DOI: 10.18195/issn.0313-122x.67.2004.257-291

Patterns in the composition of ground-dwelling araneomorph spider communities in the Western Australian wheatbelt

2 2 M. S. Harvei, J. M. Waldock', N. A. Guthrie , B. J. Durranf and N. L. McKenzie 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 Very few araneomorph spider families are South-western Australia represents a region that currently well known in southern Western is internationally recognized for its exceptional Australia. Taxonomic revisions have been diversity of terrestrial life, according it the status of published for only a few taxa, including: one of the world's top-20 biod iversity hotspots Ammoxenidae (Platnick, 2(02), Anapidae (Platnick (Myers et n/., 20(0). This rilnking was largely based and Forster, 1989), Filistatidae (Gray, 1994), upon the remarkable radiations of the vasculilr Gallieniellidae (Platnick, 2(02), Hersiliidae (Baehr flora, but it has become increasingly obvious that and Baehr, 1987, 1989, 1992, ]993, 1995, 1998), many components of the terrestrial invertebrate Lamponidae (Platnick, 20(0), Nicodamidae fauna are very species-rich and that high (Harvey, 1995), Orsolobidae (Forster and Platnick, proportions of the fauna are endemic to the region 1985), some Salticidae (e.g. Zabka, 1990,199], 1992a, (Hopper et nl., 19(6). Some terrestrial and aquatic 1992b, 2000; Zabka, 2(01), some Sparassidae invertebrates ilre also postulated as possessing high (Davies, 1994; Hirst, 1989, 199Ia,b, 1992), numbers of short-range endemic taxa (Hilrvey, Trochanteriidae (Platnick, 20(2) and most 20(2). We confined our study to araneomorph Zodariidae (BachI' and Jocquc', 2000, 2001; Jocquc' spiders which are easy to collect in pit-fall traps and Baehr, 2001; Jocquc' and Baehr, 1 . BachI' and and for which we have sufficient expertise to Churchill, BachI', in press) undertake species-levl'l identifications. Over the past 15 yeilrs, several broad-scale biotic Although sporadic descriptions of Western survevs have been conducll'd in vVestern Australia, /\ustralian spiders were milde bv 19 th century and spiders were one of the grou ps tilrgcted for European taxonomists on specimens collected from detailed analvsis to discern how the spider faun,) King Ceorge Sound and the Swan River Colonv, the WilS distributed across the landscilpe. The first \\'ilS first comprehensive publication was iln examination of the hunil of the Kimberlev Simon (I 19(9) who published on the raintorests in which spiders Wert' eXilmined in collections of a C;ermiln F:xpedition led W. detail bv Milin (1991). A rich ilssemblage of both Michaclsen and I~. Ila to south-western cursoriill and web-building types was IT'\·ealed. Australia in 1905. A great varietv of were Perhaps the most comprehensive studv was in the named, but unfortunatl'ly few of the names are southern Cilrnarvon Basin where a pitfall trapping currently in use due to the poor descriptions and program revealed a totil1 fauna of ilt le,lSt SOU paucitv of illustriltions. (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, and METHODS spatial patterns in the composition of the spider fauna were strongly correlated with climatic and Study area soil attributes. The wheatbelt study area has an area of 205 000 The dataset we investigated in this paper was km 2 and comprises all or part of five collated as part of a biodiversity survey of the biogeographical regions (Figure 1). The inland Western Australian wheatbelt (Figure 1), an inland boundary is a little to the west of the 300 mm area of some 205000 km2 ranging from Geraldton isohyet. The western boundary is east of the 600 to Esperance, to discern patterns of endemism in a mm isohyet. Along the southern coast, the southern highly fragmented and disturbed landscape. We boundary approximates the 550 mm isohyet but provide the first account of the Western Australian avoided near-coastal environments. Its semi-arid to wheatbelt's fauna of grolmd-dwelling araneomorph sub-humid climate is influenced by temperate spiders, and explore patterns in community weather systems (mainly winter rainfall, see Bureau composition in terms of measurable attributes of the of Meteorology, 2001), and its surface hydrology is area's physical environment, including salinity. characterised by low gradients, high potential salt

11 18 18 KALBARRI

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

o 100 Kilometres 118

Figure 1 Wheatbelt study area, showing the 24 survey areas and the positions of the 12 to 13 individual quadrats in each survey area. Precise quadrat co-ordinates are provided in Appendix 1. The coloured areas indicate bioregional boundaries (Environment Australia, 2000; modified from Beard, 1980). The fine lines internal to the bioregions are sub-regional boundaries defined from IBRA version 5 (Environment Australia, 2000; Morgan, 2001). Araneomorph spiders 259 loads and high variability in flows (Commander et these valley floors. Although the north-western aI., 2002; George and Coleman, 2002; Hatton and (Geraldton Sandplains) and south-eastern Ruprecht, 2002; Schoknecht, 2002). Overall, 74% of (Esperance Plains) parts of the study area are on the study area has been cleared for agriculture; its different geological basements (e.g. Morgan and remaining bushlands are isolated woodland and Peers, 1973), their geomorphic units could be shrubland mosaics embedded in fields of wheat, classified into the same landform catena according lupin and canola. Extensive areas are now affected to soil profile attributes. by dry land salinity (George et al., 1995). In phytogeographical terms the study area Field sampling strategy comprises the semi-arid part of the South-Western Sampling strategy is discussed in McKenzie et al. Botanical Province (Beard, 1980). Detailed (2004). The study area was divided into 24 survey descriptions of the vegetation in the study area are areas of similar size so that sampling was evenly provided by Beard (1980, 1981, 1990). The study dispersed across the study area's areal extent area's substrates are detailed by Chin (1986), (Figure 1). Ten to 11 quadrats each of one hectare Mulcahy and Hingston (1961) and Myers and were positioned to sample the geomorphic profile Hocking (1998). Most of it is an undulating plateau of each of 24 survey areas (Figure 1). In each survey of Tertiary duricrust overlying Archaean granitic area, aquadrat was positioned in atypical example and metamorphic rock strata of the Yilgarn Craton. (in terms of vegetation and soil) of each of the main The plateau is variously eroded into elluvial and geomorphological units (Baxter and Lipple, 1985; colluvial spillway sand deposits supporting a Chin, 1986; Morgan and Peers, 1973; Mulcahy and variety of shrub lands, and dissected to expose Hingston, 1961). An additional one or two quadrats valley-slope and -floor units characterised by were positioned to sample examples of these units duplex soils derived from the Archaean basement that were affected by groundwater salinity. The strata and supporting woodlands. Except in the quadrats chosen were the least disturbed examples south-west, chains of salt-lakes also occur along we could find. Even the sites selected to represent

