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DOI: 10.18195/issn.0313-122x.61.2000.295-321 Records of the Western Australian Museum Supplement No. 61: 295-321 (2000).

Araneomorph from the southern Carnarvon Basin, Western : a consideration of regional biogeographic relationships

Mark S. Harveyt, Alison Sampeyl,2, Paul L.J. West1,3 and Julianne M. Waldock1

1 Department of Terrestrial Invertebrates, Western Australian Museum, Francis Street, , 6000, Australia 2Present address: Lot 1984 Weller Rd, Hovea, Western Australia 6071, Australia 3Present address: Halpern Glick & Maunsell Pty Ltd, John Tonkin Centre, 629 Newcastle St, Leederville, Western Australia 6007, Australia

Abstract -A survey of the ground-dwelling araneomorph assemblages of the Southern Carnarvon Basin revealed a total of 33 families. Apart from the Gnaphosidae and Zodariidae which were not analysed due to time- constraints, we recognized a total of 285 placed in 146 genera. Very few taxa could be assigned to existing genera or species, reflecting poor taxonomic knowledge of many groups of spiders. Patterns in species composition across the study area were correlated with rainfall gradients, and a discrete claypan fauna was detected. Vicariance events seem to explain part of the patterning evident. However, strongly localised patterns in species composition were also evident.

INTRODUCTION Hartmeyer, 1907-1908). Modern authors had Araneomorph spiders constitute a large contributed only a further nine species (Baehr and proportion of total diversity, with 90 Baehr, 1987, 1992, 1993; Harvey, 1995; Hirst, 1991; recognized families and an estimated 35 000 Jocque and Baehr, 1992; Levi, 1983; Main, 1987; described species (Coddington and Levi, 1991; McKay, 1975, 1979), although numerous additional Platnick, 1997). They differ from all other spiders, species were known from the area, based mostly the which consists of the sole Recent upon specimens lodged in the Western Australian , and the which Museum (M. Harvey, unpublished data). A consists of the trap-door spiders and their relatives, recently published survey of the by the orientation of the fangs which bite inwards. (Platnick, 2000) has recorded an additional 15 The Australian fauna is represented by 68 families species from the area, mostly based upon (R. Raven, unpublished data), of which 56 have specimens collected in the present survey. been recorded from Western Australia (M. Harvey, This paper summarises the first quantitative unpublished data). Three of these - , assessment of patterns in the composition of the and - are solely represented araneomorph spider communities in the southern by introduced species, leaving a total of 53 families Carnarvon Basin, Western Australia, based upon a of indigenous species. These 53 families are detailed pitfall-trapping program conducted over a unevenly distributed across the State, with some range of major geomorphological units of the study restricted to the temperate south-west corner and area. The composition of the spider community, others known only from the northern tropical represented by species occurrences, was analysed regions. The total number of spider species in terms of a number of physical environmental expected for Western Australia is difficult to attributes to explain the variation across the study estimate, but 2000-3000 species is predicted (M. region. Harvey, unpublished data). Prior to our study, the spider fauna of the MATERIAL AND METHODS southern Carnarvon Basin had been examined only in an ad-hoc way (Table 1). The first Study Area published records were from the Michaelsen and The southern Camarvon Basin study area covers Hartmeyer Expedition of 1905, which recorded 21 some 75 000 km2 situated on the central region of species from the southern Carnarvon Basin Australia's west coast. It is centred on Shark Bay (Simon, 1908, 1909), collected at four terrestrial and extends from the Murchison River in the south localities: Denham, Brown Station (on Dirk Hartog to the Minilya River in the north, and eastwards to Island), Baba Head and Tamala (Michaelsen and beyond Junction (Figure 1). It covers the 296 M.S. Harvey, A. Sampey, P.L.J. West, J.M. Waldock

113°

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Figure 1 The southern Carnarvon Basin study area. The thirteen survey areas are indicated by two letter codes (see text), and at each area, the quadrat locations indicated by black circles. ~ Table 1 Araneomorph spiders previously recorded from the southern Carnarvon basin. ;I>... ::sll> /1) 0 Species Family General distribution Distribution within southern Authority S ...0 Carnarvon basin '"0 Argiope protensa L. Koch Araneidae Widespread across arid and semi-arid Australia. Some localities in or near study area, e.g. Levi (1983) =r r.tl Milly Milly, Newmarracarra Homestead, '"0.... Kalbarri, Gee Gie Outcamp, p.. .../1) Carrollgouda Well tIl Austracantha minax (Thorell) Araneidae Widespread and locally common in many parts of Shark Bay area. Waldock (1991) Australia, including the Shark Bay area (Waldock, 1991). Dolophones conifera (Keyserling) Araneidae Originally described from , this species was Brown Station, Dirk Hartog Island. Simon (1908) recorded from Dirk Hartog Island by Simon (1908). Larinia eburneiventris Simon Araneidae Originally described from Dirk Hartog Island and North Brown Station, Dirk Hartog Island. Simon (1908) Fremantle, WA. Paraplectanoides cerula Simon Araneidae Originally described from Dirk Hartog Island, and not Brown Station, Dirk Hartog Island. Simon (1908) subsequently recorded in the literature. Clubiona laudabilis Simon Clubionidae Known only from the locality, Denham. Denham Simon (1909) Clubiona robusta L. Koch Clubionidae Originally described from the Swan River (Koch, 1873), Brown Station, Dirk Hartog Island Simon (1909) Simon (1909) recorded it from several localities, including Dirk Hartog Island. houstoni Main Clubionidae Known only from asingle male collected 10 km ESE of Near Meedo Station Homestead Main (1987) Meedo Station Homestead Badumna candidus (L. Koch) (as The synonym P. tubicola was described Simon (1908) Phryganoporus tubicola Simon) from Denham by Simon (1908). Badumna veliferum (Simon) (as Desidae Known only from the type locality, Brown Station, Dirk Dirk Hartog Island. Simon (1908) Aphyctoschaema veliferum Simon) Hartog Island. Homoeothele micans Simon Gnaphosidae Described from Denham and Moonyoonooka, WA, and Denham Simon (1908) not subsequently recorded in the literature. Megamyrmaecion perpusillum Gnaphosidae Described from Tamala and Wooroloo, WA, and not Tamala, Edel Land Simon (1908) Simon subsequently recorded in the literature. Tamopsis depressa Baehr and Baehr Hersiliidae Known only from Badja Homestead and near Denham, Ajuvenile from near Denham was Baehr and Baehr Western Australia, and central tentatively assigned to this species by (1992, 1993) Baehr and Baehr (1992) Tamopsis occidentalis Baehr and Hersiliidae Widespread in western Pilbara, extending as far south as Crossing, near Baehr and Baehr Baehr Shark Bay Camarvon (1987, 1992, 1993) Holconia nigrigularis (Simon) Heteropodidae Originally recorded from Tamala and Northampton by Recorded from Tamala Homestead by Simon (1909), Hirst Simon (1908), Hirst (1991) found it to be widespread in the Simon (1909) (1991) semi-arid and arid regions of southern and central Australia Dlios calligaster (Thorell) Heteropodidae Originally described from eastern Australia, Simon (1908) Denham Simon (1908) recorded it from Denham, Broome Hill, and possibly the Murchison District. Asadipus ban;iwarn Platnick Lamponidae Restricted to central W.A. Bidgernia Station, Boolathana Station, Platnick (2000)

N '"'I Table 1 (cont.) N \0 00 Species Family General distribution Distribution within southern Authority Carnarvon basin Bush Bay, Cape Cuvier, Edel Land, Francois Peron National Park, Kennedy Range, Mardathuna Station, Zuytdorp Asadipus phaleratus (Simon) Lamponidae Originally described from several localities in W.A.; Big Lagoon, Cape Cuvier, Edel Land, Platnick (2000) subsequently recorded from large areas ofarid W.A., Qld Francois Peron National Park, Kennedy and SA Range, Mardathuna Station, Meedo Station, Nanga Station, Nerren Nerren Station, Zuytdorp Asadipus woodleigh Platnick Lamponidae Restricted to central W.A. Bidgemia Station, Boolathana Station, Platnick (2000) Bush Bay, Edel Land, Kennedy Range, Mardathuna Station, Meedo Station, Nanga Station, Woodleigh Station Bigenditia zuytdorp Platnick Lamponidae Widespread across arid Western and . Bidgemia Station, Boolathana Station, Platnick (2000) Bush Bay, Kennedy Range, Mardathuna Station, Nerren Nerren Station, Woodleigh Station, Zuytdorp cylindrata (L. Koch) Lamponidae Widespread across ; accidentally Boolathana Station, Zuytdorp Platnick (2000) introduced into New Zealand. lAmpona quinqueplagiata Simon Lamponidae Described from Dirk Hartog Island and Boyanup, WA; Brown Station, Dirk Hartog Island, Simon (1909); subsequently recorded from numerous localities in central Bidgemia Station, Billabong Roadhouse, Platnick (2000) WA. Boolathana Station, Bush Bay, Cape Cuvier, Peron Peninsula, Edel Land, Faure Island, Francois Peron National ~ Park, Kennedy Range, Mardathuna ~ Station, Monkey Mia, Nerren Nerren :I:~ Station, Woodleigh Station ~ lAmpona whaleback Platnick Lamponidae It) Known from Mt Whaleback and Bush Bay, WA. Bush Bay Platnick (2000) ~ lAmponata daviesae Platnick Lamponidae Widespread across Australia. Tamala Station Platnick (2000) lAmponella kimba Platnick Lamponidae ?> Widespread in scattered localities across Australia. Boolathana Station, Bush Bay Platnick (2000) en lAmponina elongata Platnick Lamponidae Widespread across southern arid Australia. Francois Peron National Park, Bidgemia Platnick (2000) ~ S Station, Boolathana Station, Bush Bay, "'0 It) Cape Cuvier, Francois Peron National ~ Park, Kennedy Range, Mardathuna ;c t"" Station, Meedo Station, Nanga Station, ~ Woodleigh Station ~ It) lAmponina scutata (Strand) Lamponidae Widespread across arid Australia. Bidgemia Station, Boolathana Station, Platnick (2000) (/) Cape Cuvier, Cape Lesueur, Peron ~ '- Peninsula, Edel Land, Faure Island, ~ Francois Peron National Park, Kennedy ~ Range, Mardathuna Station, Nanga ~ Q.. Station, Nerren Nerren Station, 0 n- Woodleigh Station, Zuytdorp ~

