Western Australian Museum | Western Australian Museum - …museum.wa.gov.au/sites/default/files/10....

35
Patterns in the composition of ground-dwelling araneomorph spider communities in the Western Australian wheatbelt M. S. Harvei, J. M. Waldock', N. A. Guthrie 2 , B. J. Durranf and N. L. McKenzie 2 Departl1H'nt of Terrestrial Invertebrates, Western ;\ustralian Museum, Francis Street, Perth, Western i\ustralia 6000, Australia T)epartl11L'nt of Conservation and Land Man,lgement, 1'0 Box 51 Wanneroo, Western Australia 6065, Australia Abstract Ground-dwelling Maneomorph spiders were sampled at 304 l]uadrats chosen to represent the geographical extent and diversity of uncleMed terrestrial l'lwironments in a 205000 km' Mea known as the Western Australian wlH'atbelt. A total of 744 species comprising 39 families were recorded, of which the families Salticidae (121 species), Zodariidae (117), Theridiidae (RO), Lycosidae (61), Oonopidae (46) and Lamponidae (41) exhibited a marked species-level radiation. For analysis, families with a high proportion of arboreal species, and l]uadrats that were often flooded or overtlv affected by secondarv s,llinitv, were excluded. Thus, a total of 622 specie's from 240 l]uadrats were analysed, with an average of 21.9 (s.d. species pCI' l]uadrat. Most of the variation obserVl'd in the patterns of species composition could be explained in terms of summer temperature, precipitation seasonality, soil salinity and pH attributes. Assemblage species richness was constrained bv soil salinity, except for the Lycosidae which showed a positive relationship. INTRODUCTION South-western Australia represents a region that is internationally recognized for its exceptional diversity of terrestrial life, according it the status of one of the world's top-20 biod i versi ty hots pots (Myers et n/., 20(0). This rilnking was largely based upon the remarkable radiations of the vasculilr flora, but it has become increasingly obvious that many components of the terrestrial invertebrate fauna are very species-rich and that high proportions of the fauna are endemic to the region (Hopper et nl., 19(6). Some terrestrial and aquatic invertebrates ilre also postulated as possessing high numbers of short-range endemic taxa (Hilrvey, 20(2). We confined our study to araneomorph spiders which are easy to collect in pit-fall traps and for which we have sufficient expertise to undertake species-levl'l identifications. Although sporadic descriptions of Western /\ustralian spiders were milde bv 19 th century European taxonomists on specimens collected from King Ceorge Sound and the Swan River Colonv, the first comprehensive publication was Simon (I 19(9) who published on the collections of a C;ermiln F:xpedition led W. Michaclsen and Ila to south-western Australia in 1905. A great varietv of were named, but unfortunatl'ly few of the names are currently in use due to the poor descriptions and paucitv of illustriltions. Very few araneomorph spider families are currently well known in southern Western Australia. Taxonomic revisions have been published for only a few taxa, including: Ammoxenidae (Platnick, 2(02), Anapidae (Platnick and Forster, 1989), Filistatidae (Gray, 1994), Gallieniellidae (Platnick, 2(02), Hersiliidae (Baehr and Baehr, 1987, 1989, 1992, ]993, 1995, 1998), Lamponidae (Platnick, 20(0), Nicodamidae (Harvey, 1995), Orsolobidae (Forster and Platnick, 1985), some Salticidae (e.g. Zabka, 1990,199], 1992a, 1992b, 2000; Zabka, 2(01), some Sparassidae (Davies, 1994; Hirst, 1989, 199Ia,b, 1992), Trochanteriidae (Platnick, 20(2) and most Zodariidae (BachI' and Jocquc', 2000, 2001; Jocquc' and Baehr, 2001; Jocquc' and Baehr, 1 . BachI' and Churchill, BachI', in press) Over the past 15 yeilrs, several broad-scale biotic survevs have been conducll'd in vVestern Australia, and spiders were one of the grou ps tilrgcted for detailed anal vsis to discern how the spider faun,) WilS distributed across the landscilpe. The first \\'ilS iln examination of the hunil of the Kimberlev raintorests in which spiders Wert' eXilmined in detail bv Milin (1991). A rich ilssemblage of both cursoriill and web-building types was IT'\·ealed. Perhaps the most comprehensive studv was in the southern Cilrnarvon Basin where a pitfall trapping program revealed a totil1 fauna of ilt le,lSt SOU (l Jilrvev ell/I., 2000; Main cI,ll,

Transcript of Western Australian Museum | Western Australian Museum - …museum.wa.gov.au/sites/default/files/10....

  • Patterns in the composition of ground-dwelling araneomorph spidercommunities in the Western Australian wheatbelt

    M. S. Harvei, J. M. Waldock ', N. A. Guthrie2, B. J. Durranf and N. L. McKenzie2Departl1H'nt of Terrestrial Invertebrates, Western ;\ustralian Museum, Francis Street,

    Perth, Western i\ustralia 6000, Australia

    T)epartl11L'nt of Conservation and Land Man,lgement, 1'0 Box 51 Wanneroo,Western Australia 6065, Australia

    Abstract ~ Ground-dwelling Maneomorph spiders were sampled at 304l]uadrats chosen to represent the geographical extent and diversity ofuncleMed terrestrial l'lwironments in a 205000 km' Mea known as theWestern Australian wlH'atbelt. A total of 744 species comprising 39 familieswere recorded, of which the families Salticidae (121 species), Zodariidae (117),Theridiidae (RO), Lycosidae (61), Oonopidae (46) and Lamponidae (41)exhibited a marked species-level radiation. For analysis, families with a highproportion of arboreal species, and l]uadrats that were often flooded orovertlv affected by secondarv s,llinitv, were excluded. Thus, a total of 622specie's from 240 l]uadrats were analysed, with an average of 21.9 (s.d.species pCI' l]uadrat. Most of the variation obserVl'd in the patterns of speciescomposition could be explained in terms of summer temperature,precipitation seasonality, soil salinity and pH attributes. Assemblage speciesrichness was constrained bv soil salinity, except for the Lycosidae whichshowed a positive relationship.

    INTRODUCTIONSouth-western Australia represents a region that

    is internationally recognized for its exceptionaldiversity of terrestrial life, according it the status ofone of the world's top-20 biod iversi ty hotspots(Myers et n/., 20(0). This rilnking was largely basedupon the remarkable radiations of the vasculilrflora, but it has become increasingly obvious thatmany components of the terrestrial invertebratefauna are very species-rich and that highproportions of the fauna are endemic to the region(Hopper et nl., 19(6). Some terrestrial and aquaticinvertebrates ilre also postulated as possessing highnumbers of short-range endemic taxa (Hilrvey,20(2). We confined our study to araneomorphspiders which are easy to collect in pit-fall trapsand for which we have sufficient expertise toundertake species-levl'l identifications.

    Although sporadic descriptions of Western/\ustralian spiders were milde bv 19 th centuryEuropean taxonomists on specimens collected fromKing Ceorge Sound and the Swan River Colonv, thefirst comprehensive publication wasSimon (I 19(9) who published on thecollections of a C;ermiln F:xpedition led W.Michaclsen and I~. Ila to south-westernAustralia in 1905. A great varietv of werenamed, but unfortunatl'ly few of the names arecurrently in use due to the poor descriptions andpaucitv of illustriltions.

    Very few araneomorph spider families arecurrently well known in southern WesternAustralia. Taxonomic revisions have beenpublished for only a few taxa, including:Ammoxenidae (Platnick, 2(02), Anapidae (Platnickand Forster, 1989), Filistatidae (Gray, 1994),Gallieniellidae (Platnick, 2(02), Hersiliidae (Baehrand Baehr, 1987, 1989, 1992, ]993, 1995, 1998),Lamponidae (Platnick, 20(0), Nicodamidae(Harvey, 1995), Orsolobidae (Forster and Platnick,1985), some Salticidae (e.g. Zabka, 1990,199], 1992a,1992b, 2000; Zabka, 2(01), some Sparassidae(Davies, 1994; Hirst, 1989, 199Ia,b, 1992),Trochanteriidae (Platnick, 20(2) and mostZodariidae (BachI' and Jocquc', 2000, 2001; Jocquc'and Baehr, 2001; Jocquc' and Baehr, 1 . BachI' andChurchill, BachI', in press)

    Over the past 15 yeilrs, several broad-scale bioticsurvevs have been conducll'd in vVestern Australia,and spiders were one of the grou ps tilrgcted fordetailed anal vs is to discern how the spider faun,)WilS distributed across the landscilpe. The first \\'ilSiln examination of the hunil of the Kimberlevraintorests in which spiders Wert' eXilmined indetail bv Milin (1991). A rich ilssemblage of bothcursoriill and web-building types was IT'\·ealed.

    Perhaps the most comprehensive studv was in thesouthern Cilrnarvon Basin where a pitfall trappingprogram revealed a totil1 fauna of ilt le,lSt SOU

    (l Jilrvev ell/I., 2000; Main cI,ll,

  • 258 M. S. Harvey, ]. M. Waldock, N. A. Guthrie, B.]. Durrant, . L. McKenzie

    Unique patterns of endemism were found, andspatial patterns in the composition of the spiderfauna were strongly correlated with climatic andsoil attributes.

    The dataset we investigated in this paper wascollated as part of a biodiversity survey of theWestern Australian wheatbelt (Figure 1), an inlandarea of some 205000 km2 ranging from Geraldtonto Esperance, to discern patterns of endemism in ahighly fragmented and disturbed landscape. Weprovide the first account of the Western Australianwheatbelt's fauna of grolmd-dwelling araneomorphspiders, and explore patterns in communitycomposition in terms of measurable attributes of thearea's physical environment, including salinity.

    METHODS

    Study areaThe wheatbelt study area has an area of 205 000

    km 2 and comprises all or part of fivebiogeographical regions (Figure 1). The inlandboundary is a little to the west of the 300 mmisohyet. The western boundary is east of the 600mm isohyet. Along the southern coast, the southernboundary approximates the 550 mm isohyet butavoided near-coastal environments. Its semi-arid tosub-humid climate is influenced by temperateweather systems (mainly winter rainfall, see Bureauof Meteorology, 2001), and its surface hydrology ischaracterised by low gradients, high potential salt

    KALBARRI

    28

    o 100

    Kilometres

    11 18 18

    IBRA Regions

    D Geraldton Sandplains 2 Se 3D Alon \Mleatbelt 1 Se 2D Janah Forest 1 Se 2D Mill1ee 1 Se 2D Esperance Plains 1 Se 2

    118

    Figure 1 Wheatbelt study area, showing the 24 survey areas and the positions of the 12 to 13 individual quadrats ineach survey area. Precise quadrat co-ordinates are provided in Appendix 1. The coloured areas indicatebioregional boundaries (Environment Australia, 2000; modified from Beard, 1980). The fine lines internal tothe bioregions are sub-regional boundaries defined from IBRA version 5 (Environment Australia, 2000;Morgan, 2001).

  • Araneomorph spiders 259

    these valley floors. Although the north-western(Geraldton Sandplains) and south-eastern(Esperance Plains) parts of the study area are ondifferent geological basements (e.g. Morgan andPeers, 1973), their geomorphic units could beclassified into the same landform catena accordingto soil profile attributes.

    Field sampling strategySampling strategy is discussed in McKenzie et al.

    (2004). The study area was divided into 24 surveyareas of similar size so that sampling was evenlydispersed across the study area's areal extent(Figure 1). Ten to 11 quadrats each of one hectarewere positioned to sample the geomorphic profileof each of 24 survey areas (Figure 1). In each surveyarea, a quadrat was positioned in a typical example(in terms of vegetation and soil) of each of the maingeomorphological units (Baxter and Lipple, 1985;Chin, 1986; Morgan and Peers, 1973; Mulcahy andHingston, 1961). An additional one or two quadratswere positioned to sample examples of these unitsthat were affected by groundwater salinity. Thequadrats chosen were the least disturbed exampleswe could find. Even the sites selected to represent

    ,.-- £'_~I~_~l! ___,

    Spillway deposits

    Qe Quailing ErosionalK KauringQd Quailing DepositionalM MonkopenBg BelmungingB BalkulingMg MalebellingY YorkA AvonMo MortlockSf SaltflatSw Swamp (fresh)

    Figure 2 Wheatbelt land forms, modified from Mulcahy and Hingston (1961). The landscape's plateau profilecomprises the duricrusted Tertiary laterite plateau and its derived spillway sands, while the dissectionprofile comprises finer textured soils derived from bedrock and pallid-zone clays beneath the duricrust.

    loads and high variability in flows (Commander etaI., 2002; George and Coleman, 2002; Hatton andRuprecht, 2002; Schoknecht, 2002). Overall, 74% ofthe study area has been cleared for agriculture; itsremaining bushlands are isolated woodland andshrubland mosaics embedded in fields of wheat,lupin and canola. Extensive areas are now affectedby dryland salinity (George et al., 1995).