,.-- ~I~ ~ ___,

Spillway deposits

Qe Quailing Erosional K Kauring Qd Quailing Depositional M Monkopen Bg Belmunging B Balkuling Mg Malebelling Y York A Avon Mo Mortlock Sf Saltflat Sw Swamp (fresh) Figure 2 Wheatbelt land forms, modified from Mulcahy and Hingston (1961). The landscape's plateau profile comprises the duricrusted Tertiary laterite plateau and its derived spillway sands, while the dissection profile comprises finer textured soils derived from bedrock and pallid-zone clays beneath the duricrust. 260 M. S. Harvey, J. M. Waldock, N. A. Guthrie, B. J. Durrant, N. L. McKenzie secondarily salt-affected sites were positioned to relevant 1:250 000 maps of surface lithology (e.g. minimise proximity to wheatfields and avoid Morgan and Peers, 1973), and arbitrarily assigned evidence of other disturbances such as rubbish to our 12-class landform catena according to their disposal, gravel extraction and previous clearing. soil profile (texture and horizon sequence), position Locations of the 304 quadrats are provided in in the relevant landscape's profile and soil origin. Appendix 1. Thus, a deep, low-level sandsheet wouid be A consistent landform model was used to assigned to the same number as the lowest of the position the individual quadrats (Figure 2), so an spillway sand units on the Craton (Monkopen = 9, equivalent cross-section of the landscape profile's in Figure 2). This approximation was considered to main components was sampled in each survey area. be acceptable because data taken from the quadrats This nested, stratified sampling allowed an themselves were used in subsequent analyses. analytical option in which each survey area was Eighteen climatic attributes were derived for each treated as a single landscape-scale releve; by quadrat using ANUCLlM (McMahon et aI., 1995). pooling the lists of species from a survey area's 12 These comprised annual and seasonal average and or 13 quadrats, we might suppress the influence of range values for temperature and precipitation local substrate on the classification structure, (Appendix 2). Soil attributes were also recorded thereby exposing broad-scale patterns for analysis. from each quadrat (Appendix 3), including The spiders on each quadrat were sampled using nitrogen, phosphorus, potassium, pH, electrical five 2-litre, glycol pits left open for one calender conductivity, organic carbon, clay-silt-sand year (1825 pit trap nights per quadrat). Pits were 25 percentages and magnesium. Eight landform and cm deep, 12 cm in diameter, contained 0.4 litres of five vegetation attributes were also generated glycol-formalin (320 ml ethylene glycol, 64 ml tap (Appendix 1), including latitude, longitude, water, 16 ml formaldehyde) and were set flush with elevation, landform unit, soil drainage category, the ground surface. Each pit was protected by a slope, salinity risk, salinity class, litter/log cover, square plate of wood (0.15 X 0.15 m) suspended 5 tree cover, shrub cover, herb cover and habitat cm above the pit's mouth. The five pits on each complexity (modified from Newsome and Catling, quadrat were positioned in different microhabitats 1979). Latitude and longitude values were - bare ground, leaf litter patches under trees, beside determined using a hand-held GPS accurate to ±30 rotting logs, under low shrubs etc. They were m. Explanation of the salinity attributes is provided always at least five metres apart. in McKenzie et al. (2004) and in Appendix 1. Quadrats in the YO, QU, KL, MN, NR, WK, KN Because landform had already been used to and HY survey areas were sampled from October position the quadrats in each survey area, it could 1997 to September 1998. NO, ML, MO, WU, JB, WH not also be used as an attribute in the compositional and BE were sampled from September 1998 to analysis. October 1999, and DA, DU, PI, LK, UN, ST, GP and ES from October 1999 to October 2000. Every four Analytical strategy to six months, specimens were removed from the The analytical approach taken in this paper was pits and fluids replenished. After washing, sorting based on the assumption that spatial distribution and identification, the specimens were lodged in reflects an underlying correlation with the collection of the Western Australian Museum. environmental factors (Austin, 1991; Clarke, 1993). It is an exploratory design, and interpretation is Physical attributes at quadrats based on deductive rather than inductive logic Landform units in survey areas on the Yilgarn (Oksanen, 2001). No experimental design has been Craton were numbered from 1 to 12 according to implemented to assess a null hypothesis (Austin their position in the landscape profile (see Figure 2, and McKenzie, 1988), so alternative hypotheses are McKenzie et al. 2004). Units belonging to the not excluded. Inferential statistics are used to test dissection profile were numbered from 1 to 7, while patterns observed in the empirical data. spillway sand and duricrust units associated with The relative responses of wheatbelt araneomorph the old plateau profile were numbered from 9 to 12. species to our pitfall traps are unknown as are the Unit Bg in which spillway sand mantles a clay year-to-year dynamics of population numbers in belonging to the dissected valley profile was these communities. We also recognise other assigned to number 8. Units at the bottom of the limitations in sampling techniques which were dissection profile included fresh water swamps aggravated by staff and time limitations. Because (unit 1) and saline flats (unit 2). The highest unit in these issues precluded reliable estimates of species the landscape was duricrust pavement of the old abundance, we chose to restrict our analyses to data Tertiary plateau (unit 12). Quadrats in survey areas on the presence-absence of taxa at individual on geological basements other than the Yilgarn quadrats (Austin, 1984; McKenzie et al., 1991). Thus, Craton (those in the Esperance Plains and Geraldton the input data utilised for the compositional Sandplains bioregions) were positioned using the analyses were survey area-x-genus, survey area-x- Araneomorph spiders 261 species and quadrat-x-species matrices. Species in the survey by 39 families, and a total of 24 640 recorded at only a single quadrat (singletons) were specimens were identified to species level. The excluded from compositional analysis because they numbers of specimens identified from each quadrat convey little information on patterns. were not particularly high despite large sampling To assess whether records of a species were more effort (average 81.1, s.d. 49.6. range 2 - 316), and geographically localised than would be expected by there are substantial differences between quadrats chance, the average distance between the quadrats that may indicate differences in sampling at which a species was recorded was calculated and effectiveness and lead to bias in comparisons of compared with 1000 trials using an equivalent quadrat richness. Some of the poorest quadrats number of randomly arrayed records (quadrat were those on salt-flats or in low-lying salt-affected intersections). The probability (P) of achieving the areas. An additional six months of sampling at a observed average distance was then assessed from sub-set of the salt-flat quadrats revealed no the distribution of the 1000 random Iv derived additional araneomorph species (Durrant and values. Cuthrie, 2004), so we proceeded with analyses of '1'0 gain a clear separation between species of non- richness on the basis that our samples adequately saline environments and those tolerant or adapted reflect trends. 111is sampling issue is investigated to saline conditions, the 51 quadrats affected by further under the heading Richness, below. secondary salinity were excluded from the quadrat- The representatives of five families were not x-species matrix (see McKenzie et aI, 2003). This identified to species-level due to time and resource separation provided a basis for interpreting the constraints, as well as uncertainty regarding the species composition of assemblages on the saline unresolved taxonomic boundaries at the species quadrats in the absence of extrinsic data on the level. These five families (Ctenidae, Cnaphosidae, habitat preferences of virtually any of the Miturgidae, Sparassidae and Zoridae) are not listed araneomorphs in the samples. Thirteen additional below and were excluded from the analyses. quadrats were eliminated from this analysis because Observations on species geographical occurrence they were flooded for long periods during the that are included in the annotated list below are trapping program (10 in ephemeral fresh water derived from Table 1. swamps, as well as PI04, UN13 and WK02). This left 240 quadrats in the quadrat-x-species matrix, Agelenidae but did not distort the stratification (see Table 1 in Two species assigned to the Agelenidae were McKenzie et ai, 2003). recorded in the survey. The first, Cenus 1 sp. 1, was Cluster analysis (from PATN, Belbin, 1995) was found at the Kt survey area, whilst the second, used to expose patterns of species composition in Cenus 1 sp. 2, was found at UN and WK. the data matrices. The Czekanowski measure (Czekanowski, 1932) was used to compare the quadrats according to their species similarities, Amaurobiidae because it is known to provide a robust measure of The family Amaurobiidae was represented in the ecological distance (Faith et aI, 1987). A modified survey by a diverse assemblage of 27 species. Four version of the unweighted pair group arithmetic species were found at more than 20 quadrats averaging (UPCMA - Belbin, 1995; Sneath and (Cenus 2 sp. 01, Cenus 3 sp. 01, Cenus 3 sp. 02 and Soka!, 1973) hierarchal clustering strategy was used, Cenus 3 sp. 16) and seven species were found at with the clustering parameter (Beta) set to -0.1. The only a single quadrat. The amaurobiid fauna of partition structure of the resulting dendrogram was Western Australia is poorly known, and the used as a summary of compositional patterns in systematic position of many of the species here araneomorph genera/species across the study area. placed in this family may reside in other families. Environmental attributes that conformed to the partition structures were assessed for statistical Amphinectidae significance using Kruskall-Wallis one way analysis The survey revealed 11 species attributed to the of variance by ranks (in the computer package Amphinectidae, most of which were found in just a STAT'ISTICA, Statsoft, 2001). The negative few quadrats. They were predominately recorded exponential smoothing function in STATISTICA from southern study sites. was used to interpolate climatic surfaces across the study area thereby illustrating geographic patterns Anapidae in climatic attributes that were found to be 'rhe anapid fauna of southern Western Australia significant. is represented by the sole genus Chasmocephalon, a genus that is also found in eastern Australia along with a variety of other genera (Platnick and Forster, RESULTS 1989). All anapids are tiny spiders less than 2-3 mm The infraorder Araneomorphae was represented in length. Three species of Chasmocephalon were N List of araneomorph taxa identified during the survey; the families Ctenidae, Gnaphosidae, Miturgidae, Sparassidae and Zoridae were omitted from the analysis due a- Table 1 N 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 quadrats from which each species was recorded.

Family Genus, species Study sites (arranged approximately north-west to south-east) No. of quadrats 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 ')'"7 Amaurobiidae Genus 3 sp. 01 2 3 1 3 2 3 1 4 6 1 1 d 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 CFJ 1 14 Amaurobiidae Genus 3 sp. 06 1 3 1 1 1 6 ::::: Amaurobiidae Genus 3 sp. 07 1 1 2 ...\"01 Amaurobiidae Genus 3 sp. 08 1 1 ro< 2 ~ Amaurobiidae Genus 3 sp. 09 2 '- 4 Amaurobiidae Genus 3 sp. 10 1 1 2 ~ Amaurobiidae Genus 3 sp. 11 1 2 1 4 1 4 3 16 13 ~ Amaurobiidae Genus 3 sp. 12 2 2 5 4 \"01 Amaurobiidae Genus 3 sp. 13 1 I 0: 0 Amaurobiidae Genus 3 sp. 14 1 1 ,., 2 1 1 3 1 1 1 24 "F Amaurobiidae Genus 3 sp. 16 2 4 4 4 ~ Amaurobiidae Genus 4 sp. 01 1 1 Amaurobiidae Genus 4 sp. 02 1 1 ?> 13 Cl Amaurobiidae Genus 4 sp. 03 1 1 1 2 1 1 2 1 3 r= Amaurobiidae Genus 4 sp. 04 1 I ;.... Amaurobiidae Genus 5 sp. 01 2 1 1 1 1 6 ,it' 2 Amaurobiidae Genus 7 sp. 01 1 1 ?:I Amaurobiidae Genus 8 sp. 01 1 1 ":""' Genus 9 sp. 01 2 1 2 1 2 3 1 1 13 0 Amaurobiidae r= Amphinectidae sp.Ol 1 1 ...... \"01 Amphinectidae sp.02 1 1 ::s 1 1 2 ~ Amphinectidae sp.03 ~ Amphinectidae sp.04 1 1 1 2 3 !'"' Amphinectidae sp.05 ~ Amphinectidae sp.06 1 1 2 ,., 6 ~ Amphinectidae sp.07 1 1 2 2 ro 7 ::s Amphinectidae sp.08 1 1 3 1 1 N Amphinectidae sp. 10 1 1 1 2 2 4 1 1 13 it· Amphinectidae sp.ll 1 1 Amphinectidae sp. 12 3 ] 4 Anapidae Chasmocephalon sp. 01 1 1 5 3 10 Anapidae Chasmocephalon sp. 02 2 1 3 6 12 sp.03 1 2 1 4 Araneidae sp.03 1 ] Araneidae sp.04 1 1 ] J 4 Araneidae sp. 05 1 1 Araneidae sp.06 1 I Araneidae sp. 07 J 1 2 Araneidae sp.08 1 I Araneidae sp.1O 1 1 2 Araneidae sp. 11 J 1 2 Araneidae sp. 12 ] 1 Araneidae sp.13 1 1 Araneidae sp.14 1 1 Araneidae sp. 16 I 1 Araneidae Argiope protensa 1 3 1 1 1 1 1 1 1 11 Araneidae Argiope trifasciatl1 1 1 Araneidae A IIstracantha minax 1 1 1 1 1 2 1 1 9 Araneidae Oolophones sp. 01 1 1 2 Araneidae Dolophones sp. 02 1 2 1 4 Araneidae Dolophones sp. 03 1 1 2 Araneidae Eriophora IJiapicata 1 1 2 Araneidae Eriophora transmarina 1 2 1 1 1 J 7 Araneidae Eriophora sp. 01 1 1 2 Araneidae Eriophora sp. 02 1 J Araneidae Genus X sp. 1 2 1 4 Araneidae Genus Y sp. 1 2 3 Araneidae Genus Z sp. 1 1 J 3 l'v1atilda sp. 01 1 1 1 1 1 6 2 3 1 1 1 1 20 Cyatholipidae Matilda sp. 02 1 1 1 1 1 2 1 1 9 Cycloctenidae Genus 1 sp. 01 1 1 Cycloctenidae Genus 1 sp. 02 1 1 Deinopidae Deinopis sp. 1 1 1 1 1 1 1 1 2 10 Desidae sp.Ol 1 1 Desidae sp.02 Desidae sp.03 1 J Desidae sp.04 1 J 1 3 Desidae sp.05 1 1 2 Desidae sp.06 1 1 2 Desidae sp.07 1 1 Desidae sp 08 1 1 Desidae sp.1O ] I Desidae sp. 11 1 1 2 N '"v.>