30Q M.S. Harvey, A. Sampey, P.L.J. West, J.M. Waldock northern half of the Irwin District of the 1995, and in August 1995 when they were removed. Southwestern Province, and the southern half of the Samples were returned to the laboratory, washed in Carnarvon District of the Eremaean Province water and stored in 75% ethyl alcohol until being (Beard, 1980). The vegetational characteristics are sorted. Target organisms were removed, labelled documented by Keighery et al. (2000), but briefly and placed in separate vials or jars. Five wet pitfall some of the southern survey sites support traps were placed in each of the 63 quadrats, the woodlands of Eucalyptus and Callitris, whereas the total number of traps utilised being 315. Over the northern survey sites support Acacia shrublands to 12-month trapping period, we had the equivalent of low open woodlands on fine textured soils and some 115 000 trap nights. The 12-month trapping hummock grasslands on red sand surfaces. The period allowed for the full sampling of all seasons, demarcation between eucalypt woodlands and and reduced the suspected bias that may occur in Acacia shrublands defines the boundary of the restricted sampling periods. Southwest and the Eremaean Botanical Provinces Live specimens were also removed from dry (Beard, 1976, 1980, 1990), and is often referred to as vertebrate pitfall traps (see McKenzie et al., 2000) in the mulga-eucalypt line. the same locations during each 5-day trapping The physical environment of the Carnarvon Basin program (September-Qctober 1994, May-June 1995) is outlined by Wyrwoll, Stoneman, Elliott and and preserved in 75% ethyl alcohol. Sandercock (2000) and the climate by Wyrwoll, Specimens were also hand collected during the Courtney and Sandercock (2000). The climate in the field program, but to date we lack the resources to southern half of the study area is semi-arid with sort and label this material. All specimens were temperate weather systems and an annual rainfall lodged in the Western Australian Museum, Perth. of c. 240-340 mm. The central and northern areas are arid tropical although temperate systems affect Data Analysis its coastal areas in winter; the annual rainfall ranges The data matrix (Table 2) comprised the presence from c. 200-240 mm (Wyrwoll, Courtney and and absence of species collected in both wet and Sandercock, 2000). dry pitfall traps at each quadrat, rather than their relative abundance. The latter technique was not Field Sampling adopted as there are few data available on the The survey was based upon 63 quadrats sampling bias in the use of pitfall traps (but see distributed across 13 survey areas (Figure 1): Churchill, 1999). The computer package PATN Boolathana (Ba), Bush Bay (BB), Cape Cuvier (CD), (Belbin, 1993) was used to explore the data matrix, Edel Land (EL), Gascoyne Junction (GJ), Kennedy exposing patterns in species composition. The Range (KE), Mardathuna (MA), Meedo (MD), Czekanowski association measure was used to Nanga (NA), Nerren Nerren (NE), Peron Peninsula compare the quadrats according to similarities in (PE), Woodleigh (WO) and Zuytdorp (ZD). At each their species composition, and the 'Two-step' survey area there were five quadrats, except for association measure was used to determine the Cape Cuvier (six quadrats) and Edel Land (two quantitative relationship between each pair of quadrats). The quadrats were positioned to cover species. The structure of the resulting association the geographical extent of the study area, represent matrices was displayed as dendrograms using a the array of stratigraphic units present and provide modified 'unweighted pair group arithmetic some pseudoreplication of its main subsfrate types. averaging' (DPGMA) hierarchical clustering Sampling was conducted with the aid of two strategy (Sneath and Sokal, 1973), and the data different types of pitfall traps: 'wet' traps and 'dry' matrix re-ordered accordingly, as a two-way table. traps. The biological patterns revealed by these analyses Wet pitfall traps consisted of 25 litre buckets (300 were investigated in terms of a set of attributes x 400 mm) dug into the ground, with the top flush related to the physical environments of the quadrats with the soil surface. Each was fitted with a lid into Wyrwoll, Stoneman, Elliott and Sandercock (2000), which five 10 cm diameter holes were cut. A piece utilising the GSTA module in PATN, and assessed of chicken wire (c. 1 cm mesh) was suspended statistically with Kruskal-Wallis K-sample tests. several cm below the lid to allow any vertebrate Eleven climatic attributes were derived for each which ventured into the trap to escape. This may quadrat using ANUCLIM (McMahon et al., 1995). have enabled larger invertebrates such as beetles Soil and geomorphic attributes were also recorded and spiders to escape although a 2.5 cm gap was from each quadrat (Wyrwoll, Stoneman, Elliott and left between the wire and the wall of the bucket. Sandercock, 2000). The 17 soil chemical and texture Each trap was equipped with approximately 3.5 values used herein were derived from sub-samples litres of preserving fluid (3 1 ethylene glycol, 40 ml collected at a depth of 5 - 10 cm from 20 - 30 formalin, 350 ml water). The traps were opened in regularly dispersed points on each quadrat, then August 1994, and the contents were cleared in bulked (Appendix D). Significant correlations September-Qctober 1994, January 1995, May-June between these physical attributes were identified Araneomorph Spiders 301

using Kendall's rank correlation coefficient Clubionidae (Kendall's tau). Physical attribute names and codes Six species currently assigned to ~ Clubionidae are listed in Appendix 2. were collected, including aspecies of and two species of Clubiona. Spiders of these genera are generally arboreal, and it is likely that they were RESULTS under-represented in the pitfall trapping program. Over 3600 vials of spiders containing more than The remaining three species are only doubtfully 12 000 specimens were sorted and identified during included in the Clubionidae, and their systematic this survey, and the data are summarized in Table 2. status is currently under revision (R.J. Raven, pers. A total of 33 families were represented which were comm.). A single species of Fissarena was taken classified into 146 genera and 285 species. As from three quadrats (B05, CU4, CU5), belonging to discussed below, time constraints precluded the agenus which has only recently been described identification of the Zodariidae and most of the from the Simpson Desert, south-western Queens- Gnaphosidae which, if included, would probably land (Henschel et al., 1995). Two species of Meedo increase the total number of species to over 350 (M. were collected, including Meedo houstoni Main fro)ll Harvey, unpublished data). Some of the species 15 northern quadrats, and an undescribed species, collected in the pitfall traps were clearly not part of M. sp. 1, from three southern quadrats (Table 2). the usual ground-dwelling fauna, and were most The former species was previously only known probably under-represented in the sampling regime from the holotype collected near Meedo Homestead utilised during this survey. Examples include web- (Main, 1987), and is now known to be Widespread building species such as Austracantha minax (Thorell) throughout the study area north of the mulga- and Nephila edulis (Labillardiere) which rarely descend eucalypt line, whereas the undescribed species is out of their webs, and species which mostly reside on restricted to the south-western portion of the study the surfaces of trees (e.g. species of Tamopsis). area. Nevertheless, these species are included in the matrix and were analysed as part of the total fauna. Many taxa could not be assigned names, at either Seven corinnid species, tentatively placed within the generic or specific levels, areflection of the six genera, were collected. 5, sp. 1was taken deficiency in our taxonomic knowledge of spiders at only asingle quadrat, but the others were more in Western Australia. It is possible that many of the widespread (Table 2). Corinnids are ground- species are undescribed, but the plethora of poorly hunting spiders. known names in the scientific literature (such as Simon, 1908, 1909), combined with alack of modem taxonomic revisions, hampers our understanding of Three undescribed species of the Australian these taxa, especially with regard to their endemic genus Matilda were collected, M. sp. 1at I biogeographic origins. three quadrats, M. sp. 2at one quadrat and M. sp. 3 at one quadrat (Table 2). These are all small spiders and may be under-represented in the study due to The Spider Taxa their lack of mobility. Araneidae Fifteen araneid species in eight genera were recorded (Table 2), including species which are Asingle species of Deinopis was collected across known to be widespread outside of the study area, five northern coastal quadrats. Deinopids cast such as Argiope protensa L. Koch (Levi, 1983) and unique catching webs, often close to the ground, Austracantha minax (Thorell) (Waldock, 1991). with which they seize prey. Araneids spin orb-webs to capture prey and are generally poorly represented in pitfall catches, as Desidae they rarely venture on to the ground. Of particular Six species attributed to the Desidae were interest was the large number of species of collected, including Badumna insignis (Thorell), a Dolophones, some represented by numerous adult species known to be Widespread across much of male specimens from avariety of quadrats. Australia, and five further species attributed to Members of this orb-weaving spider genus different putative genera. Some de sids are often generally remain in webs during the evening, and found on the bark of trees or logs, but many others remove their web to take up acamouflaged position spin small webs near to the ground. on atree trunk or branch during the day. It would appear from the results presented here that male Dolophones disperse by travelling along the ground Three dictynids, tentatively assigned to separate at least part of the time. genera, were recorded, but only Genus 2, sp. 1was Table 2 The frequency of occurrence across quadrats of all araneomorph spiders (except Gnaphosidae and Zodariidae) collected in both wet and dry pitfall traps for each of the 13 survey areas (see text for explanation of abbreviations). There were five quadrats at each survey area, except for CD which had six, and EL which had two.