    In phytogeographical terms the study areacomprises the semi-arid part of the South-WesternBotanical Province (Beard, 1980). Detaileddescriptions of the vegetation in the study area areprovided by Beard (1980, 1981, 1990). The studyarea's substrates are detailed by Chin (1986),Mulcahy and Hingston (1961) and Myers andHocking (1998). Most of it is an undulating plateauof Tertiary duricrust overlying Archaean graniticand metamorphic rock strata of the Yilgarn Craton.The plateau is variously eroded into elluvial andcolluvial spillway sand deposits supporting avariety of shrub lands, and dissected to exposevalley-slope and -floor units characterised byduplex soils derived from the Archaean basementstrata and supporting woodlands. Except in thesouth-west, chains of salt-lakes also occur along

  • 260 M. S. Harvey, J. M. Waldock, N. A. Guthrie, B. J. Durrant, N. L. McKenzie

    secondarily salt-affected sites were positioned tominimise proximity to wheatfields and avoidevidence of other disturbances such as rubbishdisposal, gravel extraction and previous clearing.Locations of the 304 quad rats are provided inAppendix 1.

    A consistent land form model was used toposition the individual quadrats (Figure 2), so anequivalent cross-section of the landscape profile'smain components was sampled in each survey area.This nested, stratified sampling allowed ananalytical option in which each survey area wastreated as a single landscape-scale releve; bypooling the lists of species from a survey area's 12or 13 quadrats, we might suppress the influence oflocal substrate on the classification structure,thereby exposing broad-scale patterns for analysis.

    The spiders on each quadrat were sampled usingfive 2-litre, glycol pits left open for one calenderyear (1825 pit trap nights per quadrat). Pits were 25cm deep, 12 cm in diameter, contained 0.4 litres ofglycol-formalin (320 ml ethylene glycol, 64 ml tapwater, 16 ml formaldehyde) and were set flush withthe ground surface. Each pit was protected by asquare plate of wood (0.15 X 0.15 m) suspended 5cm above the pit's mouth. The five pits on eachquadrat were positioned in different microhabitats- bare ground, leaf litter patches under trees, besiderotting logs, under low shrubs etc. They werealways at least five metres apart.

    Quadrats in the YO, QU, KL, MN, NR, WK, KNand HY survey areas were sampled from October1997 to September 1998. NO, ML, MO, WU, JB, WHand BE were sampled from September 1998 toOctober 1999, and DA, DU, PI, LK, UN, ST, GP andES from October 1999 to October 2000. Every fourto six months, specimens were removed from thepits and fluids replenished. After washing, sortingand identification, the specimens were lodged inthe collection of the Western Australian Museum.

    Physical attributes at quadratsLandform units in survey areas on the Yilgarn

    Craton were numbered from 1 to 12 according totheir position in the landscape profile (see Figure 2,McKenzie et al. 2004). Units belonging to thedissection profile were numbered from 1 to 7, whilespillway sand and duricrust units associated withthe old plateau profile were numbered from 9 to 12.Unit Bg in which spillway sand mantles a claybelonging to the dissected valley profile wasassigned to number 8. Units at the bottom of thedissection profile included fresh water swamps(unit 1) and saline flats (unit 2). The highest unit inthe landscape was duricrust pavement of the oldTertiary plateau (unit 12). Quadrats in survey areason geological basements other than the YilgarnCraton (those in the Esperance Plains and GeraldtonSandplains bioregions) were positioned using the

    relevant 1:250 000 maps of surface lithology (e.g.Morgan and Peers, 1973), and arbitrarily assignedto our 12-class landform catena according to theirsoil profile (texture and horizon sequence), positionin the relevant landscape's profile and soil origin.Thus, a deep, low-level sandsheet wouid beassigned to the same number as the lowest of thespillway sand units on the Craton (Monkopen = 9,in Figure 2). This approximation was considered tobe acceptable because data taken from the quadratsthemselves were used in subsequent analyses.

    Eighteen climatic attributes were derived for eachquadrat using ANUCLlM (McMahon et aI., 1995).These comprised annual and seasonal average andrange values for temperature and precipitation(Appendix 2). Soil attributes were also recordedfrom each quadrat (Appendix 3), includingnitrogen, phosphorus, potassium, pH, electricalconductivity, organic carbon, clay-silt-sandpercentages and magnesium. Eight landform andfive vegetation attributes were also generated(Appendix 1), including latitude, longitude,elevation, landform unit, soil drainage category,slope, salinity risk, salinity class, litter/log cover,tree cover, shrub cover, herb cover and habitatcomplexity (modified from Newsome and Catling,1979). Latitude and longitude values weredetermined using a hand-held GPS accurate to ±30m. Explanation of the salinity attributes is providedin McKenzie et al. (2004) and in Appendix 1.Because land form had already been used toposition the quadrats in each survey area, it couldnot also be used as an attribute in the compositionalanalysis.

    Analytical strategyThe analytical approach taken in this paper was

    based on the assumption that spatial distributionreflects an underlying correlation withenvironmental factors (Austin, 1991; Clarke, 1993).It is an exploratory design, and interpretation isbased on deductive rather than inductive logic(Oksanen, 2001). No experimental design has beenimplemented to assess a null hypothesis (Austinand McKenzie, 1988), so alternative hypotheses arenot excluded. Inferential statistics are used to testpatterns observed in the empirical data.

    The relative responses of wheatbelt araneomorphspecies to our pitfall traps are unknown as are theyear-to-year dynamics of population numbers inthese communities. We also recognise otherlimitations in sampling techniques which wereaggravated by staff and time limitations. Becausethese issues precluded reliable estimates of speciesabundance, we chose to restrict our analyses to dataon the presence-absence of taxa at individualquadrats (Austin, 1984; McKenzie et al., 1991). Thus,the input data utilised for the compositionalanalyses were survey area-x-genus, survey area-x-

  • Araneomorph spiders

    species and quadrat-x-species matrices. Speciesrecorded at only a single quadrat (singletons) wereexcluded from compositional analysis because theyconvey little information on patterns.

    To assess whether records of a species were moregeographically localised than would be expected bychance, the average distance between the quadratsat which a species was recorded was calculated andcompared with 1000 trials using an equivalentnumber of randomly arrayed records (quadratintersections). The probability (P) of achieving theobserved average distance was then assessed fromthe distribution of the 1000 random Iv derivedvalues.

    '1'0 gain a clear separation between species of non-saline environments and those tolerant or adaptedto saline conditions, the 51 quadrats affected bysecondary salinity were excluded from the quadrat-x-species matrix (see McKenzie et aI, 2003). Thisseparation provided a basis for interpreting thespecies composition of assemblages on the salinequadrats in the absence of extrinsic data on thehabitat preferences of virtually any of thearaneomorphs in the samples. Thirteen additionalquadrats were eliminated from this analysis becausethey were flooded for long periods during thetrapping program (10 in ephemeral fresh waterswamps, as well as PI04, UN13 and WK02). Thisleft 240 quad rats in the quadrat-x-species matrix,but did not distort the stratification (see Table 1 inMcKenzie et ai, 2003).

    Cluster analysis (from PATN, Belbin, 1995) wasused to expose patterns of species composition inthe data matrices. The Czekanowski measure(Czekanowski, 1932) was used to compare thequadrats according to their species similarities,because it is known to provide a robust measure ofecological distance (Faith et aI, 1987). A modifiedversion of the unweighted pair group arithmeticaveraging (UPCMA - Belbin, 1995; Sneath andSoka!, 1973) hierarchal clustering strategy was used,with the clustering parameter (Beta) set to -0.1. Thepartition structure of the resulting dendrogram wasused as a summary of compositional patterns inaraneomorph genera/species across the study area.Environmental attributes that conformed to thepartition structures were assessed for statisticalsignificance using Kruskall-Wallis one way analysisof variance by ranks (in the computer packageSTAT'ISTICA, Statsoft, 2001). The negativeexponential smoothing function in STATISTICAwas used to interpolate climatic surfaces across thestudy area thereby illustrating geographic patternsin climatic attributes that were found to besignificant.

    RESULTS

    The infraorder Araneomorphae was represented

    261

    in the survey by 39 families, and a total of 24 640specimens were identified to species level. Thenumbers of specimens identified from each quadratwere not particularly high despite large samplingeffort (average 81.1, s.d. 49.6. range 2 - 316), andthere are substantial differences between quadratsthat may indicate differences in samplingeffectiveness and lead to bias in comparisons ofquadrat richness. Some of the poorest quadratswere those on salt-flats or in low-lying salt-affectedareas. An additional six months of sampling at asub-set of the salt-flat quadrats revealed noadditional araneomorph species (Durrant andCuthrie, 2004), so we proceeded with analyses ofrichness on the basis that our samples adequatelyreflect trends. 111is sampling issue is investigatedfurther under the heading Richness, below.

    The representatives of five families were notidentified to species-level due to time and resourceconstraints, as well as uncertainty regarding theunresolved taxonomic boundaries at the specieslevel. These five families (Ctenidae, Cnaphosidae,Miturgidae, Sparassidae and Zoridae) are not listedbelow and were excluded from the analyses.

    Observations on species geographical occurrencethat are included in the annotated list below arederived from Table 1.

    AgelenidaeTwo species assigned to the Agelenidae were

    recorded in the survey. The first, Cenus 1 sp. 1, wasfound at the Kt survey area, whilst the second,Cenus 1 sp. 2, was found at UN and WK.

    AmaurobiidaeThe family Amaurobiidae was represented in the

    survey by a diverse assemblage of 27 species. Fourspecies were found at more than 20 quadrats(Cenus 2 sp. 01, Cenus 3 sp. 01, Cenus 3 sp. 02 andCenus 3 sp. 16) and seven species were found atonly a single quadrat. The amaurobiid fauna ofWestern Australia is poorly known, and thesystematic position of many of the species hereplaced in this family may reside in other families.

    AmphinectidaeThe survey revealed 11 species attributed to the

    Amphinectidae, most of which were found in just afew quadrats. They were predominately recordedfrom southern study sites.

    Anapidae'rhe anapid fauna of southern Western Australia

    is represented by the sole genus Chasmocephalon, agenus that is also found in eastern Australia alongwith a variety of other genera (Platnick and Forster,1989). All anapids are tiny spiders less than 2-3 mmin length. Three species of Chasmocephalon were

  • Table 1 List of araneomorph taxa identified during the survey; the families Ctenidae, Gnaphosidae, Miturgidae, Sparassidae and Zoridae were omitted from the analysis dueNa-

    to time constraints. The study sites (see Figure 1) are arranged approximately from north-west to south-east and each cell records the number of quad rats from whichN

    each species was recorded.

    Family Genus, species Study sites (arranged approximately north-west to south-east) No. ofquadrats

    NO ML ON MO WU WH BE JB YO QU KL MN NR WK KN HY DA UN DU ST PI LK GP ES (n=304)

    Agelenidae Genus 1 sp. 01 1 1

    Agelenidae Genus 1 sp. 02 1 1 2

    Amaurobiidae Genus 1 sp. 01 1 1 2 4

    Amaurobiidae Genus 2 sp. 01 1 1 1 5 2 3 1 1 1 4 1 2 1 24

    Amaurobiidae Genus 2 sp. 02 1 1 1 3

    Amaurobiidae Genus 2 sp. 03 1 1 2

    Amaurobiidae Genus 3 sp. 01 2 3 1 3 2 3 1 4 6 1 1')'"7d

    Amaurobiidae Genus 3 sp. 02 1 3 5 2 1 4 5 2 1 2 I 27

    Amaurobiidae Genus 3 sp. 03 1 4 2 1 8

    Amaurobiidae Genus 3 sp. 04 4 1 1 1 7 ~Amaurobiidae Genus 3 sp. 05 1 1 1 3 CFJAmaurobiidae Genus 3 sp. 06 1 3 1 1 1 6 1 14 :::::Amaurobiidae Genus 3 sp. 07 1 1 2 \"01...Amaurobiidae Genus 3 sp. 08 1 1

  • Amphinectidae sp.ll 1 1Amphinectidae sp. 12 3 ] 4Anapidae Chasmocephalon sp. 01 1 1 5 3 10Anapidae Chasmocephalon sp. 02 2 1 3 6 12

    sp.03 1 2 1 4Araneidae sp.03 1 ]Araneidae sp.04 1 1 ] J 4Araneidae sp. 05 1 1Araneidae sp.06 1 IAraneidae sp. 07 J 1 2Araneidae sp.08 1 IAraneidae sp.1O 1 1 2Araneidae sp. 11 J 1 2Araneidae sp. 12 ] 1Araneidae sp.13 1 1Araneidae sp.14 1 1Araneidae sp. 16 I 1Araneidae Argiope protensa 1 3 1 1 1 1 1 1 1 11Araneidae Argiope trifasciatl1 1 1Araneidae A IIstracantha minax 1 1 1 1 1 2 1 1 9Araneidae Oolophones sp. 01 1 1 2Araneidae Dolophones sp. 02 1 2 1 4Araneidae Dolophones sp. 03 1 1 2Araneidae Eriophora IJiapicata 1 1 2Araneidae Eriophora transmarina 1 2 1 1 1 J 7Araneidae Eriophora sp. 01 1 1 2Araneidae Eriophora sp. 02 1 JAraneidae Genus X sp. 1 2 1 4Araneidae Genus Y sp. 1 2 3Araneidae Genus Z sp. 1 1 J 3

    l'v1atilda sp. 01 1 1 1 1 1 6 2 3 1 1 1 1 20Cyatholipidae Matilda sp. 02 1 1 1 1 1 2 1 1 9Cycloctenidae Genus 1 sp. 01 1 1Cycloctenidae Genus 1 sp. 02 1 1Deinopidae Deinopis sp. 1 1 1 1 1 1 1 1 2 10Desidae sp.Ol 1 1Desidae sp.02Desidae sp.03 1 JDesidae sp.04 1 J 1 3Desidae sp.05 1 1 2Desidae sp.06 1 1 2Desidae sp.07 1 1Desidae sp 08 1 1Desidae sp.1O ] IDesidae sp. 11 1 1 2 N