N Table 1 (cont.) Cl' Cl' Family Genus, species Study sites (arranged approximately north-west to south-east) No. of quadrats 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 1 Lamponidae Lamponina sp. 03 1 1 2 1 1 6 Lamponidae Lamponusa gleneagle 3 1 4 Lamponidae Longepi woodman 1 1 7 1 4 1 5 1 1 1 3 26 Lamponidae Notsodipus domain 1 1 2 1 5 Lamponidae Notsodipus muckera 1 1 1 3 2 2 1 1 1 1 14 Lamponidae Notsodipus quobba 2 2 2 6 Lamponidae Notsodipus sp. 01 1 1 Lamponidae Prionosternum porongorup 2 2 4 Lamponidae Prionosternum scutatum 6 2 1 4 2 2 1 3 21 Lamponidae Pseudolampona boree 1 1 2 1 1 1 1 1 1 1 2 2 2 17 ~

Lamponidae Queenvic mackay 3 1 4 rJJ 2 2 4 1 6 2 4 3 5 2 1 2 4 2 1 3 2 1 2 49 Linyphiidae sp.Ol :I: Linyphiidae sp.02 4 1 1 1 1 3 2 3 3 4 4 1 1 1 3 1 34 III ::l Linyphiidae sp.03 1 1 5 2 1 2 1 3 4 3 4 4 3 3 3 1 1 1 43 nl ~ Linyphiidae sp.04 4 1 1 3 8 7 6 2 2 1 1 1 3 4 6 2 1 2 3 2 1 61 1 3 5 .. Linyphiidae sp.05 1 ~ Linyphiidae sp.06 1 3 2 1 1 1 1 1 1 1 13 36 ~ Linyphiidae sp.07 2 2 4 4 6 2 2 1 1 3 3 2 2 2 III Linyphiidae sp.08 2 2 2 1 1 1 1 1 1 1 13 c::0 Linyphiidae sp.09 4 2 1 1 1 3 4 2 3 2 23 ,., ~ 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 ~ 2 1 1 1 11 Cl Linyphiidae sp.13 1 1 4 I: Linyphiidae sp.14 3 2 3 4 1 4 4 1 4 1 1 1 4 4 1 38 ::r.... Linyphiidae 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 Cl I: Linyphiidae sp.19 2 1 1 4 .... III Linyphiidae sp.20 2 3 1 1 3 1 11 ::l , Linyphiidae sp.21 1 1 .... Z Linyphiidae sp.22 1 2 3 1 7 Linyphiidae sp.23 2 2 r' ~ Linyphiidae sp.24 1 1 ,., 1 ~ Linyphiidae sp.25 1 nl 3 ::l Linyphiidae sp.26 2 1 N Linyphiidae sp.27 1 1 ;;;. sp.28 I 1 sp. 29 ] sp. 30 1 sp.31 I sp.32 1 I sp.33 I sp. 34 3 2 3 2 1 sp.35 sp.36 sp.37 sp. 39 2 sp.40 2 2 I 1 2 3 sp.41 1 1 sp.42 sp.43 sp.44 j 1 2 sp.45 ] 4 sp.46 sp.47 I 1 sp.48 I I 2 sp.49 sp.50 I ] sp.51 1 I 2 sp. 52 I I sp. 53 sp.54 1 1 sp.55 I I I 3 sp. 56 I 1 I 2 2 2 9 57 D sp. Ol 2 Oslearius 1 1 3 2 4 8 3 12 11 11 10 Liocranidae sp. Ol 1 Liocranidae sp. 02 Liocranidae sp.04 Liocranidae sp.05 1 1 I 2 Liocranidae sp.06 I 1 1 1 4 Liocranidae sp.07 I 1 1 2 1 2 8 Liocranidae sp.09 1 1 1 3 Liocranidae 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 9 Arloria sp. 0] 4 2 1 4 1 2 4 10 8 1 1 1 5 44 Arloria sp. 02 1 1 4 1 4 1 5 5 2 24 Arloria sp. 03 2 1 7 2 1 6 3 2 1 6 6 7 2 5 6 7 1 4 3 6 5 2 5 Arloria 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 ;.. ...1:J sp.22 3 4 3 I 1 I 13 ::l rt> sp.23 1 I 2 0 Lycosidae I 3 Lt/cosa sp. 24 I 0 sp. 25 1 I '1;l... ::r Lycosidae Lycosa sp. 26 I I CJ; sp. 27 1 2 3 '1;l 0.: sp. 28 1 1 rt>... sp.29 1 2 2 I I 7 CJ; sp. 30 1 1 Lycosidae Lycosa sp. 31 1 1 Lycosidae Lycosa sp. 32 1 1 1 3 Lycosidae Lycosa storri 1 2 1 1 1 I 1 8 Venatrix arenaris 2 1 2 2 4 1 12 Lycosidae Venatrix I'II11astra 7 1 3 4 2 1 4 22 Jae Venonia sp. 01 1 1 2 4 Micropholcommatidae Micr0l'holcomma? sp. 01 2 1 1 3 1 3 3 1 I 4 2 2 24 Micropholcommatidae Micr0l'holcomma? sp. 02 I 1 1 4 3 3 1 2 4 2 2 1 25 ~ Micropholcommatidae Microl'holcomllla? sp. 03 2 I 2 1 1 I Micropholcommatidae Microl'holcomma? sp. 04 1 3 1 1 2 2 I 3 1 I 1 17 Micropholcommatidae Microl'holcolllma? sp. 06 I 2 II 5 2 1 13 Micropholcommatidae Microl'holcomma? sp. 08 2 1 1 1 2 1 1 3 I 2 1 I 1 18 Micropholcommatidae Micr0l'holcolllllla? sp. 10 1 1 1 1 4 Micropholcommatidae Micropholcomllla? sp. 11 2 1 1 3 7 Micropholcommaticlae Microl'holcolllma? sp. 12 2 1 3 Micropholcommatidae Micropholcolllma? sp. 13 1 1 2 Micropholcommatidae Textricella sp. 01 3 1 3 1 1 6 2 3 1 7 3 2 5 8 3 7 5 3 4 2 2 72 Micropholcommatidae Textricella sp. 02 I I Mysmenidae Genus 1 sp. 01 1 I Mysmenidae Genus 2 sp. 01 1 Nicodamidae AlIlbicodanllls marae 1 1 1 1 4 Nicodamidae Nicodamlls mainae 2 6 1 2 4 1 1 1 I 2 2 3 26 Orsolobidae sp.01 1 Oecobiidae Oecobills naVllS 1 1 2 Oonopidae Gamasomorpha sp. 01 Oonopidae Gamasomorpha sp. 13 Oonopidae Gamasomorpha sp. 02 1 1 2 1 S Oonopidae Gamasomorpha sp. 03 1 2 1 1 5 Oonopidae Gamasomorpha sp. 04 1 Gamasomorpha sp. 05 Oonopidae Gamasomorpha sp. 06 1 1 1 1 1 S Oonopidae Gamasomorl'ha sp. 07 3 1 1 4 2 3 1 1 1 2 6 5 2 2 34 Oonopidae 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 58 Oonopidae 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 N C!' \0

Philodromidae Tibcllus sp. 03 1 I ;;....., ::.. Philodromidae Tibcllus sp. 04 1 I ;:l Salticidae Adoxotoma chillOPOSOIl I I 3 I 6 '"0 Salticidae Adoxotoma Ilisroo/ivacca 1 I :3 ...,0 Salticidae Adoxotoma sp. 01 4 2 6 ":l ::r Salticidae Adoxotoma sp. 02 1 1 CJ; Salticidae Biallor maculatlls 1 I 2 ":l 2 Cl: Salticidae Biallor sp. Ol II .... 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 22 Salticidae Clyllotis sp. 02 2 1 1 1 1 1 2 2 1 2 14 Salticidae Cll/lwtis sp. 03 1 I 1 3 Salticidae ClYllotis sp. 04 Salticidae ClYllolis sp. 05 Salticidae Clyllotis sp. 06 1 1 Salticidae Cllfllotis sp. 07 1 1 2 4 Salticidae Clyllotis sp. 08 1 1 1 3 Salticidae Clyllotis sp. 09 1 2 2 1 6 Salticidae Clyrwtis sp. 10 1 1 2 Salticidae Clyllotis viduus 1 1 1 I 2 6 Salticidae Cytaea sp. 01 1 1 2 Salticidae Damoctas sp. 01 1 1 2 Salticidae 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 84 Salticidae Genus 1 sp. 02 1 1 2 1 3 6 2 2 I 2 1 1 2 I 26 Salticidae Genus 1 sp. 03 1 1 1 1 3 1 1 1 1 1 1 2 2 17 Salticidae Genus 1 sp. 04 1 1 1 3 SaIticidae Genus 1 sp. 05 1 1 1 3 Salticidae Genus 1 sp. 06 1 1 Salticidae Genus 1 sp. 07 1 1 1 3 Salticidae Genus 10 sp. 01 1 1 2 Salticidae Genus 12 sp. 01 1 1 2 Salticidae Genus 12 sp. 02 1 3 2 1 7 Salticidae Genus 12 sp. 03 Salticidae Genus 12 sp. 04 1 1 1 1 1 1 6 Salticidae Genus 13 sp. Ol 1 2 2 1 2 1 1 1 11 Salticidae Genus 14 sp. Ol 1 1 1 1 1 1 6 Salticidae Genus 15 sp. 01 1 1 Salticidae Genus 16 sp. Ol 1 1 2 Salticidae Genus 17 sp. 01 1 1 4 3 1 10 Salticidae Genus 18 sp. Ol 2 1 1 2 2 8 Salticidae Genus 19 sp. 01 1 1 Salticidae 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 69 Salticidae Genus 20 sp. Ol 1 1 2 Salticidae Genus 3 sp. 01 2 1 1 1 2 2 9 Salticidae Genus 3 sp. 02 2 2 1 1 1 2 9 N ....'-l Table 1 (cont.) N 'I N Family Genus, species Study sites (arranged approximately north-west to south-east) No. of quadrats 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 3sp. 03 1 1 1 3 Salticidae Genus 3sp. 04 1 1 2 Salticidae Genus 3sp. 05 1 I Salticidae Genus 3sp. 06 1 1 Salticidae Genus 5sp. 01 1 2 2 4 1 1 2 1 1 1 16 Salticidae Genus 6sp. 01 1 1 1 3 Salticidae Genus 7sp. 01 2 2 Salticidae Genus 9sp. 01 1 1 2 1 1 1 1 2 1 1 12 Salticidae Grayenulla allstralensis 2 6 2 4 2 3 9 2 1 1 5 1 38 Salticidae Grayenulla nova 5 1 6 Salticidae Holoplatys chlldalllpensis 1 1 1 3 :::: Salticidae Holoplatys dejongi 1 1 'Jl Salticidae Holoplatys sp. 01 1 1 :: Salticidae Holoplatys sp. 02 1 1 2 tJ Salticidae Holoplatys sp. 03 ~ 1 1 re Salticidae Hypoblemllm sp. 01 1 1 2 2 1 7 :< Salticidae Hypoblemllm sp. 02 1 1 2 2 6 -- Salticidae Lycidas chnjsomelas 2 1 2 2 4 6 6 3 2 3 6 6 4 1 6 4 3 4 5 3 4 2 1 1 81 :::: Salticidae Lycidas michaelseni 4 1 3 1 1 2 4 2 1 19 ~ tJ Salticidae Lycidas sp. 01 1 4 3 1 5 8 6 9 9 6 5 7 11 9 6 5 8 1 8 8 9 9 5 2 145 0: Salticidae Lycidas sp. 02 2 3 1 6 2 1 1 2 1 1 2 1 1 5 1 5 3 3 5 1 1 48 ,.,0