Familv Genus / species BB BO CU EL GT KE MD MR NA NE PE WO ZU 123451234512345 6 1212345123451234512345 123 4 5 1 2 3 4 5 123 4 5 1 2 3 4 5 123 4 5 Arnneidae A'l!.iop, prot,nsa L. Koch XXX xx X xx XX X X X X Araneidae Allstmeantha minax (fhorell) XXX X Araneidae Dolophones sp. 1 XXXXX X X XX XXX X X Arnneidae Dolophones sp. 2 XX XXX X X X Arnneidae Dolophones sp. 3 XX Araneidae Dolophones sp. 4 XXX Arnneidae Dolophones sp. 5 X Araneidae Eriophom sp. 1 XX X X Araneidae Eriophora sp. 2 X Arnneidae Eriophom sp. 3 X Araneidae Genus 1, sp. 1 X X X Araneidae Genus 1, sp. 2 X XX Araneidae Genus 2, sp. 1 XXX X X Arnneidae H'lIrod,.r sp. 1 X Araneidae Larinia sp. 1 X Clubionidae Ch,imeanthillm sp. 1 XX X Clubionidae Clllbiona sp. 1 X X Clubionidae Clllbiona sp. 2 X X XX Clubionidae Fissarena sp. 1 X XX Clubionidae Meedo hOllstoni Main X XXX X X XX X XX XX XX Clubionidae Meedo sp. 1 X XX Corinnidae Genus I, sp. 1 XX XX XX X Corinnidae Genus 1, sp. 2 XX X Corinnidae Genus 2, sp. 1 X X Corinnidae Genus 3, sp. 1 X XX XX Corinnidae Genus 4, sp. 1 X X XX XX Corinnidae Genus 5, sp. 1 X Corinnidae Genus 6, sp. 1 X X Cyatholipidae Mati/do sp. 1 X XX Cyatholipidae Matilda sp. 2 X Cyatholipidae Matilda sp. 3 X Deinopidae Deinopis sp. 1 XXX XX Desidae Badllmna insi/',nis (fhorell) X X X Desidae Genus 1, sp. 1 X XXXX XX Desidae Genus 2, sp. 1 XX X Desidae Genus 3, sp. 1 X Desidae Genus 4, sp. 1 X Desidae Genus 5, sp. 1 X X Dictynidae Genus 1, sp. 1 X X Dictynidae Genus 2, sp. 1 X XX XX XX X Dictynidae Genus 3, sp. 1 X X X Filistatidae If/ande//a sp. 1 XX XX Filistatidae Wande//a sp. 2 X X X XXX Filistatidae Wand,//a sp. 3 XXX X Filistatidae If/and,//a sp. 4 X X

Table 2 (cant.)

Familv Genus / species BB BO CV EL GT KE MD MR NA NE PE WO ZV 1 234 5 1 234 5 1 234 5 6 1 2 1 2 3 4 5 1 234 5 1 2 3 4 5 1 234 5 1 234 5 1 234 5 1 2 3 4 5 1 234 5 1 234 5 Lycosidae Lyrosa sp. 06 XXXX X Lycosidae I.,yrosa sp. 07 XXX X X X XXXX XXX XXXX Lycosidae Lyrosa sp. 08 X XXX X X XXX X XXX XX Lycosidae I.,yrosa sp. 09 X XXXX X XX X Lycosidae I.,yrosa sp. 10 XX X XX X X XX XXX XXXXX Lycostdae Lyro,a sp. 11 X X X XXX X XXX XXXX XXX XX X X Lycosidae Lyrosa sp. 12 XX X X XXXX XX X XXXXX Lycosidae Lyrosa sp. 13 XX XXX X X X X X X X XX XXXX Lycosidae Lyrosa sp. 14 X X X XXX X X XXXX X XXX X XXX X Lycosidae Lyrosa sp. 15 X XXX XX Lycosidae Lyrosa sp. 16 X X X X X XXX Lycosidae Lyrosa sp. 17 XXX XX XX X X XX XXX XX X X Lycosidae Lyrosa sp. 18 XXX XXX XX X XX Lycosidae Lyrosa sp. 19 XX X XX X Lycosidae I.,yrosa sp. 20 XX XXX X XX XX X X Lycosidae I.,yrosa sp. 21 X XX X XX Lycosidae I.,yrosa sp. 22 X XXX X X Lycosidae Lyrosa sp. 23 X XX Lycosidae I.,yrosa sp. 24 X Lycosidae I.,yrosa sp. 25 X Micropholcommatidae Genus 1, sp. 1 X Genus 1, sp. 1 X XXX X XXX X XXX Miturgidae Genus 1, sp. 2 X XX XXXXX XXX XXX Miturgidae Genus 2, sp. 1 XXXX XXXX XXX X X XXX X Miturgidae Genus 2, sp. 2 X X XXX X XX XX XXXX X X X X Miturgidae Genus 2, sp. 3 XX XX XX Miturgidae Genus 2, sp. 4 XX X XXX Miturgidae Genus 2, sp. 5 XX Miturgidae Genus 2, sp. 6 X XXXXX XXXXX X XX XXXXXX XXXX XXXX Miturgidae Genus 3, sp. 1 XX XXX X XXX XX XX XXXXX Miturgidae Genus 4, sp. 1 X Miturgidae Milllrxa aJ!.eknina (Simon) XX XX X Miturgidae Milllrxa sp. 1 XXX XX XXX XXX XXX XXX XX XX X X XX Miturgidae Milllrxa sp. 2 X XXXX X XXXXX XX XXXX XXX XX XX XXXX X Miturgidae Milllrxa sp. 3 X XXX X Miturgidae Milll'l,a sp.4 X Miturgidae Vfiodon lamnlllfinlls (L. Koch) X X X X XXXX Nirodamlls mainae Harvey X XXX X X XXX X X X sp. 1 XX XX XX X X X XX Oonopidae Gamasomorpha sp. 2 XXX X X Oonopidae Genus 1, sp. 1 XXXXX X XX Oonopidae Gtymells sp. 1 XX XXX X X XX XXXXX XX XXX XXX X Oonopidae Gtymells sp. 2 X X XX Oonopidae Gtymells sp. 3 X XX Oonopidae Gtymells sp. 4 X X X XX Oonopidae Grym'lIs sp. 5 x XX X X X Oonopidae Grym'lIs sp. 6 xx X X XXXX XX X XX Oonopidae Myrmopopa,a spp. XXXXXXXXXX XXXX XX XX XX XX XXXXXXXXXXXXXXXXXX XXX X XXXXXXXX Oonopidae Opopa,a sp. 1 XXXX X X XXXXXXXXX XXX X X X X X XX Oonopidae Opopa,a sp. 2 X X X X X XX XX X X X X XXX X X X XXX Oonopidae Opopa,a sp. 3 X XXX X XX XXX X X XX X Oonopidae Opopa,a sp. 4 X X X X X X Oonopidae Orrheslina sp. 1 X 0xyopldae O>g'opes sp. 1 XX XX X X XXXX X X XX XX XXXXX XXXX XX XXX XX Oxyopidae O>g'opes sp. 2 XXXXX XXXXX X X X X X X XXX X XXXXX XXX X X XX XXXXX X XX TrichorYc"'s sp. 1 X X XXXX XX XXXXXXX X XXXXXXXXX X XX X XXXXX X Pholcidae TrichorYe/lIs sp. 2 X Prodldomidae Cryplo,rilhlls sp. 1 XX X X X X X Prodtdomidae Cryplo,rilhlls sp. 2 XXX X XX X X Prodidomidae Cryplo,rilhlls sp. 3 XXXXX XX XX XX XX X XX XXX X X XX X Prodidomidae Cryplo,rilhlls sp. 4 X XXX X X X XXXXX X X X XX X X XX XXX X Prodtdomidae Cryplo,rilhlls sp. 5 XX X X X Prodidomidae Cryplo,rilhlls sp. 6 X X Prodidomidae Molycriinae Genus 1, sp. 1 X XXXXX XXXX X XX XXX X XXXXX XXX XXXXXXX X Prodidomidae Molycriinae Genus 1, sp. 2 XX xx XX XX XX X X Prodidomidae Molycriinae Genus 1, sp. 3 X X X XX X Prodidomidae Molycriinae Genus 2, sp. 1 XXX XXX XXXXXXXXXXX XXXXXXXX XXX XXXXX XXXXXXXX Prodtdomidae Molycriinae Genus 2, sp. 2 XXXX Prodldomidae Molycriinae Genus 2, sp. 3 X X X X Prodidomidae Molycriinae Genus 3, sp. 1 X X X XX X XXX XX X X X XXX Prodidomidae Molycriinae Genus 4, sp. 1 X XX XXXX Prodidomidae Prodidominae Genus 1, sp. 1 X X X Prodidomidae Prodidominae Genus 2, sp. 1 X X X Prodidomidae Prodidominae Genus 2, sp. 2 X Prodidomidae Prodidomlls sp. 1 X XXXXX X X XXX XXX XX X XX XX X X Prodidomidae Prodidomlls sp. 2 X Prodidomidae Prodidomlls sp. 3 X Salticidae Bianor sp. 1 X X XX XX X X X Salticidae Bianor sp. 2 X XXX XXXXX X XX XXXX XX X X X X Salticidae qynolis a/bobaroallls (L. Koch) XX XX X Salticidae G./ynolis sp. 1 X Salbcidae Cyla,a sp.1 X XX Salticidae GanJ!.lIs sp. 1 X Salticidae GanJ!.lIs sp. 2 X Salbcidae Genus 01, sp. 1 X Salbcidae Grqy