    '"v.>

  • Table 1 (cont.) NCl'Cl'

    Family Genus, species Study sites (arranged approximately north-west to south-east) No. ofquadrats

    NO ML DN MO WU WH BE JB YO QU KL MN NR WK KN HY DA UN DU ST PI LK GP ES (n=304)

    Lamponidae Lamponina sp. 02 1 1Lamponidae Lamponina sp. 03 1 1 2 1 1 6Lamponidae Lamponusa gleneagle 3 1 4Lamponidae Longepi woodman 1 1 7 1 4 1 5 1 1 1 3 26Lamponidae Notsodipus domain 1 1 2 1 5Lamponidae Notsodipus muckera 1 1 1 3 2 2 1 1 1 1 14Lamponidae Notsodipus quobba 2 2 2 6Lamponidae Notsodipus sp. 01 1 1Lamponidae Prionosternum porongorup 2 2 4Lamponidae Prionosternum scutatum 6 2 1 4 2 2 1 3 21Lamponidae Pseudolampona boree 1 1 2 1 1 1 1 1 1 1 2 2 2 17 ~Lamponidae Queenvic mackay 3 1 4 rJJLinyphiidae sp.Ol 2 2 4 1 6 2 4 3 5 2 1 2 4 2 1 3 2 1 2 49 :I:Linyphiidae sp.02 4 1 1 1 1 3 2 3 3 4 4 1 1 1 3 1 34 IIILinyphiidae sp.03 1 1 5 2 1 2 1 3 4 3 4 4 3 3 3 1 1 1 43 ::lnlLinyphiidae sp.04 4 1 1 3 8 7 6 2 2 1 1 1 3 4 6 2 1 2 3 2 1 61 ~..Linyphiidae sp.05 1 1 3 5

    ~Linyphiidae sp.06 1 3 2 1 1 1 1 1 1 1 13Linyphiidae sp.07 2 2 4 4 6 2 2 1 1 3 3 2 2 2 36 ~

    III

    Linyphiidae sp.08 2 2 2 1 1 1 1 1 1 1 13 c::Linyphiidae sp.09 4 2 1 1 1 3 4 2 3 2 23 0,.,Linyphiidae sp. 10 1 1 2 ~

    Linyphiidae sp.11 1 3 1 1 1 1 1 2 4 1 1 17 ?-Linyphiidae sp. 12 2 2 2 5 9 4 2 2 3 1 2 2 1 1 1 1 1 1 1 43 ~Linyphiidae sp.13 1 1 4 2 1 1 1 11 ClI:Linyphiidae sp.14 3 2 3 4 1 4 4 1 4 1 1 1 4 4 1 38 ::rLinyphiidae sp.15 1 1

    ....~.

    Linyphiidae sp. 16 2 1 1 1 1 2 8~

    Linyphiidae sp.17 5 6 5 4 4 1 1 1 27 ":""Linyphiidae sp.18 5 3 7 6 8 7 1 1 1 1 1 1 1 1 1 1 46 ClLinyphiidae sp.19 2 1 1 4 I:........Linyphiidae sp.20 2 3 1 1 3 1 11 III::lLinyphiidae sp.21 1 1

    ,....Linyphiidae sp.22 1 2 3 1 7 Z

    Linyphiidae sp.23 2 2 r'

    Linyphiidae sp.24 1 1 ~,.,Linyphiidae sp.25 1 1 ~nlLinyphiidae sp.26 2 1 3 ::lNLinyphiidae sp.27 1 1 ;;;.

  • sp.28 I 1sp. 29 ]sp. 30 1sp.31 Isp.32 1 Isp.33 Isp. 34 3 2 3 2 1sp.35sp.36sp.37sp. 39 2sp.40 2 2 I 1 2 3sp.41 1 1sp.42sp.43sp.44 j 1 2sp.45 ] 4sp.46sp.47 I 1sp.48 I I 2sp.49sp.50 I ]sp.51 1 I 2sp. 52 I Isp. 53sp.54 1 1sp.55 I I I 3sp. 56 I 1 I 2 2 2 9

    57D sp. Ol 2

    Oslearius 1 1 3 2 4 8 3 12 11 11 10Liocranidae sp. Ol 1Liocranidae sp. 02Liocranidae sp.04Liocranidae sp.05 1 1 I 2Liocranidae sp.06 I 1 1 1 4Liocranidae sp.07 I 1 1 2 1 2 8Liocranidae sp.09 1 1 1 3Liocranidae sp.12 2 3 3 2 2 I 1 3 1 3 2 1 3 2 2 2 I 34

    Arloria 1 1 3 2 2 9Arloria sp. 0] 4 2 1 4 1 2 4 10 8 1 1 1 5 44Arloria sp. 02 1 1 4 1 4 1 5 5 2 24Arloria sp. 03 2 1 7 2 1 6 3 2 1 6 6 7 2 5 6 7 1 4 3 6 5 2 5Arloria sp. 04 I 1 2 2 3 3 6 I 2 2 2 ,.)A rloria sp. 05 1 I I 2 1 t..l

    C1'.'1

  • , -----

    sp.21 3 3 ;.....sp.22 3 4 3 I 1 I 13 1:J::lsp.23 1 I 2 rt>0

    Lycosidae Lt/cosa sp. 24 I I 30

    sp. 25 1 I ...'1;lLycosidae Lycosa sp. 26 I I ::r

    CJ;

    sp. 27 1 2 3 '1;l0.:sp. 28 1 1 rt>...

    sp.29 1 2 2 I I 7 CJ;

    sp. 30 1 1Lycosidae Lycosa sp. 31 1 1Lycosidae Lycosa sp. 32 1 1 1 3Lycosidae Lycosa storri 1 2 1 1 1 I 1 8

    Venatrix arenaris 2 1 2 2 4 1 12Lycosidae Venatrix I'II11astra 7 1 3 4 2 1 4 22

    Jae Venonia sp. 01 1 1 2 4Micropholcommatidae Micr0l'holcomma? sp. 01 2 1 1 3 1 3 3 1 I 4 2 2 24Micropholcommatidae Micr0l'holcomma? sp. 02 I 1 1 4 3 3 1 2 4 2 2 1 25Micropholcommatidae Microl'holcomllla? sp. 03 2 I 2 1 1 ~IMicropholcommatidae Microl'holcomma? sp. 04 1 3 1 1 2 2 I 3 1 I 1 17Micropholcommatidae Microl'holcolllma? sp. 06 I 2 I I 5 2 1 13Micropholcommatidae Microl'holcomma? sp. 08 2 1 1 1 2 1 1 3 I 2 1 I 1 18Micropholcommatidae Micr0l'holcolllllla? sp. 10 1 1 1 1 4Micropholcommatidae Micropholcomllla? sp. 11 2 1 1 3 7Micropholcommaticlae Microl'holcolllma? sp. 12 2 1 3Micropholcommatidae Micropholcolllma? sp. 13 1 1 2Micropholcommatidae Textricella sp. 01 3 1 3 1 1 6 2 3 1 7 3 2 5 8 3 7 5 3 4 2 2 72Micropholcommatidae Textricella sp. 02 I IMysmenidae Genus 1 sp. 01 1 IMysmenidae Genus 2 sp. 01 1Nicodamidae AlIlbicodanllls marae 1 1 1 1 4Nicodamidae Nicodamlls mainae 2 6 1 2 4 1 1 1 I 2 2 3 26Orsolobidae sp.01 1Oecobiidae Oecobills naVllS 1 1 2Oonopidae Gamasomorpha sp. 01Oonopidae Gamasomorpha sp. 13Oonopidae Gamasomorpha sp. 02 1 1 2 1 SOonopidae Gamasomorpha sp. 03 1 2 1 1 5Oonopidae Gamasomorpha sp. 04 1

    Gamasomorpha sp. 05Oonopidae Gamasomorpha sp. 06 1 1 1 1 1 SOonopidae Gamasomorl'ha sp. 07 3 1 1 4 2 3 1 1 1 2 6 5 2 2 34Oonopidae Gamasomorl'ha sp. 08 2 2 1 1 2 2 6 2 1 1 2 7 2 3 2 5 3 8 2 1 2 1 58Oonopidae Gamasomorpha sp. 09 3 1 1 7 7 5 5 1 1 4 1 2 3 1 1 2 2 2 49

    sp. 10 1 1 NC!'\0

  • Philodromidae Tibcllus sp. 03 1 I ;;.....,Philodromidae Tibcllus sp. 04 1 I ::..;:lSalticidae Adoxotoma chillOPOSOIl I I 3 I 6 '"0Salticidae Adoxotoma Ilisroo/ivacca 1 I :30Salticidae Adoxotoma sp. 01 4 2 6

    ...,":l

    Salticidae Adoxotoma sp. 02 1 1 ::rCJ;Salticidae Biallor maculatlls 1 I 2 ":l

    Salticidae Biallor sp. Ol I I 2Cl:'"....Salticidae Brcda jovia/is 1 1 1 1 4 CJ;

    Szllticidae Clyllotis sp. Ol 1 2 1 3 2 3 I 2 1 1 2 3 22Salticidae Clyllotis sp. 02 2 1 1 1 1 1 2 2 1 2 14Salticidae Cll/lwtis sp. 03 1 I 1 3Salticidae ClYllotis sp. 04Salticidae ClYllolis sp. 05Salticidae Clyllotis sp. 06 1 1Salticidae Cllfllotis sp. 07 1 1 2 4Salticidae Clyllotis sp. 08 1 1 1 3Salticidae Clyllotis sp. 09 1 2 2 1 6Salticidae Clyrwtis sp. 10 1 1 2Salticidae Clyllotis viduus 1 1 1 I 2 6Salticidae Cytaea sp. 01 1 1 2Salticidae Damoctas sp. 01 1 1 2Salticidae Genus 1 sp. (n 2 1 4 4 6 4 4 1 9 10 7 I 6 5 1 1 3 6 1 2 3 3 84Salticidae Genus 1 sp. 02 1 1 2 1 3 6 2 2 I 2 1 1 2 I 26Salticidae Genus 1 sp. 03 1 1 1 1 3 1 1 1 1 1 1 2 2 17Salticidae Genus 1 sp. 04 1 1 1 3SaIticidae Genus 1 sp. 05 1 1 1 3Salticidae Genus 1 sp. 06 1 1Salticidae Genus 1 sp. 07 1 1 1 3Salticidae Genus 10 sp. 01 1 1 2Salticidae Genus 12 sp. 01 1 1 2Salticidae Genus 12 sp. 02 1 3 2 1 7Salticidae Genus 12 sp. 03Salticidae Genus 12 sp. 04 1 1 1 1 1 1 6Salticidae Genus 13 sp. Ol 1 2 2 1 2 1 1 1 11Salticidae Genus 14 sp. Ol 1 1 1 1 1 1 6Salticidae Genus 15 sp. 01 1 1Salticidae Genus 16 sp. Ol 1 1 2Salticidae Genus 17 sp. 01 1 1 4 3 1 10Salticidae Genus 18 sp. Ol 2 1 1 2 2 8Salticidae Genus 19 sp. 01 1 1Salticidae Genus 2 sp. 01 1 1 3 1 2 1 1 8 3 3 7 8 5 6 3 1 1 3 1 3 2 3 2 69Salticidae Genus 20 sp. Ol 1 1 2Salticidae Genus 3 sp. 01 2 1 1 1 2 2 9Salticidae Genus 3 sp. 02 2 2 1 1 1 2 9 N

    '-l....