Salticidae Lycidas sp. 03 ~ r 2 3 5 1 3 6 2 1 1 10 9 5 4 3 6 3 1 1 2 2 1 1 I~ Salticidae Lycidas sp. 04 2 3 1 3 2 2 3 1 6 4 2 4 1 1 1 1 37 Z Salticidae Lycidas sp. 05 1 8 6 1 2 9 2 3 8 8 8 2 4 62 ?- Salticidae Lycidas sp. 06 1 1 1 1 1 1 6 Cl Salticidae Lycidas sp. 07 r::: 1 1 1 1 2 1 7 ...=r Salticidae Lycidas sp. 08 1 2 1 1 2 7 ,;' Salticidae Lycidas sp. 09 1 1 ~ Salticidae Lycidas sp. 10 1 2 3 Salticidae -- Lycidas sp. 11 1 1 1 1 4 Cl Salticidae Lycidas sp. 12 1 1 2 3 1 1 1 1 2 1 14 ...r::: Salticidae Lycidas sp. 13 1 1 2 tJ Salticidae Lycidas sp. 14 1 1 2 ~ Salticidae Lycidas sp. 15 1 1 Z Salticidae Lycidas sp. 16 1 1 2 1 1 3 1 3 1 2 16 r' Salticidae Lycidas sp. 17 1 2 1 1 4 1 2 4 3 1 1 1 22 ::::,., Salticidae Lycidas sp. 18 1 1 3 2 "I / re~ Salticidae Lycidas sp. 19 1 2 1 1 1 2 8 ::l N Salticidae Lycidas sp. 20 1 1 ::t. Sillticidae Lycidas sp. 21 1 1 >- ""OJ Lycidas sp. 22 2 1 2 1 6 ::l SZllticidae ro Sillticidae Lycidas sp. 23 1 9 7 1 18 0 3 Salticidae Lycidas sp. 24 1 1 0 Sillticidae Lycidas sp. 25 1 1 '"Cl"" ::r SZllticidae Lycidas sp. 26 1 2 2 5 \J) 1 I '"Cl Salticidae Llfcidas sp. 27 0.: SZllticidae Nlaratlls mllllgaich I 1 2 ro Sillticidile A1aratlls 4 I 1 1 1 1 I I 1 I 1 14 ""U1 Sillticidae Maratlls sp. 01 Salticidae Maratlls sp. 02 Sillticidae Maratlls sp. 03 I SZllticidae 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. 02 SZllticidae 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. 01 SillticidZle Scrvaea sp. 01 ~ Salticidae Simaetha sp. 01 2 1 .) Salticidae Simaetha sp. 02 Salticidae Simaetha sp. 03 Sillticidae Simaethllla sp. 01 Salticidae Simaethll/a sp. 02 Salticidae Simaethllla sp. 03 Salticidae SOlldra sp. 01 2 1 4 1 10 2 3 8 1 1 33 Sillticidae 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 2 Genus 1 sp. 01 1 1 1 1 1 5 Segestriidae Genus 1 sp. 02 1 1 2 2 3 1 2 1 1 14 Genus 2 sp. 01 1 1 2 1 4 2 3 6 6 1 5 1 1 4 1 39 DclClla sp. 01 1 1 Genus 1 sp. (J1 1 1 Baiami volllcril'cs 3 1 3 2 9 4 2 1 25 1 1 3 22 N Comsoides sp. (J1 3 2 3 3 6 '-l W Table 1 (cont.) N ...~ Family Genus, species Study sites (arranged approximately north-west to south-east) No, of quadrats 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 10 Stiphidiidae Corasoides sp. 03 2 1 3 Stiphidiidae Corasoides? sp. 04 1 3 1 1 1 1 1 1 1 11 Stiphidiidae Corasoides? sp. 05 1 1 1 1 4 Stiphidiidae Forsterina sp. 01 1 1 1 3 Tetrablemmidae Genus? sp? 1 1 Tetragnathidae "metine" sp. 01 1 1 1 3 1 7 Tetragnathidae "metine" sp. 02 1 1 2 Theridiidae sp.01 1 1 Theridiidae sp.02 1 1 Theridiidae sp.03 1 I ~ Theridiidae sp.04 1 1 fJl Theridiidae sp.05 1 1 ..:z:: Theridiidae sp.06 1 1 < Theridiidae sp.07 1 I rtl Theridiidae sp.13 1 I ~ Theridiidae "Ctenopalpus" "hirsti" 2 2 - Theridiidae "Ctenopalpus" bicruciatus 1 1 1 1 4 1 9 ~ Theridiidae "Genus A" sp. 1 1 1 3 ..~ Theridiidae "Nico" sp. 01 1 4 2 1 4 1 2 3 1 1 20 c:: 0 Theridiidae Ariamnes sp. 1 1 2 " Theridiidae Enoplognatha sp. 4 2 1 2 1 2 2 4 1 1 2 1 1 2 1 1 28 r Theridiidae Episinus sp. 01 1 1 1 3 ~ Theridiidae Episinus? sp. 02 1 1 ?> Theridiidae Dipoena sp. 01 1 2 1 2 1 2 1 1 1 12 Cl t: Theridiidae Dipoena sp. 02 1 1 ...s:- Theridiidae EunJopis sp. 01 1 1 ~ Theridiidae Euryopis sp. 02 1 1 ~ Theridiidae Euryopis sp. 03 1 1 1 1 1 5 ':'"' Theridiidae Euryopis sp. 04 I 1 0 Theridiidae Euryopis sp. 05 1 1 ...t: Theridiidae .. ElIryopis sp. 06 1 2 1 1 1 2 2 1 1 1 1 1 15 ::l Theridiidae Euryopis sp. 07 2 2 3 4 2 2 1 1 17 ~ Theridiidae Euryopis sp. 08 2 2 2 1 1 1 1 2 1 3 1 17 Z Theridiidae ElIryopis sp. 09 2 5 2 8 4 5 9 6 2 5 1 3 3 2 5 4 1 3 1 1 3 4 79 r' Theridiidae Euryopis sp. 10 1 1 ~ Theridiidae ElIryopis sp. 11 1 1 ",.:: rtl Theridiidae Genus 1sp. 01 3 2 1 2 3 1 1 1 1 1 2 1 1 1 21 ::l N Theridiidae Genus 1sp. 02 1 1 4 6 ;;'

$ 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 ::l '"r'I> Theridiidae Genus 2 sp. 01 1 1 1 1 1 5 0 1 1 1 3 3 Theridiidae Genus 2 sp. 02 ...0 Theridiidae Genus 3 sp. 01 1 2 2 2 1 1 1 I 1 2 14 '\:l ::r' Theridiidae Genus 3 sp. 02 1 1 2 2 2 '\:l'" Theridiidae Genus 4 sp. 01 0.: 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. 04 Theridiidae Genus 4 sp. 05 1 I 2 Theridiidae Genus 4 sp. 06 1 I 2 Theridiidae Genus 5 sp. 01 1 1 1 3 Theridiidae Genus 5 sp. 02 2 1 3 Theridiidae Genus 5 sp. 03 2 J I 2 I I 8 Theridiidae Genus 6 sp. 01 1 1 I 3 I 1 I I 2 I 5 I I 20 Theridiidae Genus 6 sp. 02 J I 2 Theridiidae Genus 7 sp. 01 I I 2 Theridiidae Genus 7 sp. 02 2 2 Theridiidae Genus? sp. J 5 1 1 1 8 2 2 4 3 2 2 1 33 Theridiidae Gmogala sp. 01 1 1 2 Theridiidae Gmogala sp. 03 1 1 3 2 1 6 2 2 3 1 3 1 2 5 1 2 2 3 1 1 43 Theridiidae Gmogala sp. B 2 3 1 4 5 4 1 3 1 2 4 2 6 7 4 1 1 51 Theridiidae Gmogala sp. U 1 3 2 1 2 1 9 2 2 1 1 25 Theridiidae Hadrotarsinae Genus 1 sp. 01 1 2 2 1 1 1 8 Theridiidae Hadrotarsinae Genus 2 sp. 01 1 1 1 3 Theridiidae Hadrotarsinae Genus 3 sp. 01 1 1 2 Theridiidae Hadrotarsinae Genus 0 sp. 01 1 1 1 2 1 6 Theridiidae Hadrotarsinae Genus 0 sp. 02 1 I Theridiidae HadrotarslJs sp. 02 1 1 1 1 1 2 1 3 2 2 3 1 2 21 Theridiidae HadrotarslJs sp. 03 1 2 1 2 1 4 2 1 2 1 2 19 Theridiidae HadrotarslJs sp. C 1 1 1 1 1 1 1 1 2 5 1 16 Theridiidae Latrodectus hasseltii 11 13 5 12 13 12 13 5 5 7 4 6 2 4 5 3 5 2 9 8 5 6 155 Theridiidae Phorollcidia sp. 01 1 Theridiidae Phorollcidia sp. 02 1 1 1 3 Theridiidae Phorollcidia sp. 03 1 1 1 3 Theridiidae Phorollcidia sp. 04 1 2 1 3 1 8 Theridiidae Phorollcidia sp. 05 1 1 1 1 2 1 7 Theridiidae PllOrollcidia sp. 06 3 3 1 5 3 4 2 5 3 1 30 Theridiidae 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 2 102 Theridiidae Stealoda sp. 02 1 1 2 Theridiidae Steatoda sp. 03 2 1 1 1 1 1 1 I 1 2 12 Theridiidae Steatoda sp. 04 1 I 1 3 Theridiidae Steatoda sp. 05 1 1 1 1 4 N Theridiidae Steatoda sp. 06 1 I -...) \i1 N Table 1 (cont.) 'I 0' Family Genus, species Study sites (arranged approximately north-west to south-east) No. of quadrats 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)