Familv Genus / sDecies BB BO CU EL GT KE MD MR NA NE PE WO ZU 1 2 3 4 5 1 234 5 1 234 5 6 1 2 1 234 5 1 234 5 1 2 3 4 5 1 234 5 1 2 3 4 5 1 234 5 12345123451 234 5 Salticidae Lyddas sp. 3 X XX X X XX Salticidae Lyddas sp.4 X Salticidae Lyddas sp. 5 X XX Salticidae Marallls vesper/ilio (Simon) X X Salticidae MaT}',aromma sp. 1 X XXX X Salticidae MaT}',aromma sp. 2 X Salticidae MaT}',aromma sp. 3 X Salticidae !l!yrmaracbne sp. 1 X X Salticidae Ocrisiona Iellcocomir (L. Koch) X X X X XXX X X XX X Salticidae OpislbonCIIs sp. 1 X X XXX X XX Salticidae OpislbonCIIs sp. 2 XX XX Salticidae Paraploloides sp. 1 X XX XX XX X Salticidae Paraploloides sp. 2 X Salticidae Simaelba sp. 1 X Salticidae Simaelblllo sp. 1 XX Salticidae Unidentati, Genus 01, sp. 1 XX XXX XX XXX X X Salticidae Unidentati, Genus 02, sp. 1 XXX XXX X XX X XX X Salticidae Unidentati, Genus 02, sp. 2 X Salticidae Unidentati, Genus 03, sp. 1 X X X X X Salticidae Unidentati, Genus 04, sp. 1 X XXX XXXX XX XX XX X X Salticidae Unidentati, Genus OS, sp. 1 XX XXX XX XX XX X X XXX XX XX X X XXX Salticidae Unidentati, Genus OS, sp. 2 X X X X Salticidae Unidentati, Genus 06, sp. 1 X Salticidae Unidentati, Genus 07, sp. 1 X X Salticidae Unidentati, Genus 08, sp. 1 X Salticidae Unidentati, Genus 09, sp. 1 X XX X XX Salticidae Unidentati, Genus 10, sp. 1 X Salticidae Unidentati, Genus 11, sp. 1 XX Salticidae Unidentati, Genus 12, sp. 1 X X X X Salticidae Unidentati, Genus 13, sp. 1 X X Salticidae Unidentati, Genus 14, sp. 1 X X Salticidae Unidentati, Genus IS, sp. 1 X Salticidae Unidentati, Genus 16, sp. 1 X X XX X Salticidae Unidentati, Genus 16, sp. 2 X X Salticidae Unidentati, Genus 17, sp. 1 X X Salticidae Unidentati, Genus 18, sp. 1 X Salticidae Zebraplo!ysfraclivillala (Simon) X Salticidae Zebraplolys kryserlinl'.i Zabka XXXX XXX XX XXX XX XX XX XX XX XX X XX XXXXX X Salticidae Zebraplo!ys sp. 3 X X Salticidae Zebraplo!ys sp. 4 X Salticidae Zenodoms sp. 1 X Segestriidae Ariadna sp. 1 X X X XX Segestriidae Ariadna sp. 2 XXX Segestriidae Ariadna sp. 3 X X X X Segestriidae Ariadna sp. 4 XX X X XX XX XX X Segestriidae Ariadna sp. 5 X X XX X X ~~ X Segestriidae ATiodno sp. 6 X X ;l> xx XXX ... Segestriidae ATiodno sp. 7 X X XXXXX XXXXX x x XXXXX XXXXX XX XXX X X ll> ::l Selenops sp. 1 X rll 0 Comsoides sp. 1 X X X XXXXXX X X XXXXX XX X S Stiphidiidae ForsleTino sp. 1 XX X XX XX XX XX XX ...0 ForsleTino sp. 2 XX X XX XX '"t:l Stiphidiidae =:r' XX X XX XX XXXX Stiphidiidae ForsleTino sp. 3 X X Ul Stiphidiidae Genus 1, sp. 1 X X X X '"t:l.... XX X t:l,. Stiphidiidae Genus 2, sp. 1 ...rll Stiphidiidae Genus 3, sp. 1 XXX '" Stiphidiidae Genus 4, sp. 1 X Stiphidiidae Genus 5, sp. 1 X Stiphidiidae Genus 6, sp. 1 X Tetragnathidae Genus 1, sp. 1 X X Tetragnathidae Nepbila edlllis (Labillardiere) X Enoploi',nolho sp. 1 X X Theridiidae Ellryopir sp. 1 X X XX X X XX X X X X Theridiidae EllIyopis sp. 2 X X X XXX X X X Theridiidae Genus 1, sp. 1 X X X X X X X TheridiIdae Genus 2, sp. 1 X Theridiidae Genus 3, sp. 1 X X Theridiidae Gmoi',ola sp. 1 X X XX Theridiidae Gmoi',ola sp. 2 X X X Theridiidae Gmoi',ola sp. 3 XX TheridIidae Lalrodecllls hossellii Thorell X X X XX X X X Thendlidae Sleolorla sp. 1 X X X X XX X X X XX XX XX X Theridiidae Sleolorla sp. 2 X X X X X X Thendiidae Sleolorla sp. 3 X X X Theridiidae Sleolorla sp. 4 X X X X Theridiidae TTii',OI1obolhrys sp. 1 X Thomlsidae SlepIJonopi.r sp. 1 X Thomisidac SIphonopis sp. 2 X SIphonopis sp. 3 X X X Thomisidae Slphonopis sp. 4 X Thomisidae Slphonopis sp. 5 X X X X Thomisidae Slphonopis sp. 6 X Thomisidae Slephonopis sp. 7 X XX Thomisidae Thorf!yno sp. 1 X X XXX Thomisidae Thorpyno sp. 2 X X X X X X X X XXX Thomisidae Thorpyno sp. 3 X Trochanteriidae Genus 1, sp. 1 X XX Trochanteriidae Genus 1, sp. 2 XX X X X Trochanteriidae Genus 2, sp. 1 X X XX X X X XX XX X X XX XX Trochanteriidae Genus 3, sp. 1 X XX X XX X X XX XX Trochanteriidae Genus 4, sp. 1 XX X X X X X X X X Trochanteriidae Genus 5, sp. 1 X Trochantenidae Genus 6, sp. 1 X Zondae A1)!,oCleI1llS sp. 1 XXXXX XXX XXXXXXXX XXXXX XXXX XX XX XXXXX X XX XX XXXX XXXXX XXXXX Cl) Zoridae Heslimodemo sp. 1 X 0 Zondae Tho.rvrfJeo so. 1 XX "l 308 M.S. Harvey, A. Sampey, P.L.J. West, J.M. Waldock found at more than three quadrats. Dicytnids occur However, not all records have yet been collated in various habitats, including on the ground and in from this revision, and we are able to present data foliage. on only two species. Lampona cylindrata (1. Koch), which is widespread throughout southern Filistatidae Australia, was found at four quadrats, and The Filistatidae were represented by five Pseudolampona boree Platnick, which was found at undescribed species of Wandella, many of which PE5 and ZU3. Elsewhere this species is found in were taken from several survey areas widely other parts of southern Australia (Platnick, 2000). scattered across the study area. A recent revision of the family (Gray, 1994) showed that the Australian Filistatidae are taxonomically diverse at the species Four species of Linyphiidae were taken, including level. Filistatids often occur under bark, but many a species of Erigone. However, each species was roam across the ground in search of prey and mates. represented by very few specimens and due to their habit of spinning small sheet-webs, may be under- represented in the survey. Two species of this highly restricted and little- known family were recorded from the study area, Genus I, sp. 1 at KE3 and Genus I, sp. 2 at BB3 and Three species of were each collected at CUI. Gallieniellids are hunting spiders found close a single quadrat (ZU3, MR5 and NE4). These small to the ground. spiders are generally found in leaf litter.