  • Table 1 (cont.) N'IN

    Family Genus, species Study sites (arranged approximately north-west to south-east) No. ofquadrats

    NO ML DN MO WU WH BE JB YO QU KL MN NR WK KN HY DA UN DU ST PI LK GP ES (n=304)

    Salticidae Genus 3 sp. 03 1 1 1 3Salticidae Genus 3 sp. 04 1 1 2Salticidae Genus 3 sp. 05 1 ISalticidae Genus 3 sp. 06 1 1Salticidae Genus 5 sp. 01 1 2 2 4 1 1 2 1 1 1 16Salticidae Genus 6 sp. 01 1 1 1 3Salticidae Genus 7 sp. 01 2 2Salticidae Genus 9 sp. 01 1 1 2 1 1 1 1 2 1 1 12Salticidae Grayenulla allstralensis 2 6 2 4 2 3 9 2 1 1 5 1 38Salticidae Grayenulla nova 5 1 6Salticidae Holoplatys chlldalllpensis 1 1 1 3 ::::Salticidae Holoplatys dejongi 1 1

    'JlSalticidae Holoplatys sp. 01 1 1 ::Salticidae Holoplatys sp. 02 1 1 2 tJSalticidae Holoplatys sp. 03 1 1 ~reSalticidae Hypoblemllm sp. 01 1 1 2 2 1 7 :

  • Sillticidae Lycidas sp. 21 1 1 >-""SZllticidae Lycidas sp. 22 2 1 2 1 6

    OJ::l

    Sillticidae Lycidas sp. 23 1 9 7 1 18ro0

    Salticidae Lycidas sp. 24 1 1 30Sillticidae Lycidas sp. 25 1 1 ""'"ClSZllticidae Lycidas sp. 26 1 2 2 5

    ::r\J)

    Salticidae Llfcidas sp. 27 1 I '"Cl0.:SZllticidae Nlaratlls mllllgaich I 1 2 ro

    ""Sillticidile A1aratlls 4 I 1 1 1 1 I I 1 I 1 14 U1Sillticidae Maratlls sp. 01Salticidae Maratlls sp. 02Sillticidae Maratlls sp. 03 ISZllticidae Maratlls vespcrtilio 1 I I 2 3 3 4 2 I I 6 3 I 2 2 33

    Salticidae Margaromma sp. 01 2 1 4 4 1 3 3 5 4 1 1 4 2 3 I 1 40

    Salticicbe Margaromma sp. 02 2 2 2 1 1 3 1 1 3 2 18

    Salticidae MI/rmarachlle sp. 01 1 1 I 3

    Salticidae ivfyrmarachlle sp. 02SZllticidae Ocrisiolla /ellcocomis 1 1 2

    SZllticicbe Ocrisiolla sp. 01 1 1 1 3

    Sillticidae OpisthollCIIS sp. 01 1 2 1 1 I 1 2 1 0

    Sillticidile OpisthollCliS sp. 02 1 1 1 1 4

    Sillticidile OpisthollCliS sp. 03 1 1 1 1 4

    Sillticidae Paraplatoides sp. 01 1 1 1 1 1 1 1 1 2 1 1 1 13

    Sillticidae Prostheclilla sp. 01SillticidZle Scrvaea sp. 01Salticidae Simaetha sp. 01 2 1

    ~

    .)

    Salticidae Simaetha sp. 02Salticidae Simaetha sp. 03Sillticidae Simaethllla sp. 01Salticidae Simaethll/a sp. 02Salticidae Simaethllla sp. 03Salticidae SOlldra sp. 01 2 1 4 1 10 2 3 8 1 1 33Sillticidae Zebraplatys fractivi ttata 1 1 1 1 1 1 6

    SalticidZle Zebraplatys sp. 01 1 2 1 1 1 1 2 3 4 1 17

    Sillticidae Zebraplatys sp. 02 1 1

    Salticidae Zcbraplatys sp. 03 1 1

    Salticidae Zebraplatys sp. 04 2 2

    SZllticidae ZCllodoms sp. 01 1 1 2Genus 1 sp. 01 1 1 1 1 1 5

    Segestriidae Genus 1 sp. 02 1 1 2 2 3 1 2 1 1 14Genus 2 sp. 01 1 1 2 1 4 2 3 6 6 1 5 1 1 4 1 39DclClla sp. 01 1 1Genus 1 sp. (J1 1 1Baiami volllcril'cs 3 1 3 2 9 4 2 1 25Comsoides sp. (J1 3 2 1 3 3 6 1 3 22 N'-l

    W

  • Table 1 (cont.) N~...

    Family Genus, species Study sites (arranged approximately north-west to south-east) No, ofquadrats

    NO ML DN MO WU WH BE JB YO QU KL MN NR WK KN HY DA UN DU ST PI LK GP ES (n=304)

    Stiphidiidae Corasoides sp. 02 1 1 1 1 1 1 1 1 1 1 10Stiphidiidae Corasoides sp. 03 2 1 3Stiphidiidae Corasoides? sp. 04 1 3 1 1 1 1 1 1 1 11Stiphidiidae Corasoides? sp. 05 1 1 1 1 4Stiphidiidae Forsterina sp. 01 1 1 1 3Tetrablemmidae Genus? sp? 1 1Tetragnathidae "metine" sp. 01 1 1 1 3 1 7Tetragnathidae "metine" sp. 02 1 1 2Theridiidae sp.01 1 1Theridiidae sp.02 1 1Theridiidae sp.03 1 I

    ~Theridiidae sp.04 1 1

    fJlTheridiidae sp.05 1 1 :z::Theridiidae sp.06 1 1 ..Theridiidae sp.07 1 I

  • Theridiidae Genus 1 sp. 03 J 1 2 ;J;>...Theridiidae Genus 1 sp. 04 1 1 1 2 2 4 1 1 1 1 2 1 18 '"::lTheridiidae Genus 2 sp. 01 1 1 1 1 1 5

    r'I>0

    Theridiidae Genus 2 sp. 02 1 1 1 3 30Theridiidae Genus 3 sp. 01 1 2 2 2 1 1 1 I 1 2 14

    ...'\:l

    Theridiidae Genus 3 sp. 02 1 1 2::r'

    '"Theridiidae Genus 4 sp. 01 2 2 '\:l0.:Theridiidae Genus 4 sp. 02 1 1 1 1 4 r'I>...Theridiidae Genus 4 sp. 03 1 1 1 3 '"Theridiidae Genus 4 sp. 04Theridiidae Genus 4 sp. 05 1 I 2Theridiidae Genus 4 sp. 06 1 I 2Theridiidae Genus 5 sp. 01 1 1 1 3Theridiidae Genus 5 sp. 02 2 1 3Theridiidae Genus 5 sp. 03 2 J I 2 I I 8Theridiidae Genus 6 sp. 01 1 1 I 3 I 1 I I 2 I 5 I I 20Theridiidae Genus 6 sp. 02 J I 2Theridiidae Genus 7 sp. 01 I I 2Theridiidae Genus 7 sp. 02 2 2Theridiidae Genus? sp. J 5 1 1 1 8 2 2 4 3 2 2 1 33Theridiidae Gmogala sp. 01 1 1 2Theridiidae Gmogala sp. 03 1 1 3 2 1 6 2 2 3 1 3 1 2 5 1 2 2 3 1 1 43Theridiidae Gmogala sp. B 2 3 1 4 5 4 1 3 1 2 4 2 6 7 4 1 1 51Theridiidae Gmogala sp. U 1 3 2 1 2 1 9 2 2 1 1 25Theridiidae Hadrotarsinae Genus 1 sp. 01 1 2 2 1 1 1 8Theridiidae Hadrotarsinae Genus 2 sp. 01 1 1 1 3Theridiidae Hadrotarsinae Genus 3 sp. 01 1 1 2Theridiidae Hadrotarsinae Genus 0 sp. 01 1 1 1 2 1 6Theridiidae Hadrotarsinae Genus 0 sp. 02 1 ITheridiidae HadrotarslJs sp. 02 1 1 1 1 1 2 1 3 2 2 3 1 2 21Theridiidae HadrotarslJs sp. 03 1 2 1 2 1 4 2 1 2 1 2 19Theridiidae HadrotarslJs sp. C 1 1 1 1 1 1 1 1 2 5 1 16Theridiidae Latrodectus hasseltii 11 13 5 12 13 12 13 5 5 7 4 6 2 4 5 3 5 2 9 8 5 6 155Theridiidae Phorollcidia sp. 01 1Theridiidae Phorollcidia sp. 02 1 1 1 3Theridiidae Phorollcidia sp. 03 1 1 1 3Theridiidae Phorollcidia sp. 04 1 2 1 3 1 8Theridiidae Phorollcidia sp. 05 1 1 1 1 2 1 7Theridiidae PllOrollcidia sp. 06 3 3 1 5 3 4 2 5 3 1 30Theridiidae Sleatoda sp. 01 4 9 4 5 10 4 8 3 4 5 3 5 5 7 2 4 2 4 3 3 3 3 2 102Theridiidae Stealoda sp. 02 1 1 2Theridiidae Steatoda sp. 03 2 1 1 1 1 1 1 I 1 2 12Theridiidae Steatoda sp. 04 1 I 1 3Theridiidae Steatoda sp. 05 1 1 1 1 4Theridiidae Steatoda sp. 06 1 I N-...)

    \i1

  • Table 1 (cont.)

    Family Genus, species Study sites (arranged approximately north-west to south-east) No. ofquadrats

    NO ML ON MO WU WH BE JB VO QU KL MN NR WK KN HV OA UN DU ST PI LK GP ES (n=304)

    N'I0'

    21

    2 1 1 2 1 2 2 3...,I

    12

    TheridiidaeTheridiidaeTheridiidaeTheridiidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeThomisidaeTrochanteriidaeTrochanteriidaeTrochanteriidaeTrochanteriidae

    I .. t

    Steatoda sp. 07Steatoda sp. 08Steatoda sp. 09Steatoda sp. BDiaea sp.Sidymella sp. 01Sidymella sp. 02Sidymella sp. 03Sidymella sp. 04Sidymella sp. 05Sidymella sp. 06Sidymella sp. 08Sidymella sp. 09Sidymella sp. 10Sidymella sp. 11Sidymella sp. 12Stephanopis sp. 01Stephanopis sp. 02Stephanopis sp. 03Stephanopis sp. 04Stephanopis sp. 05Stephanopis sp. 06Stephanopis sp. 07Tharpyna sp. 01Tharpyna sp. 02Tharpyna sp. 03Tharpyna sp. 04Tharpyna sp. 05Tharpyna sp. 06Tharpyna sp. 07Tharpyna sp. 09Tharpyna sp. 10Tharpyna sp. 11Tharpyna sp. 12Tharpyna sp. 13Rebillls sp. 01Trachyspina madllraTrachyspina sp. 02Trachyspina sp. 03

    12 2

    4 1 2

    424

    21

    2

    2

    2

    3

    4 3 3

    2

    1

    21

    4 3

    2

    2

    12

    2

    2

    321

    211231249 s:1

    ';Jl16

    :t1 tJ...

  • Trochanteriidae Traehyspilla sp_ 04 1 I ;p....Zodariidae As/eron-complex sp_ 01 2 1 I 1 1 3 9 '"::lZodariidae As/eroll-complex sp_ 02 2 3 5 4 14

    re0

    Zodariidae As/eroll-complex sp_ 03 1 1 30Zodariidae As/eron-complex sp_ 04 1 3 2 1

    ..., ....I '"0

    Zodariidae As/eron-complex sp_ 05 2 2 ::r'"Zodariidae As/eron-complex sp_ 06 1 I '"00:

    Zodariidae As/eron-complex sp_ 07 1 1 1 3 re....Zodariidae A lls/ralu/ien I)uaerells 3 4 I 1 9 '"Zodariidae Aus/raluliea sp_ 01 1 3 5 6 5 5 3 5 6 7 4 7 2 4 5 3 3 1 1 2 78Zodariidae Aus/ralu/ien sp_ 02 1 1Zodariidae Aus/raluliea sp_ 03 1 IZodariidae Cavas/eron sp_ 01 6 5 6 1 2 2 1 1 1 1 1 1 28Zodariidae Chilwnena reprobans I IZodariidae Chilumena sp_ 01 3 1 1 I 1 7Zodariidae Cyrioctea sp_ 01 I IZodariidae Genus 1 sp_ 01 2 2Zodariidae Genus 1 sp_ 02 1 1 1 2 1 1 7Zodariidae Genus 2 sp_ 01 1 I 1 3Zodariidae Genus 3 sp_ 01 2 2Zodariidae Genus 4 sp_ Ol 3 3Zodariidae Habrolles/es australiensis 1 7 8Zodariidae J-Jabronestes grimwadei 1 1 1 1 1 2 1 1 3 2 14Zodariidae J-Jabronestes sp_ Ol 1 1 1 I 2 2 8Zodariidae Habronestes sp_ 02 2 1 1 5 1 2 2 5 5 5 2 1 1 1 34Zodariidae liabronestes sp_ 03 2 2 5 3 I 8 11 10 8 2 3 8 3 5 7 5 1 84Zodariidae Habronestes sp_ 04 4 4 1 6 1 3 10 2 9 9 4 5 5 4 67Zodariidae Habrollestes sp_ 05 2 1 5 4 3 4 2 1 6 4 2 3 3 1 4 1 2 48Zodariidae Habronestes sp_ 06 1 1 2 1 4 2 1 3 2 2 1 2 1 1 1 1 26Zodariidae J-Jabronestes sp_ 07 1 1 1 1 4Zodariidae Habrones/es sp_ 08 1 1 2Zodariidae Habronestes sp_ 09 4 2 2 3 9 5 6 5 7 6 6 5 3 4 1 3 3 2 76Zodariidae Habronestes sp_ 10 1 1 1 2 2 1 2 1 2 1 2 16Zodariidae Habronestes sp_ 11 1 1Zodariidae Habronestes sp_ 12 2 1 1 1 1 1 1 8Zodariidae Habrones/es sp_ 13 1 4 11 2 1 4 1 24Zodariidae Habronestes sp_ 14 1 2 1 1 1 1 7Zodariidae Ha/Jronestes sp_ 15 3 3Zodariidae Halmmestes sp_ 16 2 1 3Zodariidae Halmmes/es sp_ 17 1 1 2Zodariidae Habronestes sp_ 18 1 2 2 3 8Zodariidae Habronestes sp_ 19 1 IZodariidae Habronestes sp_ 20 1 3 3 7Zodariidae J-Jabronestes sp_ 21 2 2Zodariidae Habronestes sp_ 22 3 3 N

