care: Lockie McTaggart(Bidgemia), Bob Bettini (Boolathana), Paul Burt(Brickhouse), Dudley Maslin (Mardathuna), JamieDempster (Meedo), J. F. Sears (Nanga), HaroldCrawford (Nerren Nerren), Mike Meecham(Quobba), Mick Clausen (Woodleigh), David Smith(Dampier Salt Ltd, Lake MacLeod Division), Bruceand Carolin May (Yaringa), T.L. Kempton(Yalardy), D.P. and RM. Steadman (Wooramel),B.R and L.M. Brown (Tarnala), Noce Pty Ltd(Talisker), Frank Jasper Pty Ltd (Mt View), RJ. andL.M. Lefroy (Mooka), J.M. and N. Thorn (Minilya),G.M.H. and V.F. Baston Oimba Jimba), J.RWS. andJ.A. Hopkinson (Hill Springs), Brian Wake(Hamelin), Hakko Sunbay Pty Ltd (Gnaraloo),Mungullah Community Aboriginal Corp. (Gilroyd),Doorawarrah Pastoral Company Pty Ltd(Doorawarrah), Maslen Holdings Pty Ltd(Cooralya), JS. and M.J. Stephens (Coolcalalaya),Clough Resources (Carrarang), RJ. and MS Rogers(Carey Downs), P.A.F. Kopke and E.M. Gooch(Carbla), Bob and Dawn Porter (Riverside).

We would also like to thank Ian Cresswell ofEnvironment Australia's Reserve System Unit whonot only assisted with the administration of theCommonwealth funding but also assisted in thefield sampling program.

A number of CALM and WAM staff membersassisted with various aspects of the survey. Inparticular, we thank Keiran McNamara for his rolein negotiating with the Commonwealth concerningfunding for this and related projects, and GregLeeman, Kelly Gillen and their staff in CALM'sMidwest Region for assistance with local advice. Or

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Ken Aplin (WAM) and Professor Sandra Bowdler(University of Western Australia) assisted with thelocation of background reference material.

Funding (see above) was provided by theCommonwealth through the National ReservesSystem Co-operative Program of the AustralianNature Conservation Agency (now EnvironmentAustralia), together with funds provided by theWestern Australian Department of Conservationand Land Management and the Western AustralianMuseum.

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Manuscript received 26 March 1999; accr;pted 15 September1999.