Gnaphosidae Lycosidae Preliminary sorting indicated that the Numerous members of the ground-hunting Gnaphosidae represented one of the most diverse family Lycosidae, wolf spiders, were frequently spider families in the survey. Time constraints recorded during the survey, and 35 species were precluded species-level identifications, apart from detected, of which three could be assigned to members of the Hemicloeinae for which five species previously described species. Lycosa bicolor Hogg of Hemicloea were collected from very few localities, was recorded from 11 quadrats, which extends the possibly reflecting their corticolous nature. Other known range of the species further to the west gnaphosids are also bark-dwelling, but many others (McKay, 1973). Lycosa forresti McKay was collected are found at ground-level under leaf litter and at two quadrats in the southern portion of the amongst rocks. survey area (ELl and NA4), which is in accordance with the known distribution of the Hersiliidae species, which is generally to the south-east of the Hersiliids are tree-dwelling spiders, which rarely study area (McKay, 1973). Lycosa mainae McKay fall into pitfall traps. Tamopsis occidentalis Baehr and was collected at two quadrats on Peron Peninsula Baehr was recorded from B05, while Tamopsis sp. 1 (PE2 and PE4), extending the known range of the and Tamopsis sp. 2 were taken at one (GJ2) and two species further to the north-west (McKay, 1979). (MD5, MR2) quadrats, respectively. However, the distribution of L. mainae was found to be greater, as specimens were hand-collected Heteropodidae from burrows at Nerren Nerren Station, even Fourteen species of Heteropodidae were though no specimens were collected by pitfall collected, including single species of Delena, trapping in the area. , Holconia, Isopodella and Pediana, of which Genus I, sp. 1 and Lycosa sp. 1 were recorded the most abundant were Heteropoda kalbarri Todd mainly from coastal localities (BB, CU, EL, PE and Davies (9 quadrats) and Pediana tenuis Hogg (7 ZU). The two species of Genus 2 were quite quadrats) which have been previously recorded widespread, whereas the three species attributed to from the study area or from nearby areas (Davies, Genera 3, 4 and 5 were quite localised. The 1994; Hirst, 1989). Nine species of Neosparassus were remaining species of Lycosa were variably collected, but most were from just a few quadrats. distributed, with many species widespread ;Heteropodids usually dwell amongst foliage or throughout the study area, and others more under tree bark, but many move across the ground restricted. to find new habitat. Micropholcommatidae Lamponidae A single specimen of the Gondwanan family The ground and bark dwelling members of the Micropholcommatidae was collected at ZU3, Lamponidae were well represented in the survey, probably representing the northern limit of the with 15 species being recorded by Platnick (2000). family in Western Australia (M: Harvey, Araneomorph Spiders 309 unpublished data). These tiny litter-dwelling restricted to KE3. These spiders spin undifferen- spiders are rarely found in pitfall traps. tiated webs, but males often wander in search of females during mating season. Miturgidae Sixteen species in six genera were collected, of Prodidomidae which the most abundant were Genus I, spp. 1and The Prodidomidae were found to be particularly 2, and Genus 2, sp. 6, and two species of Miturga diverse in the study area, with 20 species in eight (spp. 1and 2). Uliodon tarantulinus (L. Koch), which genera collected. Prodidomids are ground-hunting is widespread across Australia (R. Raven, in litt.), spiders and some, such as the molycriines, was found at eight quadrats. Miturgids spin silken specialise by preying on . retreats under rocks and logs, but venture out to The Prodidominae consisted of three species of hunt and mate. Prodidomus and three species belonging to two different, possibly undescribed, genera. Prodidomus sp. 1was found to be very widespread, while P. sp. Nicodamidae 2, P. sp. 3and Genus 2, sp. 2were only found .at The sole species of Nicodamidae previously single quadrats at GJl, B03 and ZU3, respectively. known from the study area, Nicodamus mainae Genus I, sp. 1and Genus 2, sp. 1were found at Harvey, was taken from 12 quadrats from atotal of three quadrats across mutually exclusive survey 6survey areas (BB, BO, KE, NA, NE and PE) areas. scattered throughout the study area. Nicodamus The Molycriinae were represented by eight mainae is widespread throughout the southern half species, of which the three species included in of Western Australia, occupying awide variety of Genus 1were significantly larger than the others. dry habitats (Harvey, 1995). Nicodamids spin Genus 2, sp. 1was extremely widespread, and was undifferentiated webs under rocks and logs, but found at all survey areas with the exception of CU males wander in search of females during the and W. In contrast, Genus 2, sp. 2was only found mating season, often during daylight hours. at CU, and Genus 2, sp. 3only at BO. Genus 3, sp. 1 was also widespread, but Genus 4, sp. 1was Oonopidae restricted to BO and GJ. The tiny litter-dwelling spiders which constitute Six species of the little-known genus Cryptoerithus the family Oonopidae were well represented in the were recognized, of which most were relatively survey, and 15 species in six genera were collected, widespread. The subfamilial placement of this including two species of Gamasomorpha, asingle genus is currently uncertain (V. Ovtsharenko, species of , and four species of . personal communication). The genus Myrmopopaea was extremely common and widespread, and despite the presence of Salticidae morphological variation between specimens, it was The Salticidae represented the most diverse not possible to satisfactorily assign species-level family currently analysed from the survey with 58 rank to these variants. Therefore, for the purposes species placed in 39 genera. They are found in a of this study, we recognise only asingle species of variety of terrestrial habitats, including under bark, Myrmopopaea. The genus Grymeus was found to under rocks, amongst leaf litter or on low consist of six species of which G. sp. 1and G. sp. 6 vegetation. were the most widespread. A single species of an Three species of Grayenulla ~ recorded, of undescribed genus (Genus I, sp. 1) was recorded at which Grayenulla australensis Zabka, previously eight quadrats. known from the Goldfields region of Western Australia (Zabka, 1992a), was found to be Oxyopidae widespread throughout the survey area, absent Distinguishing between the various species of only from Edel Land. Two undescribed species of Oxyopes found in the survey was difficult, and we Grayenulla were collected at only one or two were only able to confidently identify two species. northern survey areas. The remaining specimens have not been identified Two species of were collected, H. to species level, and were excluded from the planissima sp. group at KE, PE and WO, and the analysis. Oxyopids are common inhabitants of other, H. sp. 2, at asingle southern quadrat (ZU). grasses and low shrub land, where they will sit and Holoplatys is an extremely diverse genus which is attempt to catch prey. widespread in Australia, New Caledonia and New Zealand (Zabka, 1991). Pholcidae vespertilio (Simon) was found at two The Pholcidae were represented by two species of quadrats on BO and NE; it has also been found in Trichocyclus, one of which (T. sp. 1) was found at other parts of southern Australia (J. Waldock, numerous quadrats, while the other (T. sp. 2) was unpublished data). 310 M.S. Harvey, A. Sampey, P.L.]. West, ].M. Waldock

Ocrisiona leucocomis (L.Koch) was found at 12 Australia, New Caledonia, New Guinea and New quadrats, representing the first specimens recorded Zealand (M. Harvey, unpublished data). Similarly, north of Perth. The species is widespread the only other tetragnathid collected, Genus 1, sp. 1, throughout eastern and southern Australia (Zabka, was represented by a single male from W05. 1990). Two species of the genus Paraplatoides were Theridiidae recorded, one at eight quadrats (P. sp. 1) and the Fifteen species of Theridiidae attributable to 10 other at one quadrat (P. sp. 2). This is the first genera were collected, including four species of record of the genus from outside eastern Australia (16, 6, 3 and 4 quadrats, respectively), (Zabka, 1992a). hasseltii Thorell (8 quadrats), a single The endemic Australian genus Zebraplatys was species of (2 quadrats, CUI, PE1) and represented by four species, Z. fractivittata (Simon), single species representing three unidentified Z. keyserlingi Zabka and two undescribed species. genera (7, 1 and 2 quadrats, respectively). The Zebraplatys fractivittata was collected once at ZU5, Hadrotarsinae were represented by two species of extending its known distribution further north (12 and 9 quadrats, respectively), three (Zabka, 1992b), while Z. keyserlingi, previously species of Gmogala (4, 3 and 2 quadrats, known from only the male holotype from respectively) and a single species of Trigonobothrys Woodstock Station in the northern Pilbara (Zabka, (1 quadrat, NE4). As part of a recent revision of the 1992b), was found to be widespread throughout the genus Gmogala (M. Harvey and J. Waldock, unpUbl- survey area at 33 quadrats. Two undescribed ished data), Gmogala sp. 1 was found throughout species of Zebraplatys were found at NA1, NE4 (Z. Western Australia and South Australia, Gmogala sp. sp. 3) and ZU3 (Z. sp. 4). 2 was not found outside of the study area, and Little can be said of the remaining salticid species, Gmogala sp. 3 was found to be extremely presumably undescribed, detected in the survey, Widespread throughout southern Australia. but many taxa were recorded from few quadrats, Theridiids generally spin tangle webs and are not indicating that they are either rarely collected in particularly amenable to capture in pitfall traps, but pitfall traps or that there is significant local the hadrotarsines occur in leaf litter and at the bases endemism. of low vegetation, thus making them more susceptible to capture. Segestriidae Seven species of Ariadna were discerned, of which Thomisidae A. sp. 7 was the most abundant, being found at 36 Only two genera of Thomidae were recorded: quadrats. Segestriids dig burrows into the soil, into Stephanopis with seven species and Tharpyna with tree bark, or between rock fissures, but males roam three species. Both genera are generally found in search of females during the mating season. under the bark of trees, particularly eucalypts.