Theridiidae Steatoda sp. 07 3 Theridiidae Steatoda sp. 08 2 2 Theridiidae Steatoda sp. 09 1 1 ..., Theridiidae Steatoda sp. B 2 1 1 2 1 2 2 3 I 21 Thomisidae Diaea sp. 1 1 Thomisidae Sidymella sp. 01 2 2 Thomisidae Sidymella sp. 02 3 Thomisidae Sidymella sp. 03 1 Thomisidae Sidymella sp. 04 2 Thomisidae Sidymella sp. 05 1 2 4 Thomisidae Sidymella sp. 06 2 2 9 s: Thomisidae Sidymella sp. 08 1 ';Jl sp. 09 4 1 2 2 16 Thomisidae Sidymella :t Thomisidae Sidymella sp. 10 1 ...tJ <: Thomisidae Sidymella sp. 11 1 ro Thomisidae Sidymella sp. 12 2 :< '- Thomisidae Stephanopis sp. 01 424 2 3 2 27 Thomisidae Stephanopis sp. 02 1 I s: Thomisidae Stephanopis sp. 03 2 2 5 ~ tJ Thomisidae Stephanopis sp. 04 2 4 3 3 17 z:: Thomisidae Stephanopis sp. 05 1 5 ,..,0 Thomisidae Stephanopis sp. 06 2 r Thomisidae Stephanopis sp. 07 8 z Thomisidae Tharpyna sp. 01 3 ~ Thomisidae Tharpyna sp. 02 1 CJ t: Thomisidae Tharpyna sp. 03 2 2 ::r Thomisidae Tharpyna sp. 04 1 3 ::l. ~ Thomisidae Tharpyna sp. 05 4 3 2 14 ?:1 Thomisidae Tharpyna sp. 06 1 '- Thomisidae Tharpyna sp. 07 1 1 0 Thomisidae Tharpyna sp. 09 2 6 ...t: Thomisidae Tharpyna sp. 10 2 tJ ~ Thomisidae Tharpyna sp. 11 Thomisidae Tharpyna sp. 12 Z Thomisidae Tharpyna sp. 13 r' Trochanteriidae Rebillls sp. 01 2 s:,.., Trochanteriidae Trachyspina madllra ~ ro Trochanteriidae Trachyspina sp. 02 ::l N Trochanteriidae Trachyspina sp. 03 ;;.

I .. t ;p Trochanteriidae Traehyspilla sp_ 04 1 I .... Zodariidae As/eron-complex sp_ 01 2 1 I 1 1 3 9 ::l re'" Zodariidae As/eroll-complex sp_ 02 2 3 5 4 14 0 Zodariidae As/eroll-complex sp_ 03 1 1 3 ..., ....0 Zodariidae 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 '"0'" 0: 1 3 re Zodariidae As/eron-complex sp_ 07 1 1 .... 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 78 Zodariidae Aus/ralu/ien sp_ 02 1 1 Zodariidae Aus/raluliea sp_ 03 1 I Zodariidae Cavas/eron sp_ 01 6 5 6 1 2 2 1 1 1 1 1 1 28 Zodariidae Chilwnena reprobans I I Zodariidae Chilumena sp_ 01 3 1 1 I 1 7 Zodariidae Cyrioctea sp_ 01 I I Zodariidae Genus 1 sp_ 01 2 2 Zodariidae Genus 1 sp_ 02 1 1 1 2 1 1 7 Zodariidae Genus 2 sp_ 01 1 I 1 3 Zodariidae Genus 3 sp_ 01 2 2 Zodariidae Genus 4 sp_ Ol 3 3 Zodariidae Habrolles/es australiensis 1 7 8 Zodariidae J-Jabronestes grimwadei 1 1 1 1 1 2 1 1 3 2 14 Zodariidae J-Jabronestes sp_ Ol 1 1 1 I 2 2 8 Zodariidae Habronestes sp_ 02 2 1 1 5 1 2 2 5 5 5 2 1 1 1 34 Zodariidae liabronestes sp_ 03 2 2 5 3 I 8 11 10 8 2 3 8 3 5 7 5 1 84 Zodariidae Habronestes sp_ 04 4 4 1 6 1 3 10 2 9 9 4 5 5 4 67 Zodariidae Habrollestes sp_ 05 2 1 5 4 3 4 2 1 6 4 2 3 3 1 4 1 2 48 Zodariidae Habronestes sp_ 06 1 1 2 1 4 2 1 3 2 2 1 2 1 1 1 1 26 Zodariidae J-Jabronestes sp_ 07 1 1 1 1 4 Zodariidae Habrones/es sp_ 08 1 1 2 Zodariidae Habronestes sp_ 09 4 2 2 3 9 5 6 5 7 6 6 5 3 4 1 3 3 2 76 Zodariidae Habronestes sp_ 10 1 1 1 2 2 1 2 1 2 1 2 16 Zodariidae Habronestes sp_ 11 1 1 Zodariidae Habronestes sp_ 12 2 1 1 1 1 1 1 8 Zodariidae Habrones/es sp_ 13 1 4 11 2 1 4 1 24 Zodariidae Habronestes sp_ 14 1 2 1 1 1 1 7 Zodariidae Ha/Jronestes sp_ 15 3 3 Zodariidae Halmmestes sp_ 16 2 1 3 Zodariidae Halmmes/es sp_ 17 1 1 2 Zodariidae Habronestes sp_ 18 1 2 2 3 8 Zodariidae Habronestes sp_ 19 1 I Zodariidae Habronestes sp_ 20 1 3 3 7 Zodariidae J-Jabronestes sp_ 21 2 2 3 N Zodariidae Habronestes sp_ 22 3 'I 'I N Table 1 (cont.) '-l 00 Family Genus, species Study sites (arranged approximately north-west to south-east) No. of quadrats 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 29 Zodariidae Habronestes sp_ 24 1 3 4 Zodariidae Habronestes sp_ 25 1 1 Zodariidae Habronestes sp_ 26 1 1 Zodariidae Habronestes sp_ 27 1 1 Zodariidae Habronestes sp_ 28 1 1 Zodariidae Habronestes sp_ 29 1 1 Zodariidae Habronestes sp_ 30 1 1 Zodariidae Habronestes sp_ 31 1 1 Zodariidae Habronestes sp_ 32 2 2 Zodariidae Habronestes sp_ 33 1 1 Zodariidae Habronestes sp_ 34 1 1 s: fJl Zodariidae Habronestes sp_ 35 1 1 :t Zodariidae HalJronestes sp. 36 1 1 2 III ...<: Zodariidae Hal)ronestes sp. 37 1 1 ~ Zodariidae Habronestes sp_ 38 6 5 2 2 1 16 ':< Zodariidae Habronestes sp. 39 1 1 -~ Zodariidae Heptasteron sp_ 01 1 1 2 3 1 2 2 1 2 1 2 2 20 Zodariidae 1 2 2 4 2 1 17 ~ Hetaerica harveyi 5 III Zodariidae Hetaerica sp_ 01 2 4 1 1 3 3 1 15 Q: Zodariidae Kerasteron sp_ 01 2 2 9 8 7 2 4 5 4 1 1 2 47 ,.,0 Zodariidae Leptasteron sp_ 01 1 1 2 r Zodariidae Neostorena sp_ 01 3 4 4 6 5 2223424152 4 1 3 57 Z Zodariidae Neostorena sp_ 02 4 5 4 5 2 1 3 2 4 3 2 35 ~ Zodariidae Neostorena sp_ 03 1 1 1 2 3 2 1 1 4 2 1 3 22 C'l l: Zodariidae Neostorena sp_ 04 2 1 4 1 2 1 11 Er Zodariidae Neostorena sp. 05 2 2 2 6 ~.... Zodariidae Neostorena sp_ 06 1 1 ?' Zodariidae Neostorena sp_ 07 1 1 1 1 2 2 8 ':""' Zodariidae Neostorena sp_ 08 1 1 Cl Zodariidae Neostorena sp_ 09 1 1 1 3 ...l: Zodariidae Neostorena sp. 10 1 1 2 III ~=' Zodariidae Neostorena sp. 11 1 4 1 1 7 Zodariidae Neostorena sp. 12 2 1 1 2 1 1 1 1 2 1 1 14 ~ Zodariidae Neostorena sp_ 13 1 1 2 r' Zodariidae Neostorena sp. 14 1 1 1 2 1 6 s:,., 7: Zodariidae Neostorena sp_ 15 1 1 2 ~ 16 1 1 5 Zodariidae Neostorena sp_ 1 2 N=' Zodariidae Neostorena sp. 17 1 1 it- Araneomorph spiders 279