    'I'I

  • Table 1 (cont.) N'-l00

    Family Genus, species Study sites (arranged approximately north-west to south-east) No. ofquadrats

    NO ML DN MO WU WH BE JB YO QU KL MN NR WK KN HY DA UN DU ST PI LK GP ES (n=304)

    Zodariidae Habronestes sp_ 23 1 1 3 2 2 1 1 3 4 1 1 2 4 2 1 29Zodariidae Habronestes sp_ 24 1 3 4Zodariidae Habronestes sp_ 25 1 1Zodariidae Habronestes sp_ 26 1 1Zodariidae Habronestes sp_ 27 1 1Zodariidae Habronestes sp_ 28 1 1Zodariidae Habronestes sp_ 29 1 1Zodariidae Habronestes sp_ 30 1 1Zodariidae Habronestes sp_ 31 1 1Zodariidae Habronestes sp_ 32 2 2Zodariidae Habronestes sp_ 33 1 1 s:Zodariidae Habronestes sp_ 34 1 1

    fJlZodariidae Habronestes sp_ 35 1 1 :tZodariidae HalJronestes sp. 36 1 1 2 III...Zodariidae Hal)ronestes sp. 37 1 1

  • Araneomorph spiders

    In~i 0~ In '02· In

    1""-

    279

    Ln

    Ln cr-; N ("I"')

    - - 01 .~- N 01 In

    Cfj cC) "f

    N -"f - ("'·1 ("1

    % 0'. C' .....-< (,J er; ry In '-.J::; t",,,- '--- ,........ ("'J [',J ("',J \',J r'~ rJ ("'.j (""J

    6..ci..ci.o...o...6...ci..ci..C-D..f f f f f ~ ~ J f f

    ci..o...o...ci.ci-.n..ci.ci..ci..ci..6...J f f f r f r: f f f f

    -0... ...::::r: ~-

    cc:~ r,e.2Z 'JG -

    ,Lj

    ,~ en N

    :J :."r:; r::;

    ""':J

    r::; r::;?j2~~

    '~ Sr ':;.; :J :; '1; ;'.J 'J,.i :J ::lJ :.. :J '"

  • 280 M. S. Harvey, J. M. Waldock, N. A. Guthrie, B. J. Durrant, N. L. McKenzie

    found during the survey, which all appear to differfrom the species revised by Platnick and Forster(1989): ChaslIlocephaloll sp. 1 (DA, JB, UN and YO),ChasllIocephalo/l sp. 2 (DA, JB, NR and UN) andChaslIlocephaloll sp. 3 (DA, ST and UN). Anapidswere generally absent from the northern andeastern quadrats and were more abundant in thesouthern quadrats.

    AraneidaeWe recorded a variety of araneids, but usually in

    very low numbers. The majority of araneid speciesare arboreal and, because they are not reliablycaptured in pitfall traps, were excluded from thequantative analyses.

    CyatholipidaeThe family Cyatholipidae occurs in the majority

    of the southern continents (Forster et al., 1988;Griswold, 2001) and the Australian fauna isrepresented by only a few named genera. The soft-bodied genera such as Tee1l1ellaarus, Tekel/atus andToddiana are generally restricted to temperatewoodland or rainforest habitats, while members ofthe genus Matilda occur throughout mainlandAustralia usually in open woodland habitats.Harvey et al. (2000) found three species of Matildain the Carnarvon Basin region, and this study foundtwo species of this widespread and abundant genus.

    CydoctenidaeCycloctenid spiders are restricted to Australia and

    New Zealand and are vagrant hunters occurring inleaf litter or under stones and rocks. The WesternAustralian fauna consists of small spiders ofuncertain generic affinity. In this survey, Genus 1sp. 1 was found once at DA and Genus 2 sp. 1 wasfound once at LK.

    DeinopidaeMembers of the cosmopolitan family Deinopidae

    construct spectacular webs that they use to ensnaretheir prey (Raven et aI., 2002). A single species ofDeinopis was found sporadically throughout thestudy area at 10 quadrats. Most deinopids aregenerally arboreal and, because they are not reliablycaptured in pitfall traps, were excluded from thequantative analyses.

    DesidaeSpiders of the family Desidae are commonly

    found in a wide variety of Western Australianterrestrial habitats. This survey recorded 39 species,the majority of which were found in few quad rats.

    FilistatidaeFilistatid spiders are found in a variety of regions

    in mainland Australia, and are represented by

    several species of Walldel/a and by a single cave-dwelling species of Yardiel/a (Gray, 1994). Mostspecies of Walldel/a occur under the bark of trees,although Harvey et al. (2000) recorded five speciesin pitfall traps in the Carnarvon Basin regionindicating that some species either live permanentlyon ground habitats (such as in litter or understones) or that they move from tree trunks onto theground at certain times of the year. The presentsurvey recorded Walldel/a barbarella Gray.

    GallieniellidaeThe Australian members of the gnaphosoid

    family Gallieniellidae were recently revised by(Platnick, 2002) who found a small fauna of fivegenera and 27 species. Harvey et al. (2000)encountered five species in the Carnarvon survey,including two species of Meedo. The present surveyrecorded six species, including Meedo houstoni Mainwhich was also found in the Carnarvon survey.

    HahniidaeHahniids are tiny litter or bark dwelling spiders.

    Ten species were found in low numbers during thepresent survey, while no hahniids were found inthe Carnarvon survey (Harvey et al., 2000).

    HersiliidaeH.ersiliid spiders are generally found on tree

    trunks where they lie cryptically in wait for theirprey. Baehr and Baehr (1987,1989, 1992, 1993, 1995,1998) have recorded a total of 25 species of Ta1l1opsisfrom Western Australia, with a further four speciesof Hersilia from northern Western Australia. Ninespecies of Ta1l1opsis were recorded in this survey,most from just a handful of localities, suggestingthat they are infrequently captured in pitfall trapsdue to their arboreal lifestyle. For this reason thefamily was excluded from the quantative analyses.

    LamponidaeThe Lamponidae are restricted to Australia, New

    Caledonia and New Guinea, with two speciesaccidentally introduced into New Zealand; the vastmajority of lamponids are Australian (Platnick,2000). The Lamponidae were revised by Platnick(2000) who recognised numerous Australian generaspecies.

    During this study 41 species in 13 generaencompassing all three subfamilies were identified.The Lamponinae was represented by six genera,Lalllpolla, Lalllpollata, Lalllpollega, Lalllponclll1,Ll1l1lpOllinl1 and Ll1ll1pollusa, and 26 species, of which17 were Ll1l1lpolla species. The Centrothelinae wererepresented by six genera, Asadipus, Bigellditil1,LOllgcpi, NotsodipllS, Prionostemulll and QlICClluic, and14 species. The Pseudolamponinae wererepresented by Pscudoll1l1lpolll1 boree Platnick. The

  • ~ -~-- "'~-----------------------------------------,

    LiocranidaeLiocranid spiders are small to medium sized

    hunting spiders that are moderately diverse in theAustralian region. We recorded nine species, andonly sp. 12 was found at more than 10 sites.

    LinyphiidaeLinyphiids were abundant at many sites, but were

    excluded from the quantitative analyses because ofuncertainties in associating male and femalespecimens and in interpreting the taxonomicsignificance of slight morphological differences inthe structure of the male and female genitalia. Thefamily is largely unstudied in Australia andremaInS an enormous challenge in ecologicalanalvses.

    LycosidaeThe spider family Lycosidae is a diverse group

    that is poorly known in Australia. The generaArtoria and \/ellatnx have been recentlv revised(Framenau and Vink, 2001; Framenau, 2(02) but theremaining genera are still in flux. The vast majorityof the Australian wolf spider fauna has beenattributed to the genus Lycosa but these species bearlittle affinity to the type species of Lycosa (Vink etaI., 2(02). The systematics of the Australian wolfspiders is currently being revised (Framenau, pers.comm.). We identified 61 lycosid species in thepresent survey, but the case for some of theidentifications is tenuous and the fauna may besmaller.

    Araneomorph spiders

    genera Pnollostemllm andrestricted to the southern quadrats.

    )/11'1'117)11' were

    281

    consistent with the broad distributional rangerecorded Harvey (1995). Ambicodamus marae wasonly found at DA07, JBI0 and LKI2, eachrepresenting range extensions for this species whichwas previously found to be restricted to highrainfall areas, predominately in the karri forests ofthe south coast (Harvey, 1995) and in outlyingpopulations in the Darling Range (Brennan, 1999).Ambicodamlls kochi was found at NOlI.

    OecobiidaeThe spider family Oecobiidae is represented in

    the Australian region by introduced species (seeSantos and Conzaga, 2(03), and this study foundOecobws Ill/ViiS at several sites. Unfortunately thisspecies was coded as "Annulipes sp. 1" and"Annulipes sp. 2" in the analysis.

    OonopidaeThe tiny spiders of the family Oonopidae are an

    extremely abundant and diverse component of theAustralian fauna, and we found 46 species in avarietv of genera including Cmnasomorpha, Cn/meus,iV1:11/71l0!'70r7aea, Opopaea, Orchestilla and a new genus.

    OrsolobidaeThe Condwanan family Orsolobidae is

    represented in the Australian fauna by members ofseveral genera, of which Tasmanoonops andAlIstraloblls have been found in Western Australia(Forster and Platnick, 1985). Further unnamedspecies of both genera are known (Harvey,unpublished data). A single specimen of anunnamed species was found at WH02.

    Micropholcommatidae'The tiny spiders comprising the family

    Micropholcommatidae are abundant in Australianforests but have largely remained unstudied exceptfor the work of Hickman (1944, 1945, 1979) andForster (1959). We found several species tentativelyplaced in the genus Micropholcolllma.

    MysmenidaeThe tinv spiders of the Mysmenidae are only

    rarely collected in pitfall traps, and only two specieswere found in this survey, each from a single site.

    NicodamidaeThe Nicodamidae were revised by Harvey

    who found two genera in New Zealand and fivegenera in Australia, one of which also occurs inNe,v Cuinea. The Western Australian fauna isrepresented by three species, Nicodamlls maillaeHarvey, Ambicodamus marae Harvey and A kochiHarvev. Nlcodamus maillae was found to beWidespread throughout the survey area, which is

    PararchaeidaeThe family Pararchaeidae is restricted to Australia

    and New Zealand where they occur in forested andwoodland habitats. Two unnamed species ofPararchaea were found in the survey. Pararchaea sp.1 was found at one quadrat whilst Pararchaea sp. 2was found at a variety of sites throughout thesurvey area.

    PhilodromidaeThe Australian philodrornid fauna is small (Raven

    et aI., 2(02) with just a handful of recognised speciesin two genera, Philodromlls and Tibellus. Werecognised four species of Tibellus in the presentsurvey, each collected in very low numbers,presumably due to their primarily arborealpreferences.

    SaIticidaeThe Salticidae are the largest spider family

    (Platnick, 20(4) and the Australian fauna isimperfectly known (Davies and Zabka, 1989). Werecorded 121 species in 41 genera, of which the vast

  • 282 M. S. Harvey, J. M. Waldock, N. A. Guthrie, B. J. Durrant, N. L. McKenzie

    majority are undescribed. The Salticidae were thesubject of a separate analysis in this volume(Guthrie and Waldock, 2004).

    SegestriidaeOnly three species of Segestriidae were

    recognised, in contrast to the seven species found inthe Carnarvon survey.

    StiphidiidaeThe Australasian spider family Stiphidiidae was

    represented in the survey by a single species ofBaiami, three species of Corasoides, a species ofForsterina, and two species tentatively placed inCorasoides.

    TetrablemmidaeThe spider family Tetrablemmidae is currently

    represented within Australia by a single namedspecies, Tetrablemma okei from Victoria, butnumerous unnamed species occur in eastern andnorthern Australia (Harvey, unpublished data). Asingle female tetrablemmid was collected at M002,which represents the first record of this family fromsouthern Western Australia.

    TetragnathidaeTetragnathid spiders are represented in south-

    western Australia by several genera such as Nephilaand Tetragnatha, as well as a series of poorly knownmetines. This survey found two such metine speciesin low numbers.

    TheridiidaeTheridiid spiders are a ubiquitous part of the

    spider fauna of most regions of the world, and werecorded 80 species.

    ThomisidaeWe recorded 31 species of Thomisidae during the

    survey, primarily in the genera Sidymella, Stephanopisand Tharpyna. As the majority of these species areconsidered to be arboreal or corticolous and, becausethey are not reliably captured in pitfall traps, theywere excluded from the quantative analyses.

    TrochanteriidaePlatnick (2002) has recorded a rich trochanteriid

    fauna in Australia, represented by 14 indigenousgenera. We found five species of this family,including four species of Trachyspina and a singlespecies of Rebilus, each from a single quadrat.