Selenopidae Trochanteriidae Only a single species of Selenops was taken at BB3. Seven species of the Gondwanan family It is unlikely that pitfall trapping would adequately Trochanteriidae were collected, mostly represented sample for selenopids, as they are mostly restricted by adult males presumably moving about on the to the underside of stones and rocks, and are rarely ground in search of mates. found outside of this habitat. Zodariidae Stiphidiidae Preliminary sorting indicated that the Zodariidae A single species of Corasoides was taken at three were extremely diverse in the study area, but time quadrats (NA1, NE5 and PE3), and three species of constraints precluded species-level identifications. Forsterina of which F. sp. 1 was Widespread across the study area. Six species, placed in putatively Zoridae separate genera, were collected at a few quadrats. Three zorids were found in the study area: single Stiphidiids spin webs, ranging from poorly species of Argoctenus, Hestimodema and Thasyraea. differentiated structures in Forsterina to elaborate The latter two genera were very rarely collected, 'sheet webs in Corasoides. whilst Argoctenus sp. 1 was found at most survey areas. Zorids are ground-hunting spiders. Tetragnathidae A single female specimen of Nephila edulis Species Richness and Assemblage Composition (Labillardiere) was collected in a pitfall trap at NEl. We identified 285 species across the 63 quadrats, These web-building spiders are abundant ranging from 12 to 56 species per quadrat. Species throughout the study area, as well as elsewhere in richness was generally poorest on quadrats situated Araneomorph Spiders 311 on saline claypans (PE1 with 12 species, CUI with type. Analysis confirmed significant differences in 14 species and NA1 with 28 species), but at BB3, various climatic and substrate ~ 'annual another saline claypan, 44 species were recorded. average precipitation' (Pann), 'soil potassium' Other depauperate quadrats included the two Edel [K(HC03)], 'soil exchangeable magnesium' (exMg) Land quadrats which were situated on fine and 'exchangeable sodium' (exNa) provided the calcareous dunes, with 14 and 16 species, best statistical separation of the five groups defined respectively. The five richest quadrats were B03-5 in Figure 2(Figure 3), although Pann and K(HC03) and PE2-3 with 49-56 species each, all situated in were tightly inter-correlated (R2 =-0.43, P<0.0001), Acacia shrub land. The overall species richness was as were exMg and exNa (R2 = 0.52, P<0.0001). little affected by regular grazing by introduced When the dendrogram clusters at the 17-group level herbivores (e.g. stock, feral goats and/or rabbits), were examined, we noted that many of the groups and even the quadrats with obvious sheet erosion, comprised quadrats from asingle survey area minimal leaf litter, and/ or no Al soil-horizon, (Table 3). In contrast, the structure of the retained high species richness [e.g. B05 (52 species), dendra gram derived from the species classification GJ4-5 (25, 35), KE4 (36), MDl-2, 4-5 (22, 20, 33, 34), was the only basis used to partition the 285 species PE4 (43) and W01 (33)]. in the data matrix into 30 assemblages; unfort- In Table 3 the data have been re-ordered unately, detailed ecological data on the ground- according to the quadrat and species classification dwelling spiders of the region are too scant to analyses. provide an extrinsic rationale for this partitioning When the 63 quadrats were classified according decision. to similarities in their species composition (Figure Three distinct types of species assemblage were 2), five dendrogram partitions were recognisable in apparent in Table 3. Two were related to ecological terms of their geographical location and substrate or biogeographical gradients, and one to the

Group Levels BB1,3 1 1 BOl-5 J.I J.I 1. Central and northern GJl-5 1 1 areas (non-coastal) as KE3-5 \ 1 well as saline claypans, MDl-5 I I except for PE3-4 and NA2. MRl-5 1 1 NAl-2 I I PE3-4 1 1 WOl-5 -1-1--.- BB2,4,5 21 21 2. Central coastal sands CU2-6 -1-1- ELl-2 _31_ I 3. Edelland dunes KEl-2 I I PE2,5 4\ I 4. Southern sands, NEl-5 I 1 except for KEl-2 NA3-5 I I ZUl-5 -1-1 CU1 51 41 l5. Saline claypans high in gypsum PE1 -, 1-1--\ ---I 0.8 1.0 1.2

Coefficient of dis-similarity

Figure 2 Quadrats classified according to similarities in their species composition. Dendrogram structure is displayed to the 8-group level, and cut at the 4- and 5-group levels. 312 M.S. Harvey, A. Sampey, P.L.]. West, ].M. Waldock strongly localised patterns of occurrence in their To reduce the influence of localised 'endemism' component species. on the analysis outcome, assemblages 5 to 27 and 30 1. Species assemblages 1 to 4 comprised species were removed from the data-set. When the reduced that were widespread in the study area; matrix was re-analysed, the allocation of quadrats 2. Assemblages 5 to 27 and assemblage 30 among dendrogram partitions (Figure 4) were more comprised species with strongly localised geographically consistent than the allocation distributions in the study area. Their component derived using the entire data-matrix (Figure 2): KEl species were almost always confined to one or and KE2 were allocated with the remainder of the two survey areas. Where they occurred at two central and northern quadrats, while PE3, PE4 and or more survey areas, these were generally NA2 were allocated with the rest of the southern adjacent. The scattered singleton exceptions quadrats. may indicate taxonomic problems which could 'Precipitation in the coldest Quarter' (PcldQ) in not be resolved with the available collections; conjunction with 'soil exchangeable Sodium' (exNa) and provided the best statistical separation of the five 3. Assemblages 28 and 29 occurred on saline groups defined in Figure 4 (see Figure 5), although claypans at widely separated locations. the gradient in 'soil carbon' conformed most closely

Annual Average Precipitation (mm) (Kruskal-Wallis H = 35.7 df= 4 P = 0.0000)

200 230 261 291 315 GRP N +------+------+------+------+ 1 34 ======M======2 8 ======M======3 2 =M== 4 17 ======M======5 2 ===M==

Soil Potassium (ppm) (Kruskal-Wallis H = 41.8 df= 3 P = 0.0000)

12 174 330 500 660 GRP N +------+------+------+------+ 1 34 ======M======2 8 ====M=== 3 19 ==M== 4 2 ======M======

Soil Exchangeable Magnesium (me%) (Kruskal-Wallis H = 23.3 df= 3 P = 0.0000)

0.04 1.2 2.4 3.6 4.8 GRP N +------+------+------+------+ 1 34 ======M======2 8 ===M=== 3 19 ==M= 4 2 ======M=====

Soil Exchangeable Sodium (me%) (Kruskal-Wallis H = 11.8 df= 3 P = 0.0081)

0.01 7.3 14.6 21.8 29.2 GRP N +------+------+------+------+ 1 34 M= 2 8 M= 3 19 M 4 2 ======M======Figure 3 Average environmental attribute values for the quadrat groups defined in Figure 2. Bars indicate standard deviations about the mean (M); GRP = classification group number in Figure 2; N =number of quadrats in each classification group. Araneomorph Spiders 313

BB1,3 1 BOl-5 1 GJl-5 1 KEl-5 11 1. Central and northern MDl-5 I areas (non-coastal) as MRl-5 I well as saline claypans. NA1 1 WOl-5 -1------BB2,4,5 12 2. Central coastal sands CU2-6 _1_ NA2-5 1 NEl-5 13 3. Southern sands PE2-5 I ZU1-5 -I-I---...L..------r CU1 14 4. Saline claypans high in gypsum PE1 _1 _ ELl-2 _15 1 _ 5. Edel Land dunes IIII 0.89 1.01 1.19

Coefficient ofdis-similarity

Figure 4 Quadrats classified according to similarities in their species composition, excluding species assemblages-5 to -27 and assemblage-3D (the strongly localised species; see text). Dendrogram structure is displayed to the 5- group level.

with dendrogram structure at this level. Although The two species of Miturgidae Genus 1 were well PcldQ and exNa were not inter-correlated (R2 = 0.02, represented amongst the samples and were P = NS), 'soil potassium' (K(HC03» was tightly completely allopatric, the former in five northern inter-correlated with PcldQ (R2 = -0.36, P < 0.001), survey areas (Ba, KE, GJ, BB and MD) and the and 'soil exchangeable Magnesium' (exMg) with latter in four southern survey areas (PE, WO, NA exNa (R2 = 0.52, P < 0.0001). Since PcldQ and and NE). No specimens of this genus were collected 'annual average precipitation' (Pann) are also at CU, MR, EL or ZU. tightly inter-correlated (R2 = 0.71, P < 0.0001), these The six species of Miturgidae Genus 2 were found results are consistent with the physical attributes in varying regions of the study area. Genus 2, sp. 1 identified using the entire data-set (described was disjunctly distributed in the north (CU and MR) above). and the south (WO, NA, NE and ZU). Genus 2, sp. 2 and sp. 6 were widespread throughout the survey Vicariant Patterns areas north of the mulga-eucalypt line, with the Spiders have been the focus of many broad-scale most southerly occurrence of sp. 2 at NA2 and that biogeographic studies, some of which have of sp. 6 at NAl, which are situated on the boundary demonstrated significant vicariant distribution between the Southwest and the Eremaean Botanical patterns, especially at the continental scale (e.g. Provinces. Conversely, Genus 2, sp. 3 was found at Platnick, 1976). The data presented here was two near-coastal survey areas within the Southwest scrutinised to determine whether any vicariant Botanical Province at EL and ZU. Genus 2, sp. 4 patterns were evident between two or more species and 5 were restricted to northern areas, with the within individual genera. former at CU and MR, and the latter at Ba. The two Carnarvon species of the clubionid Meedo Species of Miturga also possessed varying were found to be disjunctly distributed with M. distribution patterns. Miturga sp. 2 was widespread houstoni found in the central and northern quadrats in all survey areas except GJ, MD and EL. Miturga (BB, Ba, GJ, KE, MD, MR, NA and WO) with the sp. 3 was patchily distributed in MR, KE and BB, most southerly record at NAl, and M. sp. 1 only in while M. agelenina (Simon) was found to be disjunct the southern quadrats at EL and ZU. This pattern at BB and ZU. Miturga sp. 1 was found in all survey follows the division between the Southwest and the areas of the Eremaean Botanical Province except Eremaean Botanical Provinces. CU, with the most southerly occurrence at NA2 314 M.S. Harvey, A. Sampey, P.L.J. West, J.M. Waldock