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r::; r:: ;?j2 ~~ 280 M. S. Harvey, J. M. Waldock, N. A. Guthrie, B. J. Durrant, N. L. McKenzie found during the survey, which all appear to differ several species of Walldel/a and by a single cave- from the species revised by Platnick and Forster dwelling species of Yardiel/a (Gray, 1994). Most (1989): ChaslIlocephaloll sp. 1 (DA, JB, UN and YO), species of Walldel/a occur under the bark of trees, ChasllIocephalo/l sp. 2 (DA, JB, NR and UN) and although Harvey et al. (2000) recorded five species ChaslIlocephaloll sp. 3 (DA, ST and UN). Anapids in pitfall traps in the Carnarvon Basin region were generally absent from the northern and indicating that some species either live permanently eastern quadrats and were more abundant in the on ground habitats (such as in litter or under southern quadrats. stones) or that they move from tree trunks onto the ground at certain times of the year. The present Araneidae survey recorded Walldel/a barbarella Gray. We recorded a variety of araneids, but usually in very low numbers. The majority of araneid species Gallieniellidae are arboreal and, because they are not reliably The Australian members of the gnaphosoid captured in pitfall traps, were excluded from the family Gallieniellidae were recently revised by quantative analyses. (Platnick, 2002) who found a small fauna of five genera and 27 species. Harvey et al. (2000) Cyatholipidae encountered five species in the Carnarvon survey, The family Cyatholipidae occurs in the majority including two species of Meedo. The present survey of the southern continents (Forster et al., 1988; recorded six species, including Meedo houstoni Main Griswold, 2001) and the Australian fauna is which was also found in the Carnarvon survey. represented by only a few named genera. The soft- bodied genera such as Tee1l1ellaarus, Tekel/atus and Hahniidae Toddiana are generally restricted to temperate Hahniids are tiny litter or bark dwelling spiders. woodland or rainforest habitats, while members of Ten species were found in low numbers during the the genus Matilda occur throughout mainland present survey, while no hahniids were found in Australia usually in open woodland habitats. the Carnarvon survey (Harvey et al., 2000). Harvey et al. (2000) found three species of Matilda in the Carnarvon Basin region, and this study found Hersiliidae two species of this widespread and abundant genus. H.ersiliid spiders are generally found on tree trunks where they lie cryptically in wait for their Cydoctenidae prey. Baehr and Baehr (1987,1989, 1992, 1993, 1995, Cycloctenid spiders are restricted to Australia and 1998) have recorded a total of 25 species of Ta1l1opsis New Zealand and are vagrant hunters occurring in from Western Australia, with a further four species leaf litter or under stones and rocks. The Western of Hersilia from northern Western Australia. Nine Australian fauna consists of small spiders of species of Ta1l1opsis were recorded in this survey, uncertain generic affinity. In this survey, Genus 1 most from just a handful of localities, suggesting sp. 1 was found once at DA and Genus 2 sp. 1 was that they are infrequently captured in pitfall traps found once at LK. due to their arboreal lifestyle. For this reason the family was excluded from the quantative analyses. Deinopidae Members of the cosmopolitan family Deinopidae Lamponidae construct spectacular webs that they use to ensnare The Lamponidae are restricted to Australia, New their prey (Raven et aI., 2002). A single species of Caledonia and New Guinea, with two species Deinopis was found sporadically throughout the accidentally introduced into New Zealand; the vast study area at 10 quadrats. Most deinopids are majority of lamponids are Australian (Platnick, generally arboreal and, because they are not reliably 2000). The Lamponidae were revised by Platnick captured in pitfall traps, were excluded from the (2000) who recognised numerous Australian genera quantative analyses. species. During this study 41 species in 13 genera Desidae encompassing all three subfamilies were identified. Spiders of the family Desidae are commonly The Lamponinae was represented by six genera, found in a wide variety of Western Australian Lalllpolla, Lalllpollata, Lalllpollega, Lalllponclll1, terrestrial habitats. This survey recorded 39 species, Ll1l1lpOllinl1 and Ll1ll1pollusa, and 26 species, of which the majority of which were found in few quadrats. 17 were Ll1l1lpolla species. The Centrothelinae were represented by six genera, Asadipus, Bigellditil1, Filistatidae LOllgcpi, NotsodipllS, Prionostemulll and QlICClluic, and Filistatid spiders are found in a variety of regions 14 species. The Pseudolamponinae were in mainland Australia, and are represented by represented by Pscudoll1l1lpolll1 boree Platnick. The ~ ~ ~

Araneomorph spiders 281 genera Pnollostemllm and )/11'1'117)11' were consistent with the broad distributional range restricted to the southern quadrats. recorded Harvey (1995). Ambicodamus marae was only found at DA07, JBI0 and LKI2, each Linyphiidae representing range extensions for this species which Linyphiids were abundant at many sites, but were was previously found to be restricted to high excluded from the quantitative analyses because of rainfall areas, predominately in the karri forests of uncertainties in associating male and female the south coast (Harvey, 1995) and in outlying specimens and in interpreting the taxonomic populations in the Darling Range (Brennan, 1999). significance of slight morphological differences in Ambicodamlls kochi was found at NOlI. the structure of the male and female genitalia. The family is largely unstudied in Australia and Oecobiidae remaInS an enormous challenge in ecological The spider family Oecobiidae is represented in analvses. the Australian region by introduced species (see Santos and Conzaga, 2(03), and this study found Liocranidae Oecobws Ill/ViiS at several sites. Unfortunately this Liocranid spiders are small to medium sized species was coded as "Annulipes sp. 1" and hunting spiders that are moderately diverse in the "Annulipes sp. 2" in the analysis. Australian region. We recorded nine species, and only sp. 12 was found at more than 10 sites. Oonopidae The tiny spiders of the family Oonopidae are an Lycosidae extremely abundant and diverse component of the The spider family Lycosidae is adiverse group Australian fauna, and we found 46 species in a that is poorly known in Australia. The genera varietv of genera including Cmnasomorpha, Cn/meus, Artoria and \/ellatnx have been recentlv revised iV1:11/71l0!'70r7aea, Opopaea, Orchestilla and anew genus. (Framenau and Vink, 2001; Framenau, 2(02) but the remaining genera are still in flux. The vast majority Orsolobidae of the Australian wolf spider fauna has been The Condwanan family Orsolobidae is attributed to the genus Lycosa but these species bear represented in the Australian fauna by members of little affinity to the type species of Lycosa (Vink et several genera, of which Tasmanoonops and aI., 2(02). The systematics of the Australian wolf AlIstraloblls have been found in Western Australia spiders is currently being revised (Framenau, pers. (Forster and Platnick, 1985). Further unnamed comm.). We identified 61 lycosid species in the species of both genera are known (Harvey, present survey, but the case for some of the unpublished data). A single specimen of an identifications is tenuous and the fauna may be unnamed species was found at WH02. smaller. Pararchaeidae Micropholcommatidae The family Pararchaeidae is restricted to Australia 'The tiny spiders comprising the family and New Zealand where they occur in forested and Micropholcommatidae are abundant in Australian woodland habitats. Two unnamed species of forests but have largely remained unstudied except Pararchaea were found in the survey. Pararchaea sp. for the work of Hickman (1944, 1945, 1979) and 1was found at one quadrat whilst Pararchaea sp. 2 Forster (1959). We found several species tentatively was found at avariety of sites throughout the placed in the genus Micropholcolllma. survey area.

Mysmenidae Philodromidae The tinv spiders of the Mysmenidae are only The Australian philodrornid fauna is small (Raven rarely collected in pitfall traps, and only two species et aI., 2(02) with just ahandful of recognised species were found in this survey, each from asingle site. in two genera, Philodromlls and Tibellus. We recognised four species of Tibellus in the present Nicodamidae survey, each collected in very low numbers, The Nicodamidae were revised by Harvey presumably due to their primarily arboreal who found two genera in New Zealand and five preferences. genera in Australia, one of which also occurs in Ne,v Cuinea. The Western Australian fauna is SaIticidae represented by three species, Nicodamlls maillae The Salticidae are the largest spider family Harvey, Ambicodamus marae Harvey and A kochi (Platnick, 20(4) and the Australian fauna is Harvev. Nlcodamus maillae was found to be imperfectly known (Davies and Zabka, 1989). We Widespread throughout the survey area, which is recorded 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 the Theridiidae subject of a separate analysis in this volume Theridiid spiders are a ubiquitous part of the (Guthrie and Waldock, 2004). spider fauna of most regions of the world, and we recorded 80 species. Segestriidae Only three species of Segestriidae were Thomisidae recognised, in contrast to the seven species found in We recorded 31 species of Thomisidae during the the Carnarvon survey. survey, primarily in the genera Sidymella, Stephanopis and Tharpyna. As the majority of these species are Stiphidiidae considered to be arboreal or corticolous and, because The Australasian spider family Stiphidiidae was they are not reliably captured in pitfall traps, they represented in the survey by a single species of were excluded from the quantative analyses. Baiami, three species of Corasoides, a species of Forsterina, and two species tentatively placed in Trochanteriidae Corasoides. Platnick (2002) has recorded a rich trochanteriid fauna in Australia, represented by 14 indigenous Tetrablemmidae genera. We found five species of this family, The spider family Tetrablemmidae is currently including four species of Trachyspina and a single represented within Australia by a single named species of Rebilus, each from a single quadrat. species, Tetrablemma okei from Victoria, but numerous unnamed species occur in eastern and Zodariidae northern Australia (Harvey, unpublished data). A Recent systematic research on the Australian single female tetrablemmid was collected at M002, zodariid fauna has revealed a highly diverse fauna which represents the first record of this family from consisting of numerous genera (e.g. Jocque and southern Western Australia. Baehr, 1992, 2001; Jocque, 1995a,b; Baehr, 2003a,b, in press; Baehr and Jocque, 2000, 2001; Jocque and Tetragnathidae Baehr, 2001; Baehr and Churchill, 2003;). The 117 Tetragnathid spiders are represented in south- species of Zodariidae recorded during the survey western Australia by several genera such as Nephila were the subject of a separate analysis in this and Tetragnatha, as well as a series of poorly known volume (Durrant, 2004), in which there was no metines. This survey found two such metine species discernible relationships between species in low numbers. composition and substrate data.

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

~ l1J c 28 ~ 24 (/) l1J ·0 l1J 24 27 27 16 Et r- 20 Cl. L- 0 l' E 20 26 0 11'1 l1J C ~ l' (ij i' O'i 16 l1J L- 0 16 .Cl L- ~, c 12 z0 IH =48.3, n =294, df =11, P « 0.0001

8 2 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 are displayed. Means are labelled with the number of quadrats sampled in each landform. H = Kruskall-Wallis coefficient, n = number of quadrats, df = degrees of freedom, and P = probability. Because of trap-flooding problems, data from landform unit 1 (freshwater swamps) are excluded. AraneOlIlorph spiders 283

Richness correlation (Kendals Tau -0.26, p = 0.006; r -0.37, A total of 622 ground-dwelling araneomorph P 0.0001; r 0.31, P 0.001, respectively). A spider species were recorded from the 304 quadrats, significant negative correlation was also found an average of 21.9 species per quadrat (s.d. = 8.3). when all araneomorph families, except lycosidae, Figure 3 provides a graphical representation of were included (r = -0.27, P 0.004). Although species richness within each landfonn unit. Natural richness was tightly correlated with numer of saltflats (unit 2) had the fewest species, while specimens (r = 0.56, P < 0.0001), the latter was not dissection valley slopes (units 4 to 8) and durierust significantly correlated with salinity risk. This pavements (unit 12) were the richest. suggests that the significance of the salinity risk The relationship between salinity risk (SAL, as a correlation was related to species richness rather measure of secondary salinity) and species richness than being an artefact of the relatively low numbers was quantified for the ground-dwelling of identified specimens per quadrat (87, s.d. = 60, n araneomprphs overalL and for the three most 55). Scatter-plots of species richness versus EC speciose families separately: Salticidae (121 species), showed that individual outlying points did not Zodariidae (117) and Lycosidae (61). Together, determine the significance of these correlations. these comprised 48(};) of the araneomorph species Using a larger data-set of arachnids and vertebrates recorded during the survey. This analysis was that encompassed the araneomorph data being restricted to the 55 quadrats sampled on landform investigated in this paper, McKenzie et al. (2003) unit 3 (dissection valley floors), the unit with showed that these secondarily salt-affected quadrats sufficient quadrats sampled in each of the four held a species-poor sub-set of the assemblages salinity risk categories to offer a balanced recorded on their unaffected counterparts, as well comparison (McKenzie et 2003). Together the as a few natural salt-flat species. three families analysed comprised 47% of the araneomorph richness recorded on landfonn unit 3. Singleton species and local endemism Salticids and zodariids showed a significant One hundred and eighty-five species of negative correlation with electrical conductivity, araneomorph spiders were recorded at only one while lvcosids showed a significant positive quadrat, representing 29.7°!cl of the entire data-set.