    ZodariidaeRecent systematic research on the Australian

    zodariid fauna has revealed a highly diverse faunaconsisting of numerous genera (e.g. Jocque andBaehr, 1992, 2001; Jocque, 1995a,b; Baehr, 2003a,b,in press; Baehr and Jocque, 2000, 2001; Jocque andBaehr, 2001; Baehr and Churchill, 2003;). The 117species of Zodariidae recorded during the surveywere the subject of a separate analysis in thisvolume (Durrant, 2004), in which there was nodiscernible relationships between speciescomposition and substrate data.

    32 r-....----.,----.---.,..---.....--..,----.,.---.---,..---.....--....---.

    ~l1Jc 28 24~

    l'(/)

    11'1l1J·0 27 27 16l1J 24Et l' 20r- i'Cl.L-0E 20 260l1JC~

    (ij

    O'i 16l1JL- 160.ClL-~,c 120z IH =48.3, n =294, df =11, P « 0.0001

    82 3 4 5 6 7 8 9 10 11 12

    Landform unit

    Figure 3 Relationship between quadrat species richness and landform unit. Mean and 95% confidence intervals aredisplayed. Means are labelled with the number of quadrats sampled in each landform. H = Kruskall-Walliscoefficient, n = number of quadrats, df = degrees of freedom, and P = probability. Because of trap-floodingproblems, data from landform unit 1 (freshwater swamps) are excluded.

  • AraneOlIlorph spiders

    RichnessA total of 622 ground-dwelling araneomorph

    spider species were recorded from the 304 quadrats,an average of 21.9 species per quadrat (s.d. = 8.3).Figure 3 provides a graphical representation ofspecies richness within each landfonn unit. Naturalsaltflats (unit 2) had the fewest species, whiledissection valley slopes (units 4 to 8) and durierustpavements (unit 12) were the richest.

    The relationship between salinity risk (SAL, as ameasure of secondary salinity) and species richnesswas quantified for the ground-dwellingaraneomprphs overalL and for the three mostspeciose families separately: Salticidae (121 species),Zodariidae (117) and Lycosidae (61). Together,these comprised 48(};) of the araneomorph speciesrecorded during the survey. This analysis wasrestricted to the 55 quadrats sampled on landformunit 3 (dissection valley floors), the unit withsufficient quadrats sampled in each of the foursalinity risk categories to offer a balancedcomparison (McKenzie et 2003). Together thethree families analysed comprised 47% of thearaneomorph richness recorded on landfonn unit 3.Salticids and zodariids showed a significantnegative correlation with electrical conductivity,while lvcosids showed a significant positive

    283

    correlation (Kendals Tau -0.26, p = 0.006; r -0.37,P 0.0001; r 0.31, P 0.001, respectively). Asignificant negative correlation was also foundwhen all araneomorph families, except lycosidae,were included (r = -0.27, P 0.004). Althoughrichness was tightly correlated with numer ofspecimens (r = 0.56, P < 0.0001), the latter was notsignificantly correlated with salinity risk. Thissuggests that the significance of the salinity riskcorrelation was related to species richness ratherthan being an artefact of the relatively low numbersof identified specimens per quadrat (87, s.d. = 60, n

    55). Scatter-plots of species richness versus ECshowed that individual outlying points did notdetermine the significance of these correlations.Using a larger data-set of arachnids and vertebratesthat encompassed the araneomorph data beinginvestigated in this paper, McKenzie et al. (2003)showed that these secondarily salt-affected quadratsheld a species-poor sub-set of the assemblagesrecorded on their unaffected counterparts, as wellas a few natural salt-flat species.

    Singleton species and local endemismOne hundred and eighty-five species of

    araneomorph spiders were recorded at only onequadrat, representing 29.7°!cl of the entire data-set.

    BEWU 1-DN I 1 1ML 1-1- north band except JBWH 1-MO INO 1-HY east central IKN I IKL I 2 IQU I-- I 2 Central BandMN I I INR west central ,WK 3 ,YO -I ,DA 1 IDU I IJB 1 IST 4 I 3 Southern Band + JBIGP 1 ,LK I I IPI -I I I IUN 11- -I ,ES 5_1 1_4_1 I

    I I 1 I 1 I I0.19 0.26 0.32 0.39 0.46 0.52

    Coefficient of Dis-similarity

    Figure 4 Dendrogram structure derIved by classifying the 24 survey areas according to the combined generacornpC)Sltlon of their component quad rats (a total of 166 non-arboreal araneomorph genera). Singleton specieswere retamed in the 304 quadrat versus 622 species matrix from which the survey area genera lists werederived. Genera that were recorded m only a single survey area were included in the analysis.

  • 284 M. S. Harvey, J. M. Waldock, N. A. Guthrie, B. J. Durrant, N. L. McKenzie

    Figure 5 The four partitions in Figure 4 were most clearly separated by warmest period maximum temperature. Meanand 95% confidence intervals are based on climatic values averaged across the 12 to 13 quadrats within eachsurvey area (H ; Kruskall-Wallis coefficient, n ; number of quadrats, df degrees of freedom, and P ;probability).

    On average, each quadrat had only one singleton(sd = 1.01, n = 304). Of the 400 non-singleton speciesrecorded on the 240 quadrats that were bothunaffected by secondary salinity and adequatelysampled, 38% (152/400) were more geographicallylocalised than would be expected by chance alone(P < 0.05), compared to 1000 trials using anequivalent number of randomly arrayed records(quadrat intersections), randomly arrayed, for eachspecies. Overall, 65% of species (261/400) wererecorded at between 2 and 10 quadrats and 26% ofthese (67/261) were more geographically localisedthan would be expected by chance (P < 0.05). These67 species were recorded in an average of 3.3 surveyareas (s.d. = 1.9), and most were recorded only inthe same or adjacent survey areas (51/67 species).This should not be interpreted as unambiguouslyimplying that all 67 species are locally endemic.Records of 66 of the 67 species extended into surveyareas on the periphery of the study area and furthersurvey work will probably show that many of thesespecies extend beyond this periphery. Because ofthe elongate shape of the study area (Figure 1), 18of the 24 survey areas were peripheral.

    Patterns in genus compositionBy classifying survey areas in terms of their

    genera composition we could minimise the effectsof species endemism and substrate, and reduce theinfluence of study area edge effects and samplingnoise, on the partition structure of the resultingdendrogram. The compositional pattern, assummarised by the resulting dendrogram structure

    at the four partition-level (Figure 4), was closest tothe warmest period maximum temperature (seeFigure 5). At the five partition-level significantseparation of all partitions was best achieved by acombination of temperature diurnal range andwarmest quarter mean temperature (Table 2).Together, these two climatic attributes yielded aclimatic topography and axis only slightly differentfrom warmest period maximum temperature.

    To see whether substrate attributes emerged assignificant correlates with compositional patterns inaraneomorph genera, the 240 quadrats wereclassified in terms of similarities in the presence andabsence of their 162 araneomorph genera. The fourgenera that occurred at only a single quadrat areexcluded from this analysis. The resultingdendrogram structure could be interpreted down tothe seven partition-level, where six of the sevengroups could be separated statistically using acombination of four quadrat attributes: SAL(salinity risk), soil pH, warmest quarter meantemperature and precipitation seasonality (Table 2).Group 5 could not be separated uniquely, butcomprised just three of the 240 quadrats.

    Patterns in species compositionFigure 6 is the dendrogram structure derived

    when the survey areas were classified according totheir total araneomorph species composition. Thisdendrogram reveals a strong geographical patternrelated to rainfall and temperature gradients. Whenthe structure was examined statistically againstsurvey area climatic attributes, the tightest

  • Araneomorph spiders

    BE I 1ML --I-- I IMO I I North-east IWU I 1- I I 1 NorthWH I I INO I I_NW extreme IDN __NW_I IHY 1K.N 1KL I Central-east 1QU 1- 1MN I I 2 CentralJB I INR Central-west IYO I IWK I 1--1DA 1ST South-west 1UN 1DU I I 3 SouthLK South-central IPI & east I IGP 1- I IES SE extreme___ 1_4____ 1

    I I I 1 I0.39 0.46 o 54 0.61 0.69 0.76

    Coefficient of Dis-similarity

    285

    Figure 6 Dendrogram structure derived by classifying the 24 survey areas according to the combined speciescomposition of their component quad rats. Singleton species were retained in the 304 quadrat versus 622species matrix from which the survey area species lists were derived.

    relationship was with warmest period maximumtemperature (Table 2), a north-north-east to south-south-west axis related to the hotter summers in thenorth-north-east (Figure 7).

    When the 240 quad rats were classified in terms of

    similarities in their species composition, thedendrogram could be interpreted down to theeight-partition level (Figure 8). All partitions couldbe separated statistically using a combination offour quadrat attributes: soil salinity, soil pH,

    Table 2 Environmental attributes that best separated the classification partition structures derived from the fourcompositional analyses (A survey areas, C = genera, Q = quadrats, S species, I1 = Kruskall-Walliscoefficient, P = probability, mxTwml' = warmest period maximum tt'mperature'C, Tdir temperature diurnalrange, TwmQ W

  • 286 M. S. Harvey, J. M. Waldock, N. A. Guthrie, B. J. Durrant, N. L. McKenzie

    Warmest Period Maximum Temperature-28

    -29

    -

  • Araneomorph spiders 287

    BEOI-5,7-9,12,13, DNOI,2,6,8

    KNIO, MLOI-13, MN03,4,7,1O,

    MOOI,3,5,6,9-12, NOOI-7,1O,11

    PI05, QU04, WH04,5,7,8,11,13

    WUOI-6,9-12, Y002,1O

    DN05, DU02-4,9-13, ES07,9

    GPOI-4,6,12, HY01,3-7,9,1O

    JBOl,2,5-IO,12,13, KL03 -12

    KN02-6, 11, LK02,4-6,8, 10-13

    MN01,5,6,11, NROI-4,6,8-12

    PIOI,6-9,12, QUOl-3,6,9-12

    ST09,10, WH03,6,9,IO,

    WKOI,6-11,13, Y001,4-8,12,13

    DN03,4,9-12

    111 1

    11

    -1-IIII 2111

    1 1-1--

    3

    6

    4

    5

    1

    1

    III111

    1 I

    -------1-1-----I II 7 1

    1 1-1----- I_1_8 1

    1 I I1 1.24 1.52 1.79

    Dis-sirnilarity Coefficient

    DAI2, ESOI,2,5,8,11

    GP13, PI13, ST02-5,7

    UNO I,2,5-7,9-12

    GPIO, PIll

    DAOI,3,4,6,8-11,13, DUOI

    GP08,9, LK07, NR05,7

    STOI ,6, 13, UN03, Y003

    GP05

    BE 11, DU05, ES06, HY02,

    KLOI, LK03, MLl2, MN09,

    M004, ST08, WU08

    Figure 8 Dendrogrum derived cbssifymg 240 quudruts uccording to their species composition. Structure to the 8-partition level IS shown. Matrix was presence/ubsence data for 400 species und excluded singleton species,fresh-'Nater swamps (LF 1), severely salt-affected quudrats (SAL 3 and 4) other thun natural salt flats (LF2), and the three that were inunduted for long periods during the trup program (PI04, UN13 undWK02)

    axis. As pointed out in Results, these two climaticattributes in combination yield a climatictopography and axis only slightly different fromwarmest period maximum temperature. However,the climatic components of this pattern could alsobe explained in terms of broadscale landscaperelationships. As pointed out in Methods, the ESand part of the ep survey areas are not actually onthe Yilgarn Craton; nor are NO and DN in thenorth. Furthermore, Mulcahy and liingston (1961)divided the Craton in the central part of the studyarea into eastern and western zones because the

    Tertiary Plateau is much more dissected in the westdue to their different paleoclimatic histories. Also,the western zone has a rejuvenated drainage systemthat flows westwards, while the eastern zone'sancient drainage system comprises chains of saltlakes and follows Tertiary paleodrainage lines thatare orientated northwards and/or eastwards (e.g.Beard, 2000). These broad landscape differencesmore or less correspond to the dendrogrampartitions analysed (rable 2); deductive studiessuch as this do not eliminate alternative hypotheses(Oksanen, 2001).

  • 288 M. S. Harvey, J. M. Waldock, N. A. Guthrie, B. J. Durrant, N. L. McKenzie

    It was clear that the saltflats support an array ofspecialist halophilic araneomorphs (species group18 in Appendix 4). This is not surprising becausegroundwater dating in south-western Australia(Commander et ai, 1994) and sedimentary studieson gypsum accumulation in Lake Lefroy by Zhenget al (1998) suggest that most halite lakes have beensaline for up to a million years (and earlier phasesof salt lake formation in this old and geologicallystable landscape are probable).