Coldest Quarter Precipitation (mm) (Kruskal-Wallis H = 47.4 df = 4 P = 0.0000) 69 101 133 165 180 GRP N +------+------+------+------+- 1 33 ======M======2 8 =M= 3 18 ======M======4 2 ======M======5 2 =M=

SoilExchangeableSodium(me%) (Kruskal-WallisH= 10.7 df= 4p=O.03) 0.01 7 15 22 29 GRP N +------+------+------+------+ 1 33 M= 2 8 M= 3 18 M 4 2 ======M======5 2 M

Soil Exchangeable Magnesium (me%) (Kruskal-Wallis H = 21.0 df = 4 P = 0.0004) 0.04 1.2 2.4 3.6 4.8 GRP N +------+------+------+------+ 1 33 ======M======2 8 ===M=== 3 18 ==M= 4 2 =====M===== 5 2 =M=

Soil Organic Carbon (ppm) (Kruskal-Wallis H = 30.7 df = 4 P = 0.0000)

0.09 0.28 0.48 0.67 0.87 GRP N +------+------+------+------+ 1 33 ======M======2 8 ======M======3 18 ======M======4 2 ======M======5 2 ===M==

Figure 5 Average environmental attribute values for the quadrat groups defined in Figure 4. Bars indicate standard deviations about the mean (M); GRP =classification group number in Figure 4; N =number of quadrats in each classification group.

which is situated on the boundary of the Southwest These patterns ~ that varying historical and the Eremaean Botanical Provinces. Miturga sp. factors have probably played asignificant role in 4was only found at ZUl within the Southwest spider speciation patterning across the southern Botanical Province and may represent amember of Carnarvon Basin, with the divergence between amore southerly distributed taxon. southern and northern species the most obvious Molycriinae Genus 2(Prodidomidae) contained pattern. Less frequently observed was awest-east three species of which Genus 2, sp. 2was found vicariant pattern, with the Prodidomus species the only at the northerly CU, demonstrating total most obvious example. allopatry with Genus 2, sp. 1which was widespread at all survey areas except CU and ZU. The third ~ Genus 2, sp. 3, was less informative, being DISCUSSION found sporadically at Ba, PE, NA and ZU. Of the 53 araneomorph spider families known to The three species of Prodidomus possess varying possess indigenous representatives in Western distributions with P. sp. 1Widespread across all Australia, eight are restricted to the temperate survey areas except GJ, PE, NA and EL, and the south-western corner (, , allopatric P. sp. 2found only at GJ, the most Cycloctenidae, , Mimetidae, Orsolob- westerly site examined as part of the survey. The idae, Pararchaeidae and Toxopidae), while three are third species is sympatric with P. sp. 1at B03. known only from the northern tropiCal regions Araneomorph Spiders 315

(, Scytodidae, Tetrablemmidae -an of data on the ecological niches of most Carnarvon introduced scytodid species has been found in Basin araneomorph spiders, the ~ south-western Australia). Others, such as the correlates with their patterns of occurrence across , are patchily distributed in the study area that are identified in this paper Western Australia, occurring either in forest should not be treated as causal. habitats in the temperate south-west (Harvey, 1992) It is clear that the most disparate quadrats, in or in caves of Cape Range peninsula (Harvey, 1998). terms of species composition, were situated at As expected, none of these 12 families were the southern end of the study area. This is recorded during the present survey of the semi-arid entirely consistent with the floral attributes of southern Carnarvon Basin. the region (Beard, 1980), and indicates that the Of the remaining 41 families which are thought araneomorph spider assemblages of this region to be relatively widely distributed in Western shows marked differences to that in the Eremean Australia, 33 were recorded in pitfall traps during Botanical Province to the north. More work is the current survey (Table 2). Pitfall traps sample needed to analyse the spider fauna in relation to those portions of the fauna that are active on the areas to the south of the study area, but it. is ground surface, and the data presented here are likely that assemblages detected in the southern not designed to represent acomplete survey of the quadrats will also be found further south of the araneomorph spiders of the region. As noted study area. above, representatives of many families which are As noted above, many pairs of species were known to occur in the survey area are under- found to possess allopatric distribution patterns, represented in the species recorded (Table 2). suggesting that they have evolved in situ due to These include members of the which either low mobility or in response to local soil are unlikely to be found in pitfall traps due to their and climatic variation. Examples include the web-building habit, and the Hahniidae which clubionid Meedo, the miturgids Miturga, Genus 1 probably do not venture far from their natural and Genus 2, and the prodidomids Prodidomus habitat of tree bark. However, pitfall traps contain and Genus 2. These strongly localised patterns of ahigh proportion of adult males which present the endemism were particularly noticeable in the most useful diagnostic features for species southern survey areas and in quadrats with separation, and alow proportion of juveniles extreme environmental conditions, such as the which are virtually unidentifiable due to the lack saline claypans and the scree slopes of the of sexual features. Kennedy Ranges. However, the wider impli- The wide variety of taxa identified to date (285 cations of these observed trends in disjunct species placed in 146 genera) does not include two distributions and local endemism cannot be fully major families, Zodariidae and Gnaphosidae, for assessed without more detailed collecting of which numerous taxa exist in the survey collections. spiders in regions adjacent to the study area, and The addition of these taxa may well alter the results the completion of suitable taxonomic revisions of presented here. We estimate that the total diversity the taxa involved. of ground-dwelling araneomorph spiders for the The total patterning found in the assemblages survey region is likely to exceed 350 species, which (Table 3) were related to environmental gradients is nearly three times the diversity of the reptile as well as in situ evolutionary processes which can fauna of the region (McKenzie et al., 2000). These only be more fully assessed as the identity, data are far in excess of our initial estimates of the historical relationships and ecological niches of the fauna of the region, and highlight the relatively Australian spider fauna become more fully high diversity of terrestrial invertebrates in the understood. semi-arid and arid zones of Australia. Geographical patterns in the species composition Spiders in biogeographical surveys of araneomorph spider communities across the Spiders are adominant component of the study area were found to be correlated with of most terrestrial habitats (Coddington environmental gradients such as rainfall, modified and Levi, 1991) and are a useful tool in by aclaypan floor effect (indicated by either 'soil biogeographic and ecological studies (Churchill, exchangeable Magnesium' or salinity as 'soil 1996). They are easy to collect in astandardised exchangeable Sodium') (Figure 5). Magnesium fashion utilising pitfall traps and most of the accumulates in lake sediments, such as the saline specimens collected in this way are adults, which claypan quadrats that comprise Group-4. Together, can then be efficiently classified at the species level, these correlations point to productivity as ascalar despite the lack of taxonomic revisions and of spider assemblage composition, summarised by publications for many groups. The results presented the observed gradient in soil organic carbon (see herein suggest that araneomorph spiders can Figure 5) which is asurrogate for other variables provide afine-grained perspective of biodiversity such as vegetational cover. However, in the absence patterns across awide geographic area. 316 M.S. Harvey, A. Sampey, P.L.}. West, }.M. Waldock

Table 3 Entire data matrix reordered according to species co-occurrences at the same quadrats and similarities in the overall species composition of the quadrats. Quadrat codes are printed vertically, and dendrogram partitions at the 17-group level are coded'A-Q'.