BE WU 1- DN I 1 1 ML 1-1- north band except JB WH 1- MO I NO 1- HY east central I KN I I KL I 2 I QU I-- I 2 Central Band MN I I I NR west central , WK 3 , YO -I , DA 1 I DU I I JB 1 I ST 4 I 3 Southern Band + JBI GP 1 , LK I I I PI -I I I I UN 11- -I , ES 5_1 1_4_1 I I I 1 I 1 I I 0.19 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 genera cornpC)Sltlon of their component quadrats (a total of 166 non-arboreal araneomorph genera). Singleton species were retamed in the 304 quadrat versus 622 species matrix from which the survey area genera lists were derived. 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. Mean and 95% confidence intervals are based on climatic values averaged across the 12 to 13 quadrats within each survey area (H ; Kruskall-Wallis coefficient, n ; number of quadrats, df degrees of freedom, and P; probability).

On average, each quadrat had only one singleton at the four partition-level (Figure 4), was closest to (sd = 1.01, n = 304). Of the 400 non-singleton species the warmest period maximum temperature (see recorded on the 240 quadrats that were both Figure 5). At the five partition-level significant unaffected by secondary salinity and adequately separation of all partitions was best achieved by a sampled, 38% (152/400) were more geographically combination of temperature diurnal range and localised than would be expected by chance alone warmest quarter mean temperature (Table 2). (P < 0.05), compared to 1000 trials using an Together, these two climatic attributes yielded a equivalent number of randomly arrayed records climatic topography and axis only slightly different (quadrat intersections), randomly arrayed, for each from warmest period maximum temperature. species. Overall, 65% of species (261/400) were To see whether substrate attributes emerged as recorded at between 2 and 10 quadrats and 26% of significant correlates with compositional patterns in these (67/261) were more geographically localised araneomorph genera, the 240 quadrats were than would be expected by chance (P < 0.05). These classified in terms of similarities in the presence and 67 species were recorded in an average of 3.3 survey absence of their 162 araneomorph genera. The four areas (s.d. = 1.9), and most were recorded only in genera that occurred at only a single quadrat are the same or adjacent survey areas (51/67 species). excluded from this analysis. The resulting This should not be interpreted as unambiguously dendrogram structure could be interpreted down to implying that all 67 species are locally endemic. the seven partition-level, where six of the seven Records of 66 of the 67 species extended into survey groups could be separated statistically using a areas on the periphery of the study area and further combination of four quadrat attributes: SAL survey work will probably show that many of these (salinity risk), soil pH, warmest quarter mean species extend beyond this periphery. Because of temperature and precipitation seasonality (Table 2). the elongate shape of the study area (Figure 1), 18 Group 5 could not be separated uniquely, but of the 24 survey areas were peripheral. comprised just three of the 240 quadrats.

Patterns in genus composition Patterns in species composition By classifying survey areas in terms of their Figure 6 is the dendrogram structure derived genera composition we could minimise the effects when the survey areas were classified according to of species endemism and substrate, and reduce the their total araneomorph species composition. This influence of study area edge effects and sampling dendrogram reveals a strong geographical pattern noise, on the partition structure of the resulting related to rainfall and temperature gradients. When dendrogram. The compositional pattern, as the structure was examined statistically against summarised by the resulting dendrogram structure survey area climatic attributes, the tightest Araneomorph spiders 285

BE I 1 ML --I-- I I MO I I North-east I WU I 1- I I1 North WH I I I NO II_NW extreme I DN __ NW_I I HY 1 K.N 1 KL I Central-east 1 QU 1- 1 MN I I2 Central JB I I NR Central-west I YO I I WK I 1--1 DA 1 ST South-west 1 UN 1 DU I I3 South LK South-central I PI & east I I GP 1- I I ES SE extreme ___ 1_4 ____ 1 I I I 1 I 0.39 0.46 o54 0.61 0.69 0.76 Coefficient of Dis-similarity

Figure 6 Dendrogram structure derived by classifying the 24 survey areas according to the combined species composition of their component quad rats. Singleton species were retained in the 304 quadrat versus 622 species matrix from which the survey area species lists were derived. relationship was with warmest period maximum similarities in their species composition, the temperature (Table 2), anorth-north-east to south- dendrogram could be interpreted down to the south-west axis related to the hotter summers in the eight-partition level (Figure 8). All partitions could north-north-east (Figure 7). be separated statistically using acombination of When the 240 quad rats were classified in terms of four quadrat attributes: soil salinity, soil pH,

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

------_._---- Data matrix Dendrogram Physical Partitions Separated H (P) Comment Partition Level Attribute

4(Figure 4) mxTwml' all IS (0.0004) Iigurl' (Figure 4) Tdir 1&2vs3vs4vs5 17 (O.0()2) TwmQ Ivs 2 20 (O.O()06)

24Ax622S 4(Figure 6) mxTwml' all 20 (lIlJ002)

240Qx 162C, 7 Sr\L 1-5\s6&7 31 (0.0001) I'

X) H' 1-6vs7&S 37 (()OO()()I) I'll ~ vsI' 35 (O.OOO() I) 1\\ m() I&3 vs 2vs 4\s 5; 6 I' 147 (O.O()()O) ~ 1\s 3 56 (O.OOO()) 286 M. S. Harvey, J. M. Waldock, N. A. Guthrie, B. J. Durrant, N. L. McKenzie Warmest Period Maximum Temperature -28

-29

-<'"--" .... --" -30 '\1l /"-" .. ' ~ ~.

. ~ _...----"""

", ~ . ~ ~ - ~ 0--31 .. ------.--,r--- . Cl) "C , ::J :i: ~RI -32 ...J i ~ ~ i -33 t ,~ ~ ~ i -34 Ui i -38 34 -35 30 114 115 116 117 118 119 120 121 122 123 26 - 22 Longitude (0)

Figure 7 Results of the survey area classification (Figure 6) displayed geographically. Warmest period maximum temperature contours (QC) are superimposed on the survey area map. Colours indicate relationships down to the eight partition-level in the classification dendrogram and are ordered from yellow to blue down the dendrogram (Figure 6). Highest temperature values are in the north. warmest quarter mean temperature and rainfall considerable component of the fauna that ranges into seasonality (Figure 8, Table 2). other areas of southern Australia or into the central Warmest quarter mean temperature is tightly or northern regions of Western Australia. intercorrelated with warmest period maximum Regrettably, many other families are taxonomically temperature (Kendall's Tau = 0.92, P<0.001). Thus, poorly known and the identifications made during the four classification analyses yielded similar this study cannot be readily correlated with the results (Table 2), although salinity emerged as an faunae found in other regions of the country. important factor in the two quad rat classifications There is limited extrinsic knowledge available on (either as salinity risk or as soil electrical patterning in Western Australia araneomorph conductivity), along with soil pH. spider communities, which precluded a multivariate statistical approach to the analysis. Relationships revealed by such an analysis could be DISCUSSION artefacts of sampling design. Also, many of the D1e spider fauna of southern Western Australia environmental attributes recorded for each study consists of rich assemblages (Brennan and Majer, in quad rat were strongly intercorrelated. Thus, to press; Brennan et al., in press), with numerous genera generate hypotheses for future studies we restricted and species showing varying degrees of regional our analyses to aunivariate approach. endemicity. Knowledge of the detailed taxonomy Overall araneomorph diversity patterns across the and distributions of this fauna is restricted to just a wheatbelt study area conformed most closely to a few families (see Introduction). Many of these combination of soil salinity, asummer mean studies show that whilst species endemism in temperature gradient with anorth-south axis, and a southern Western Australia is high, there is also a precipitation seasonality gradient with an east-west Araneomorph spiders 287

BEOI-5,7-9,12,13, DNOI,2,6,8 1 KNIO, MLOI-13, MN03,4,7,1O, 1 MOOI,3,5,6,9-12, NOOI-7,1O,11 1 1 PI05, QU04, WH04,5,7,8,11,13 1 WUOI-6,9-12, Y002,1O 1 -1- DN05, DU02-4,9-13, ES07,9 I GPOI-4,6,12, HY01,3-7,9,1O I JBOl,2,5-IO,12,13, KL03 -12 I KN02-6, 11, LK02,4-6,8, 10-13 I 2 MN01,5,6,11, NROI-4,6,8-12 1 PIOI,6-9,12, QUOl-3,6,9-12 1 ST09,10, WH03,6,9,IO, 1 WKOI,6-11,13, Y001,4-8,12,13 1 1 -1-- DN03,4,9-12 3

DAOI,3,4,6,8-11,13, DUOI 1 GP08,9, LK07, NR05,7 4 1 STOI ,6, 13, UN03, Y003 I I DAI2, ESOI,2,5,8,11 I GP13, PI13, ST02-5,7 5 1 UNO I,2,5-7,9-12 1 1 GPIO, PIll 6 1 I ------1-1----- BE 11, DU05, ES06, HY02, I I KLOI, LK03, MLl2, MN09, I 7 1 M004, ST08, WU08 1 1 -1----- I GP05 _1_8 1 1 II 1 1.24 1.52 1.79 Dis-sirnilarity Coefficient