    This study revealed an inverse relationship(correlation) between araneomorph species richnessand soil salinity, both natural salinity and thesecondary salinization induced by agriculturalclearing during the last 100 years. Given that salt-affected sites in our study contained acompositional sub-set of their unaffectedcounterparts (McKenzie et al., 2003), the effect ofsecondary salinity on araneomorph richness (andcomposition) is essentially negative. While thearaneomorphs belonging to the family Lycosidae(wolf spiders) showed a positive correlation, therelationship is probably not causal. Individuallycosid species, both in Australia and elsewhere, areoften confined to particular habitats such as riverinemargins, sand dunes and salt lakes (McKay, 1979;Hudson and Adams, 1996; Moring and Stewart,1994). Generally, Main (2001) characterised lycosidsas adaptive opportunists that favour openenvironments, and predicted that the family wouldincrease in lands cleared for agriculture. As withclearing, rising ground-water salinity results in anet loss of vegetation cover and, in the wheatbelt,our data suggest that these conditions favourlycosids, at least initially. Thus, salt affected sites(mainly the woodland sites on the dissection valleyfloor, landform 3) hold their lycosid diversity and,at least in the initial stages of secondary salinity,may be further enriched through colonisation bysaltflat specialist species such as Lycosa salifodina.

    The only substrate patterns that emerged dearlyfrom the quadrat analyses were related to salinityattributes. No distinct, study area-wide patterns incomposition emerged that related to other substrateattributes (e.g. soil texture, chemical attributes etc).A plausible explanation is that local endemism, asnoted in the salticid and zodariid components(Durrant, 2004; Guthrie and Waldock, 2004),perhaps exaggerated by local covariance inpopulation densities etc, masked any suchsubtleties.

    Broader relationships with soil-landscape zonesand climatic gradients were discussed earlier. Inany case, clear-cut patterns should not be expectedbecause a species' presence or absence at a site isactually determined by physiological limits andavailability of suitable resources in combinationwith guild, predation and other ecologicalprocesses. Rosensweig (1992) pointed out that

    species define habitats, not vice versa. Correlationswith environmental parameters such as thoserevealed in this study, are indirect consequences ofspecies niche requirements rather than directlycausal to their presence/absence at sites.

    ACKNOWLEDGEMENTS

    We thank J.K. Rolfe, W.P. Muir, P. van Heurckand N. Hall for assistance in installation andoperation of the pit traps; L. King, E. Ladhams andP. van Heurck for assistance with sorting, datacompilation and field sampling; J.K. Rolfe for database development and interrogation, B. Baehr forassistance in the identification of some Zodariidae;and M.R. Williams and B. Brand for statisticaladvice. We thank two anonymous referees for theircomments on the manuscript. Funding for thisstudy was provided as part of the State SalinityStrategy.

    REFERENCES

    Austin, M.P. (1984). Problems of vegetation analysis fornature conservation. In Myers, K. and Margules, eR(eds), Survey methods for nature conservation, vo!. 1:101-130. CSIRO Division of Water and LandResources, Canberra.

    Austin, M.P. (1991). Vegetation theory in relation to cost-efficient survey. In Margules, CR. and Austin, M.P.(eds), Nature conservation: cost effective biological surveysand data analysis: 17-22. CSIRO Division of Wildlifeand Ecology, Canberra.

    Austin, M.P. and McKenzie, N.L. (1988). Data analysis.In Gunn, RH., Beattie, ].A., Reid, RE. and van derGraaff, RH.M. (eds), Australian soil and land surveyhandbook: guidelines for conducting surveys: 210-232.lnkata Press, Melbourne.

    Baehr, B. (2003a). Revision of the tropical genusTropasteron gen. novo of north Queensland (Araneae,Zodariidae): a new genus of the Asteron-complex.Memoirs of the Queensland Museum 49: 29-64.

    Baehr, B. (2003b). Three new endemic genera of theAsteron-complex (Araneae: Zodariidae) fromAustralia: Basasteron, Euasteron and Spinasteron.Memoirs of the Queensland Museum 49(1): 1-27.

    Baehr, B. (in press). Revision of the new Australian genusHolasteron (Araneae, Zodariidae): taxonomy,phylogeny and biogeography. Memoirs of theQueensland Museum.

    Baehr, B. and Baehr, M. (1987). The AustralianHersiliidae (Arachnida: Araneae): taxonomy,phylogeny, zoogeography. Invertebrate Taxonomy 1:351-437.

    Baehr, B. and Baehr, M. (1989). Three new species ofgenus Tamopsis Baehr & Baehr from WesternAustralia (Arachnida, Araneae, Hersiliidae). Secondsupplement to the revision of the AustralianHersiliidae. Records of the Western Australian Museum14: 309-320.

    Baehr, B. and Baehr, M. (1992). New species and new

  • CSIRO,

    Araneomorph spiders

    recmds ot genus B,ll'hr & B'll'hr (Arachmda,Ar,1Iw, R. (2000), Revisions of genera in the/ble}(lll~complcx (Araneae: Zodariidae), The newgenera Cm'l/,;leroll and Milll/,;Ieroll. Records of lIleWe.slem A/lslmliall Muse/lll/ 20: 1-30.

    Baehr, B. and Jocqut>, R, (2001). Revisions of genera in theAsleroil~complex (Araneae: Zodariidae): new generaPell la,; It'roil, Pheilasleroil, Lel'lasll'roll and S/lIJI/Sleroil.,Mell/oirs of the Q/lel'II,;laild M./lSeUIII 46: 359-385.

    Baxter, J.L. and Lipple, S.L. (1985), Perell!ori, We,;temA/I,;lmlia 1:250 ()()O seolosiclIl ,;eries. Geological Surveyof Western Australia, Perth.

    Beard, J5. (19S0). A new phytogeographic map forWestern Australia, Weslel'll Allslmliall Ilerl)lIri/l1l/Re,;earch Nole,; 3: 37-5S.

    Beard, J.5. (19SI), Vesetatioll ';/lri'elj of We,;lt'm A/I,;tmlia~·Swail 1 1 000 O()() series. Universitv ofWestern Australian Press, Nedlands.

    Beard, J.5. (1990). 1'111111 lit" of ~Vl'slem A list 1'171 ill. KangarooPress, Kenthu rst.

    Beard, J.5. (2000). Drainage evolution in the MooreMongn system, Western Australia.lollrJ/al of Ihe 1\01/111

    of Wes/('rJ/ AII,;lmlill 83: 29-3S,

    Belbin, L. (1995). PATN Il'(//I/lcalCanbnra.

    Brennan, K. E.C. (I IJiscoven of the spiderAII/IJicodlllllll'; IIlame (Araneae: Nicodamidae) in thenorthern jarrah forest of \Nestern Australia. Rl'(ords ofthl' Wl'slem Allslmlillil All1sellll/19: 323-325.

    Brennan, K.E.C. and Majer, J.D. (in press). Conservationand biodiversitv of spiders in Western Australianjarrah forest: untangling multiple disturbance effectsfrom burning and mining. III Lunney, D. (ed.),C)II';I'l'i'allOlI of Allslmlill''; forest Falllll/, 2,,,1 edn. RoyalZoological Socidy of New South Wait's, Svdney.

    Brennan, K.E.C., Majn, J.D. and Moir, !'v1.l .. (in

    289

    Biodiversitv of jarrah torest spidl'l's at larr'lhdalc',Western Australia: richness, endemicit\, andtaxonomic knowledge. COIISl'n'lltltlll [i1l)lo,,,I;'

    Bureau ot ML'tl'orologv (2001). What is till' wl'atherusu,lllv like) Climate averages fnr We,;tern Australiansitl's. htt ww.bom.gov.,lu/clim,ltc/averatablcs/ca_wananws,sht ml.

    Chin, RJ (19Sh). Kt'llerl)arill, Wl'stem AlIstmllll, I 25()()(I(I .sail'.s. C;eologic,ll Survey ot Wl'slL'rnAustr'llia, Perth.

    CIMke, K.I\. (J993), Non~p,H,lmetric multivariate'lnalvses of changes in communit" structure,Au.sfmlillll 10uI'II1l1 of 6: 163-17-1.

    Commander, [).I'., Fifield, L.K., Thorpe, I'M, IJavie, 1\.1'.,Bird, J,I\. andlurIlL'r, J.V. (1994). Chlorine~36 andcarbon~1-1 measurements on hvpersaline groundwaterin Tertiarv paleochannels near Kalgoorlie, WesternAustralia. Professional I'apers 37: .53·~60. GeologicalSurvey of Western Australia, Perth,

    Commander, D.I'., 5choknecht, N" Verboom, W, andCaccetta, I) (2002). The geology, physiology and soilsof wheatbelt valleys. 111 V. Read & Associates (eds),Deillillg wilh slllillilll ill iclIClll/)c!1 ProICSSI'';,I'rosl'cct,; lllld I'mlliml 0l'tiOIl';: 1-21, Waters and RiversCommission, Perth (Available on CD),

    Czekanowski, J. (1932). Coefficient of racial likeness, unddurchschnittliche differenz. AlIlhrol'ologis(/Ii'l' flll_CI""9: 227-249.

    Davies, V.'F. (1994), The huntsman spiders flctcrol'odaLatreille and Yiilllhi gen. novo (Araneae:Heteropodidae) in Australia, Mellloirs of IlleQueellsllllld MuseulII 35: 75-122,

    Davies, V.T. and Zabka, M. (19S9). lIlustrated keys to thegenera of jumping spiders (Araneae: Salticidae) inAustralia. l\:lclllOirs of' fhe QueclIslnlld Musc/l1I/ 27: IS9-266,

    Durrant, B.J. (2004), Biogeographical patterns of zodariidspiders (Araneae: Zodariidae) in the wheatbeltregion, Western Australia. RClord,; of' Ihe WesfernAuslmlinll AlusCU1I1 S/lI'I'ICIIIClIt 67: 217-230

    Durrant, B.J. and Guthrie, N.A. (2004). Faunas ofunflooded saline wetland floors of the WeskrnAustralian wheatbelt. [\ccords of Ihc Wcstern AustmliallMuseulII S/ll'l'lclllclIl 67: 231-256.

    Environment Australia (2000). I\l'i'isioll of'th,' illlerillll>iogcogral'hil rcgiollilli';llfioll Auslrlllin I/BI\A) nllddCi'c!ol'lIlelll of' Z'('/',;ioll 1. Environment Australia,Canberra.

    Faith, D.I), Minchin, 1',1\. and Belbin, L. (19S7).Compostional dissimilarity as a robust measure ofl'cological distance: a theoretical model and computersimulations, 69: 57-6S.

    Forslcr, R.R. (1959). The spiders of the famil"Symphvtognathidae. TmllSllc'liolls of thc R()jlnlof NCil' Zellllllld 86: 269-329.

    Fmster, R.R., Millidge, A.F, and Court, DJ (19SS). Thespiders of New Zealand. Part VI. Mu'eUIII13u/lcllII 6: 1-124,

    Forster, R.R. and Platnick, N.1. (1985). A review of theaustral spider family Orsolobidae (Arachnida,Araneae). with nntes on the super!amih'Dysderoid,'a. Bulletlll of tllC A,llcri(llll MuseUl1I ofNlltural 181: I-DO.

  • 290 M. S. Harvey, J. M. Waldock, N. A. Guthrie, B. J. Ourrant, N. L. McKenzie

    Framenau, V.W. (2002). I{evision of the wolf spider genusArtoria Thorell (Araneae: Lycosidae). IlluertelnateSystelllatics 16: 209-235.

    Framenau, V.W. and Vink, Cl. (2001). Revision of thewolf spider genus Vellatrix Roewer (Araneae:Lycosidae). Illuertebrate TaxoIIOII1l/15: 927-970.

    George, Rand Coleman, M. (2002). Hidden menace oropportunity - groundwater hydrology, playas andcommercial options for salinity in wheatbelt valleys.III V. Read & Associates (eds), Dealillg with salillilt/ illwheatbelt valleys. Processes, prospects alld practicaloptiolls: 1-21. Waters and Rivers Commission, Perth(Available on CD).

    George, R.J., McFarlane, D.J. and Speed, RJ. (\995). Theconsequences of a changing hydrologic environmentfor native vegetation in Western Australia. IIISaunders, OS, Craig, J.L. and Mattiske, E.M. (eds),The role of lIe/works: 9-22. Surrey Beatty & Sons:Chipping Norton, NSW.

    Cray, M.R. (\994). A review of the filistatid spiders(Araneae: Filistatidae) of Australia. Records of theAustraliall Museulll 46: 39-61.

    Griswold, CE. (2001). A monograph of the living worldgenera and Afrotropical species of cyatholipid spiders(Araneae, Orbiculariae, Araneoidea, Cyatholipidae).Mellloirs ol!he Calif()rnia AcadelllY ofSciellces 26: 1-251.

    Guthrie, N.A. and Waldock, J.M. (2004). Patterns in thecomposition of the jumping spider (Arachnida:Araneae: Salticidae) assemblage from the Wheatbeltregion, Western Australia. Records of the WesternAustraliall Museulll, Supplelllellt67: 203-216.

    flarvey, MS (1995). The systematics of the spider familyNicodamidae (Araneae: Amaurobioidea). IllvertebrateTaxollolllY 9: 279-386.

    Harvey, M.5. (2002). Short-range endemism in theAustralian fauna: some examples from non-marineenvironments. Illvertebrate Systelllatics 16: 555-570.

    Harvey, MS, Sampey, A., West, P.L.J. and Waldock, J.M.(2000). Araneomorph spiders from the southernCarnarvon Basin, Western Australia: a considerationof regional biogeographic relationships. Records of theWestern Australiall Museulll Supplelllellt 61: 295-321.

    Hatton, T.J. and Ruprecht, J. (2001). Watching the riversflow: hydrology of the wheatbelt. III V. Read &Associates (eds), Dealillg with salillity ill wheatbeltmllel/s. Processes, prospects alld practical OptiOIlS: 1-15.Waters and Rivers Commission, Perth (Available onCD).

    Hickman, V.V. (1944). On some new AustralianApneumomorphae with notes on their respiratorysystems. Papers alld Proceedillgs of the ROlfal Society ofTaslllallial943: 179-195.

    Hickman, V.V. (\945). A new group of apneumonespiders. Trallsactiolls of the COllllecticut AmdelllY of Artsalld Scieuces 36: 135-157.

    f1ickman, V.V. (1979). Some Tasmanian spiders of thefamilies Oonopidae, Anapidae and Mysmenidae.Papers alld Proceedillgs of the Royal Society of Taslllallia113: 53-79.

    Hirst, 0.13. (1989). A revision of the genus Pedialla Simon(Heteropodidae: Araneae) in Australia. I{ecords of theSouth Australiall Museulll 23: 113-126.

    L _

    Hirst, 0.13. (199Ia). Revision of Australian species of thegenus F!olcollia Thorell (Heteropodidae: Araneae).I{ecords of the South Australiall Museulll 24: 9 I-109.

    f-Iirst, 0.13. (1991 b). Revision of the Australian generaEodclella Hogg and Zachria L. Koch (Heteropodidae:Araneae). Records of the South Allstraliall lvluseulll 24:91-109.

    Hirst, 0.13. (1992). Revision of the genus Isopeda Koch(Heteropodidae: Araneae) in Australia. IllvertebrateTaxollolllY 6: 337-387.

    Hopper, S.D., Harvey, MS, Chappill, J.A., Main, A.R.and Main, B.Y. (\996). The Western Australian biotaas Gondwanan heritage a review. III Hopper, 5.0.,Chappill, J.A, Harvey, M.5. and George, AS (eds),COlldwallall heritage: past, presellt alld future of theWestern Australiall biota: 1-46. Surrey 13eatty & Sons,Sydnev.

    Hudson, P. and Adams, M. (1996). Allozymecharacteristics of the salt lake spiders (Lycosa:Lycosidae: Araneae) of southern Australia: systematicand population genetic implications. Austl'l1liallJournal of Zoology 44: 535-567.

    Jocque, R. (1995a). Notes on Australian Zodariidae(Araneae), I. New taxa and key to the genera. Recordsof thc Australiall MUSCUIII 47: 117-140.

    Jocque, R. (1995b). Notes on Australian Zodariidae(Araneae), 11. Redescriptions and new records. Recordsol!hc Australiall Musculll 47: 141-160.

    Jocque, R. and Baehr, B. (2OCJI). Revisions of genera in theAstcroll-complex (Araneae: Zodariidae). AstcrollJocque and the new genus Pseudas!eroll. Records of theAustraliall M USCUIll 53: 21-36.

    Jocque, R. and 13aehr, M. (1992). A revision of theAustralian spider genus Storella (Araneae:Zodariidae). Illvertcbrate TaxollolllY 6: 953-1004.

    Main, B.Y. (1991). Kimberley spiders: rainforeststrongholds. III McKenzie, N.L., Johnston, R13. andKendrick, P.C. (eds), Killlberley raillforests: 271-293.Surrey 13eatty & Sons, Sydney.

    Main, B.Y. (2001). Historical ecology, responses to currentecological changes and conservation of Australianspiders. Journal of III sect COllserl1atioll 5: 9-25.

    Main, B.Y., Sampey, A. and West, P.L.J. (2000).Mygalomorph spiders of the southern CarnarvonBasin, Western Australia. Records of the WesternAustraliall Museulll Supplelllcllt 61: 281-293.

    McKay, R.J. (1979). The wolf spiders of Australia(Araneae: Lycosidae): 12. Descriptions of someWestern Australian species. Mellloirs of the QueellslalldMuseulll19: 241-275.

    McKenzie, N.L., Burbidge, A.H. and Rolfe, J.K. (2003).Effect of salinity on small, ground-dwelling animalsin the Western Australian wheatbelt. Australiall/ourllal of Botally 51: 725-740.

    McKenzie, N.L., Gibson, N., Keighery, G.J. and Rolfe, J.K.(2004). Patterns in the biodiversity of terrestrialenvironments in the Western Australian wheatbelt.Records of the Western Australiall Museulll Supplelllellt67: 293-335.

    McKenzie, N.L., Robinson, A.C and 13elbin, D.L. (1991).Biogeographic survey of the Nullarbor district,Australia. III Margules, CR. and Austin, M.I'. (eds),

  • Araneol11orph spiders 291

    Rdven, R[, Baehr, B,C and Ilarvev, MS (2002)of Australia: interactive identification to ~ubfamily.CSIRO/ABRS, Melbourne,

    Rosensweig, M,I.. (1992). Species diverSity gradIents: wcknow more and less than wc thought. Jo 11 rI/al ofMammalogy 73: 715-730,

    Santos, AJ. and Gonzaga, M.o. (2003) On the spIdergenus Oecobills Lucas, 1846 In South AmerIca(Araneae, Oecobiidae), JOllrnal of Natllml Histon! 37:239-252.

    Schoknecht, N, (2002). Soil groups of We~tern Au~tralia,edn 3, Resource Management Technical Report 246,Department of Agriculture, Perth,

    Sll110n, E. (1908), Araneae, 1" partie. [n Michaelsen, W.and Hartmeyer, R, (eds), Dlt' Fauna lidwest-AlIstmliens, vof. I: 359-446. Custav Fischer, Jena.

    Simon, E, (1909). Araneae, 2me partie, [n Michaelsen, W.and Hartmeyer, R. (eds), Die Fauna Si/dwest-AlIstmliens, vof. 2: 155-212, Custav Fischer, lena,

    Sneath, P.H,A. and Sokal, R,R, (1973), Nllmericaltaxonomy: the principles and practice or nlllllericalclassification. Freeman, San Francisco,

    Statsoft (2001). STA TISTICA System Rererence Statsoft,Tulsa, USA.

    Vink, Cl., Mitchell, AD, and Paterson, AM, (2002) Amolecular of phylogenetic relationships ofAustralaSian wolf spider genera (Araneae:Lycosidae). JOllrnal or Arachnology 30: 227-237_

    Zabka, M. (1990), Salticidae (Arachnida: Araneae) ofOriental, Australian and Pacific regions, IV, GenusOcrisiona Simon, 1901. Records of the AllstralianMllsellm 42: 27-43.

    Zabka, M. (1991). Salticidae (Arachnida: Araneae) ofOriental, Australian and Pacific regions, V, GenusHoloplatys Simon, 1885, Records of the AllstralianMllsellm 43: 171--240.

    Zabka, M, (1992a), Salticidae (Arachnida: Araneae) ofOriental, Australian and Pacific regions, VIII , A newgenus from Australia. Records of' the ~Vestern AlIstralianMllsellm 15: 673-684,

    Zabka, M. (1992b). Salticidae (Arachnida: Araneae) ofOriental, Australian and Pacific regions, VII.Paraplatoides and Grl1yenlllla - new genera fromAustralia and New Caledonia. Records or theAlIstralian MllsCtlm 44: 165--183.

    Zabka, M. (2000). Salticidae (Arachnida: Araneae) of theOriental, Australian and Pacific regions, XIII: thegenus Sandl1lodes Keyserling, Il/uertebrate Taxol/omy 14:695-~704_

    Zabka, M_ (2001), Salticidae (Arachnida: Araneae) ofOriental, Australian and Pacific regIOns, XIV, Thegenus Adoxotoma Simon. Records or the Westerl/Allstral/an Mllsellm 20: 323-332,

    Zheng, H,B" Wyrwoll, K,H., Li, CM, and Powell, CM.(1998), Onset of aridity m southern Western Australta

    a prelIminary palaeomagnetic appraIsal. Global and18 175-187,

    Platnick, N,I. and For~ter, R,R, (1989), A reVISIon of thetemperate South American and AustralaSIan spidersof the fam Anapldae (Araneae,1311111'111/ of tht' American A'luseum of Natllml1-139

    Natllrt' COII'cri'atlOnand Dala A 109~126. CSIRO I)IvI~Ion ofWIIdltfe and . Canberra.

    McMahon, [1', llutchmson, M.F., Nix, FLA. and Ord,K.D. (1995). ANUCii;\:l users gllide, version 1. Centrefor Re~ource and EnvIrCmmental Studie~, Au~trahanNatIonal Canberra.

    LII,'ld,:ca/'e health in Australia. A rapidassesslllent the relallvt' condition of Allstralln'sbioregions and subregions. Environment Australia andNatIonal Land and Water Resources Audit, Canberra.

    Morgan, K.H. and Peers, R. (1973). Espeml/ce-Mondmin[sllll/d, Wt'stern AlIstralill, I 250 000 geological scries

    of Western Australia, Perth.

    tVlonng, [.B and Stewart, K.W. (1994) IlabItatpartltlmllng the wolf spider (Araneae, Lyco~Idae)guilds m streamsIde and npanan vegetatIon zone~ ofthe Conelo~ River, Colorado. Journlll of Amclwology 22:205-217.

    Mulcahy, M.[, and Hingston, F.[. (1961). Thedevelopment and distribution of the soil~ of the York-Quairading area, We~tern Au~tralia, in relation tolandscape evolution. Soil Publication 17. CSIRO,Canberra, Au~tralia.

    [S and Rtd. (1Weslerll llslmlill I 2,S()(),()()()

    of Western Au~traIIa, Perth.

    tv1yers, N, tvltttermeIer, RA., MittermeIer, C.G., Fonseca,, G.A.B. da and Kent, J. (2000). BIodiversIty hots pots

    for conservation priorities. Nature 403: 853-858.

    Newsome, A.E. and Catling, P.C (1979), Habitatpreference of mammals mhabIting heathlands ofwarm temperate coastal, Inontane and alpine region~of southeastern Australia, [n Specht, R.L (cd.),ECOSI!stellls of the world, vof. 9A. Heathlands al/d relatedshmiJlal/ds of the world: 301-316, Elsevier, Amsterdam.

    Oksanen, L. (2001). of experiments in ISpseudorephcation a pseudoissue! Oikos 94: 27-38,

    Platnick, N ,I. (2000), A relimltation and revi~Ion of theAustralasian ground family LamponIdae(Araneae: Gnaphosoidea). Bulletlll of the AlIlerlcanMllseUIll of Natllml History 254: 1-330,

    Platnick, N.I. (2002). A revision of the Australasianground spIders of the families Ammoxenidae,CithaeronIdae, Calheniellidae and Trochanteriidae(Araneae: Cnaphosoidea), Bulletin of the AmericanMllseum of Natuml 271: 1--243,

    Platnick, N,I. (20(Jl), The World Spider Catalog, Version4,5, American Museum of Naturall'listorv, New Yorkat rch,amnh org/(Cnl:ornolo;gv/sf)IClers/

    .htrn!.

    Electronic appendices are on CD inside the back cover

    Supplement67.pdfCOVERSTATE SALINITY STRATEGY BIOLOGICAL SURVEY OF THE WESTERN AUSTRALIAN WHEATBELT BACKGROUNDAQUATIC INVERTEBRATE ASSEMBLAGES OF WETLANDS AND RIVERS IN THE WHEATBELT REGION OF WESTERN AUSTRALIAWETLAND FLORA AND VEGETATION OF THE WESTERN AUSTRALIAN WHEATBELTSOILS SAMPLED DURING A BIOLOGICAL SURVEY OF THE WESTERN AUSTRALIAN WHEATBELTBIOGEOGRAPHIC PATTERNS IN SMALL GROUND-DWELLING VERTEBRATES OF THE WESTERN AUSTRALIAN WHEATBELTTERRESTRIAL FLORA AND VEGETATION OF THE WESTERN AUSTRALIAN WHEATBELTPATTERNS IN THE COMPOSITION OF THE JUMPING SPIDER (ARACHNIDA ARANEAE SALTICIDAE) ASSEMBLAGE FROM THE WHEATBELT REGION, WESTBIOGEOGRAPHICAL PATTERNS OF ZODARIID SPIDERS (ARANEAE ZODARIIDAE) IN THE WHEATBELT REGION, WESTERN AUSTRALIA.FAUNAS OF UNFLOODED SALINE WETLAND FLOORS OF THE WESTERN AUSTRALIAN WHEATBELTPATTERNS IN THE COMPOSITION OF GROUND-DWELLING ARANEOMORPH SPIDER COMMUNITIES IN THE WESTERN AUSTRALIAN WHEATBELTPATTERNS IN THE BIODIVERSITY OF TERRESTRIAL ENVIRONMENTS IN THE WESTERN AUTRALIAN WHEATBELTBIODIVERSITY PATTERNS AND THEIR CONSERVATION IN WETLANDS OF THE WESTERN AUSTRALIAN WHEATBELTTOWARDS IDENTIFICATION OF AN EFFICIENT SET OF NATURAL DIVERSITY RECOVERY CATCHMENTS IN THE WESTERN AUSTRALIAN WHEATBELT