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 saltflats (LF 2), and the three that were inunduted for long periods during the trup program (PI04, UN13 und WK02) axis. As pointed out in Results, these two climatic Tertiary Plateau is much more dissected in the west attributes in combination yield a climatic due to their different paleoclimatic histories. Also, topography and axis only slightly different from the western zone has a rejuvenated drainage system warmest period maximum temperature. However, that flows westwards, while the eastern zone's the climatic components of this pattern could also ancient drainage system comprises chains of salt be explained in terms of broadscale landscape lakes and follows Tertiary paleodrainage lines that relationships. As pointed out in Methods, the ES are orientated northwards and/or eastwards (e.g. and part of the ep survey areas are not actually on Beard, 2000). These broad landscape differences the Yilgarn Craton; nor are NO and DN in the more or less correspond to the dendrogram north. Furthermore, Mulcahy and liingston (1961) partitions analysed (rable 2); deductive studies divided the Craton in the central part of the study such as this do not eliminate alternative hypotheses area into eastern and western zones because the (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 of species define habitats, not vice versa. Correlations specialist halophilic araneomorphs (species group with environmental parameters such as those 18 in Appendix 4). This is not surprising because revealed in this study, are indirect consequences of groundwater dating in south-western Australia species niche requirements rather than directly (Commander et ai, 1994) and sedimentary studies causal to their presence/absence at sites. on gypsum accumulation in Lake Lefroy by Zheng et al (1998) suggest that most halite lakes have been saline for up to a million years (and earlier phases ACKNOWLEDGEMENTS of salt lake formation in this old and geologically We thank J.K. Rolfe, W.P. Muir, P. van Heurck stable landscape are probable). and N. Hall for assistance in installation and This study revealed an inverse relationship operation of the pit traps; L. King, E. Ladhams and (correlation) between araneomorph species richness P. van Heurck for assistance with sorting, data and soil salinity, both natural salinity and the compilation and field sampling; J.K. Rolfe for data secondary salinization induced by agricultural base development and interrogation, B. Baehr for clearing during the last 100 years. Given that salt- assistance in the identification of some Zodariidae; affected sites in our study contained a and M.R. Williams and B. Brand for statistical compositional sub-set of their unaffected advice. We thank two anonymous referees for their counterparts (McKenzie et al., 2003), the effect of comments on the manuscript. Funding for this secondary salinity on araneomorph richness (and study was provided as part of the State Salinity composition) is essentially negative. While the Strategy. araneomorphs belonging to the family Lycosidae (wolf spiders) showed a positive correlation, the relationship is probably not causal. Individual REFERENCES lycosid species, both in Australia and elsewhere, are Austin, M.P. (1984). Problems of vegetation analysis for often confined to particular habitats such as riverine nature conservation. In Myers, K. and Margules, eR margins, sand dunes and salt lakes (McKay, 1979; (eds), Survey methods for nature conservation, vo!. 1: Hudson and Adams, 1996; Moring and Stewart, 101-130. CSIRO Division of Water and Land 1994). Generally, Main (2001) characterised lycosids Resources, Canberra. as adaptive opportunists that favour open Austin, M.P. (1991). Vegetation theory in relation to cost- environments, and predicted that the family would efficient survey. In Margules, CR. and Austin, M.P. increase in lands cleared for agriculture. As with (eds), Nature conservation: cost effective biological surveys clearing, rising ground-water salinity results in a and data analysis: 17-22. CSIRO Division of Wildlife and Ecology, Canberra. net loss of vegetation cover and, in the wheatbelt, Austin, M.P. and McKenzie, N.L. (1988). Data analysis. our data suggest that these conditions favour In Gunn, RH., Beattie, ].A., Reid, RE. and van der lycosids, at least initially. Thus, salt affected sites Graaff, RH.M. (eds), Australian soil and land survey (mainly the woodland sites on the dissection valley handbook: guidelines for conducting surveys: 210-232. floor, landform 3) hold their lycosid diversity and, lnkata Press, Melbourne. at least in the initial stages of secondary salinity, Baehr, B. (2003a). Revision of the tropical genus may be further enriched through colonisation by Tropasteron gen. novo of north Queensland (Araneae, saltflat specialist species such as Lycosa salifodina. Zodariidae): a new genus of the Asteron-complex. The only substrate patterns that emerged dearly Memoirs ofthe Queensland Museum 49: 29-64. from the quadrat analyses were related to salinity Baehr, B. (2003b). Three new endemic genera of the attributes. No distinct, study area-wide patterns in Asteron-complex (Araneae: Zodariidae) from composition emerged that related to other substrate Australia: Basasteron, Euasteron and Spinasteron. attributes (e.g. soil texture, chemical attributes etc). Memoirs of the Queensland Museum 49(1): 1-27. A plausible explanation is that local endemism, as Baehr, B. (in press). Revision of the new Australian genus noted in the salticid and zodariid components Holasteron (Araneae, Zodariidae): taxonomy, (Durrant, 2004; Guthrie and Waldock, 2004), phylogeny and biogeography. Memoirs of the Queensland Museum. perhaps exaggerated by local covariance in population densities etc, masked any such Baehr, B. and Baehr, M. (1987). The Australian subtleties. Hersiliidae (Arachnida: Araneae): taxonomy, phylogeny, zoogeography. Invertebrate Taxonomy 1: Broader relationships with soil-landscape zones 351-437. and climatic gradients were discussed earlier. In Baehr, B. and Baehr, M. (1989). Three new species of any case, clear-cut patterns should not be expected genus Tamopsis Baehr & Baehr from Western because a species' presence or absence at a site is Australia (Arachnida, Araneae, Hersiliidae). Second actually determined by physiological limits and supplement to the revision of the Australian availability of suitable resources in combination Hersiliidae. Records of the Western Australian Museum with guild, predation and other ecological 14: 309-320. processes. Rosensweig (1992) pointed out that Baehr, B. and Baehr, M. (1992). New species and new Araneomorph spiders 289

recmds ot genus B,ll'hr & B'll'hr (Arachmda, Biodiversitv of jarrah torest spidl'l's at larr'lhdalc', Ar,1Iw<1(', Iil'rsiliidae). Ihird suppil'ml'nt to tlw Western Australia: richness, endemicit\, and revision of till' Austr,lIian Ikrsiliidal'. I\.eco,d.s o[ lill' taxonomic knowledge. COIISl'n'lltltlll [i1l)lo,,,I;' WI',;lerll AII.'lmlillll M11.'1'11 III 16: ~ . Bureau ot ML'tl'orologv (2001). What is till' wl'ather Baehr, B. and Baehr, M. (1993). New species ,11ld Ill'\\' usu,lllv like) Climate averages fnr We,;tern Australian rel'ords of Iil'rsiliidae tTom ;\ustr,llia, with an sitl's. htt ww.bom.gov.,lu/clim,ltc/avera updalL'd kev to all Australian specil's (Arachnid'l: tablcs/ca_ wananws,sht ml. ;\raneae: Iil'rsiliidae). Fourth supplement to the Chin, RJ (19Sh). Kt'llerl)arill, Wl'stem AlIstmllll, I 25() revision ot the Australian Hersiliidae. 1\I'(o,d,; of till' ()(I(I .sail'.s. C;eologic,ll Survey ot Wl'slL'rn We,;terll A/I,;lmlil/lI /\:II1SI'IIIII 16: ~ . Austr'llia, Perth. B;lL'hr, B. and Baehr, M. (1995). New specil's and new CIMke, K.I\. (J993), ~ multivariate records of Ilersiliidae from Australia, with an 'lnalvses of changes in communit" structure, updated key to all Australian species (Arachnida: Au.sfmlillll 10uI'II1l1 of 6: 163-17-1. Ara ne;1(': Ilcrsi Iiid ,le). Fifth su pp lenwn tto the Commander, [).I'., Fifield, L.K., Thorpe, I'M, IJavie, 1\.1'., revision of till' ;\ustralian Ilersiliidae. I\l'cord' 01 Iill' Bird, J,I\. andlurIlL'r, J.V. (1994). ~ and We,;ll'm AII,;lml/l1l1 /Vll1';l'/11I1, S/lI'I'Il'II/l'1I1 52: 107-IIS. ~ 1-1 measurements on hvpersaline groundwater Baehr, B. and Baehr, !'v1. (I 99S). New species and new in Tertiarv paleochannels near Kalgoorlie, Western records of Hersiliidae from Australia, with an Australia. Professional I'apers 37: . ~ . Geological updated key to all Australian species (Arachnida: Survey of Western Australia, Perth, Araneae: Hersiliidae), Sixth supplement to the Commander, D.I'., 5choknecht, N" Verboom, W, and revision of the Australian Hersiliidae, I\ecord,; of Ihe Caccetta, I) (2002). The geology, physiology and soils Wl',;lt'm A/I,;lrl/lil/lI A1/1,;e/l1l/ 19: U-3S, of wheatbelt valleys. 111 V. Read & Associates (eds), Deillillg wilh slllillilll ill iclIClll/)c!1 ProICSSI'';, Baehr, B. and Churchill, '1',13, (2003). Revision of the I'rosl'cct,; lllld I'mlliml 0l'tiOIl';: 1-21, Waters and Rivers endemic Australian genus (Araneae, Commission, Perth (Available on CD), Zodariidae): taxonomy, phylogeny and Czekanowski, J. (1932). Coefficient of racial likeness, und biogeography.llli'erli'lJrtllt' 17: 641-h65. durchschnittliche differenz. AlIlhrol'ologis(/Ii'l' flll_CI"" Baehr, B. and Jocqut>, R. (2000), Revisions of genera in the 9: 227-249. ~ (Araneae: Zodariidae), The new Davies, V.'F. (1994), The huntsman spiders flctcrol'oda genera Cm'l/,;leroll and Milll/,;Ieroll. Records of lIle Latreille and Yiilllhi gen. novo (Araneae: We.slem A/lslmliall Muse/lll/ 20: 1-30. Heteropodidae) in Australia, Mellloirs of Ille Baehr, B. and Jocqut>, R, (2001). Revisions of genera in the Queellsllllld MuseulII 35: 75-122, ~ (Araneae: Zodariidae): new genera Davies, V.T. and Zabka, M. (19S9). lIlustrated keys to the Pell la,; It'roil, Pheilasleroil, Lel'lasll'roll and S/lIJI/Sleroil. genera of jumping spiders (Araneae: Salticidae) in ,Mell/oirs of the Q/lel'II,;laild M./lSeUIII 46: 359-385. Australia. l\:lclllOirs of' fhe QueclIslnlld Musc/l1I/ 27: IS9- Baxter, J.L. and Lipple, S.L. (1985), Perell!ori, We,;tem 266, A/I,;lmlia 1:250 ()()O seolosiclIl ,;eries. Geological Survey Durrant, B.J. (2004), Biogeographical patterns of zodariid of Western Australia, Perth. spiders (Araneae: Zodariidae) in the wheatbelt Beard, J5. (19S0). A new phytogeographic map for region, Western Australia. RClord,; of' Ihe Wesfern Western Australia, Weslel'll Allslmliall Ilerl)lIri/l1l/ Auslmlinll AlusCU1I1 S/lI'I'ICIIIClIt 67: 217-230 Re,;earch Nole,; 3: 37-5S. Durrant, B.J. and Guthrie, N.A. (2004). Faunas of Beard, J.5. (19SI), Vesetatioll ';/lri'elj of We,;lt'm I ~ unflooded saline wetland floors of the Weskrn Swail 1 1 000 O()() series. Universitv of Australian wheatbelt. [\ccords of Ihc Wcstern Austmliall Western Australian Press, Nedlands. MuseulII S/ll'l'lclllclIl 67: 231-256.

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Electronic appendices are on CD inside the back cover