Quadrat Codes Group A BC D EF GH I JKL M NO P Q BBBKKINIMMMIBBBBNPplGGGMGGMMWWWWWIMMMMIKIBBBlccccclEEIKKlpplNNNNININNNlzzzzzIcp BOBEEIAIDDDloOOOAEEIJJJRJJDDOOOOOIRRRRIEIBBBluuuuulLLIEEIEEIEEEEIEIAAAluuuuuluE 1234311112311345243113512445142351253415124512345611211212511345121345112453111 Assemblage-t Araneidae Argiope protensa * II I*I I *I* 1** I ** I*I*I 1* I I* I* Prodidomidae Molycriinae Genus I, sp. 2 ** II 1* I I **1*1 1* I 1**1 *1 II 1** Araneidae Dolophones sp. 1 ** 1*1 I** I I*II*** II*I*I*IIII Lycosidae Lycosa sp. 20 II *I** I 1* *II*** IIIIIIII Oonopidae Opopaea sp. 3 * II *I ** *1 1* * I 1 ** 1 *** 1 I I I 1 I I Prodidomidae Cryptoerithus sp. 2 I 1 1* *I 1* * 1 I ** I* I I I 1 1 I I Nicodamidae Nicodamus mainae II I**I 1 1 I ** I I I * I 1** I** II Salticidae Unidentati, Genus 04, sp. 1 I* I 1*** **I *III ** IIII*I ** I 1 * I Prodidomidae Cryptoerithus sp. 1 ** 1 I II* *1 I 1* 1 1 I I 1* 1* II Salticidae }otussp. I 1 I II* *1 *1 I * I * 1 I 1 1 I 1* 1* 1 Oonopidae Grymeus sp. 5 II II* I*II*I ** IIIIIIII Segestriidae Ariadna sp. 5 II I 1* 1* * 1*1 I I I I I 1 1 *1 *I Clubionidae Meedo houstoni 1*I ** I 1* *1*1I I IIII 1I I I Miturgidae Genus I, sp. 1 **I 1* *I 1* 1 I I I I I** I I I 1 I I Filistatidae Wandella sp. 5 ** II ** 1**** I** I I 1*** I 1 *1 I 1 I I I I Lycosidae Lycosa sp. 5 *****1*1** 1****** I ***********1****1*1** 1* 1**1 1 1*1 1 I Miturgidae Genus 2, sp. 6 ** * 1*I*** 1**** I************ I *** I*I I I*II 11I I Miturgidae Miturga sp. 1 * ***1 1 *1****** 1**** **** 1** 1*1* *1 I 1* I 1 1 1 1 Salticidae Bianorsp.2 *****1 1* 1**** 1* ** * 1** 1*1 **1 II 1* *1 I 1 I Prodidomidae Cryptoerithus sp. 3 * *** I 1 1 *** 1* * *** **I** * I 1*** III*I*II** II Prodidomidae Prodidomus sp. 1 ****1 1 **1**** 1 * ***1* *II 1* *I *1 I **1*1 1 *1 Lycosidae Lycosa bicolor * 1 I * I * * I ** ** I*1II II1*1II I Lycosidae Lycosa sp. 02 III ** ** 1 *******1*** 1 I 1* II 1*** 1*1 I 1 Miturgidae Genus 2, sp. 2 1 I* 1** *** I* ** *I**** I 1* 1* * I I I I 1 1 I Lycosidae Genus 2, sp. 2 *1*1* *1 ******1 *********** I ***1 1* *1**** 1* 1**1 *I 1***1 I* Oonopidae Myrmapopaea spp. **** 1*1- *1*******1** * *** * **1****1*1***1 **** *1**1**1****1*1***1*****1 Salticidae Grayenulla australensis ***** II*** I******* I************* I**** I* I * 1** ** I * I** I**** I* I*** I*****I Zoridae Argoctenus sp. 1 *****1*1** 1** * **1*** *********1****11***1***** * 1**1**1* **1*1 * 1*****1* Oxyopidae Oxyopes sp. 1 * *1 I* 1** **** 1* ** I**** I*1* * I**** I*I*I ** 1 1*** 1** 1 Oxyopidae Oxyopes sp. 2 *****1*1* *1 ******1**** *******1** 11***1 ** * I *1 I * II 1*** 1** Salticidae Zebraplatys keyserlingi *** *1 I * I***** *I** ** ****** I*** II** I *** I*1*I *II 1* 1 Pholcidae Trichocyclus sp. 1 * I 1 **1 ***** 1*************1****1*1* *1** 1**1 1 **1 II *1 Segestriidae Ariadna sp. 7 **** 11***1**** *1********** *1****1 I 1 I *1 I ***1 1**1 *1 Prodidomidae Molycriinae Genus 2, sp. 1 ***1 I *1 ******1************ 1****1*1** I *1**1**1****1*1** II Prodidomidae Molycriinae Genus 1, sp. I **III***** 1** *** ***** I*** I*I* I I* I 1**** 1 1*** I** *I Oonopidae Opopaea sp. I * **1 1 1*** * 1 ** ** **1****1*1 I 1**1* 1 I I I * * Salticidae Unidentati, Genus 05, sp. I *1 I 1**** *1 **1** *1*1 *1 1* I 1 1 1** 1 Lycosidae Lycosa sp. 04 1*1 **1 ** **1 ***1* 1 I 1 I 1**1* *1*1 *I Prodidomidae Cryptoerithus sp. 4 III ** **I ** * * *1 ** IIII** I** I * 1 1 I Miturgidae Miturga sp. 2 II 1** I ****1****1 I **1***** I *1**1* 1*1***1 Lycosidae Lycosa sp. 17 II 1* 1 * ** **I*III ** I*II**** II*I* Stiphidiidae Forsterina sp. I I 1** 1* I *****1 **1*1 1 **1 1* 1****1*1* *1*** Oonopidae Opopaea sp. 2 I I *1** **** 1* *III*III**** II*I ** * Trochanteriidae Genus 2, sp. 1 I 1 1**** ***1 **1 I 1 1* 1 1* 1** 1* Heteropodidae Neosparassus sp. 8 II*I*I** I 1 1 I** IIII Theridiidae Latrodectus hasseltii I I * 1 I*II I II*I 1*I Theridiidae Steatoda sp. 1 *I*I ** *1 * I ** I I I I* I* I I 1 Heteropodidae Heteropoda lralbarri II* **** I I 1 1 1* 1 I 1 I * I Lycosidae Lycosa sp. 08 I 1******* 1 *I III 11*1* I** I Salticidae Unidentati, Genus 02, sp. 1 I 1******* 1 *I I I ** 1 1 I 1 * I* I Lycosidae Lycosa sp. 13 I * 1****** I* ** 1 I1*I 1II* I*I **** I Oonopidae Grymeus sp. I II******* I ** III*II*II*I*** I** ** I Salticidae Lycidas sp. 2 I 1**** ** 1 * * *I** 1 I*** I I * 1 I* I 1 ** I Lycosidae Genus 2, sp. I *II ****** I I I I 1 * I** 1** 1** I*** I ** I Lycosidae Lycosa sp. 11 1** 1 * ** *1 I ***1*1 1 1 *1**1 ** *1***1 * 1 Oonopidae Grymeus sp. 6 I*I ***** I 1 I I I I I I I*** I *** I Miturgidae Genus 1, sp. 2 II *** 1 I I I I I 1* 1**** *I ** I 1 Salticidae Lycidas sp. 1 I 1 * * *I I I*I I I 1 1* ** 1 ** I** *I Theridiidae Euryopis sp. 2 II ** I 1 I I 1* * I * I* 1 *I** II Salticidae Ocrisiona leucocomis II**I ** ** *I 1*1 III*II*II Theridiidae Euryopis sp. 1 *II 1* ** I I III I*I1 I*I* I -----+-+---+------+------+----+-+---+-----+--+--+--+----+-+---+-----+------.

Araneomorph Spiders 317

Table 3 (cant.)

Group A BeD EFGHI J K L M N 0 P Q BBBKKINIMMMIBBBBNPplGGGMGGMMWWWWWIMMMMIKIBBBlccccclEEIKKlpplNNNNININNNlzzzzzjcp BOBEEIAIDDDIOOOOAEEIJJJRJJDDOOOOolRRRRIEIBBBIUUUUUILLIEEIEEjEEEEIEIAAAluuuuulUE 1234311112311345243113512445142351253415124512345611211212511345121345112453111 Assemblage-2 - Araneidae Dolophones sp. 2 I 1** I *I **I III I I 1 III I Lycosidae Lycosa sp. 03 * 1 I*** I I***I I I 1 I II III 1* Lycosidae Lycosa sp. 18 ***** III 1* *** *1 I I 1 *1 II III I Miturgidae Genus 3, sp. 1 ** j 1* *I I ********1* I* II I II 1 ** II 1 Prodidomidae Molycriinae Genus 3, sp. 1 *1*1***1 1* ** * * * **1 III I I 1 * I * II I Stiphidiidae Forsterina sp. 3 III * 1** ** ****1* I I 1 I I * I I ** II I Lycosidae Lycosa sp. 16 II 1* 1* * * ** I III I II III I Thomisidae Tharpyna sp. 2 * II* I 1** * ** **1 III I II 1 I I I Trochanteriidae Genus 4, sp. 1 I I 1 1** * **1 III 1 1 I I Prodidomidae Molycriinae Genus 1, sp. 3 I I 1* I *** I I 1 I I I 1 III'* II I Corinnidae Genus 1, sp. 1 I* II I * ** I I* I * I I II I I 1 I Heteropodidae Pediana tenuis III I **** I I* I * I I II III I Salticidae Paraplatoides sp. 1 I j I I ** 1 II ** I I 1 *1 I 1 I I Corinnidae Genus 3, sp. 1 1 I I 1* ** 1* III I I III I Lycosidae Lycosa sp. 19 III 1**** 1* III I I I 1 I 1 Gnaphosidae Hemicloea sp. 1 I 1 I 1* ** *I III I I 1 1 1 I Prodidomidae Prodidominae Genus 2, sp. 1 1 I I 1* *I III I I 1 * II 1 Lycosidae Genus 1, sp. 2 1*1 1** I 1 * III I I III I Salticidae Opisthoncus sp. 1 II 1** * I I I 1 * I 1 I III 1* Oonopidae Genus I, sp. 1 II 1**** I* 1* III I I I 1 I *1 Salticidae Bianorsp.1 1* I 1*** I 1 III I 1** * I 1 I * I Lycosidae Lycosa sp. 09 1*1 1*** 1* * 1 III I I III I Prodidomidae Molycriinae Genus 4, sp. 1 II 1*** I** I III I I I 1 1 I Theridiidae Genus 1, sp. 1 ** I*II * I*** I III 1 I III I Miturgidae Miturga sp. 3 *** III I * I I* II 1 I I I 1 I Oonopidae Opopaea sp. 4 **II *I * I* I 1* II I I 1 1 I I Oonopidae Grymeus sp. 2 *II *I * I I I 1* I I I III I -----+-+---+------+------+----+-+---+-----+--+--+--+----+-+---+-----+-- Assemblage-3 Araneidae Austracantha mrnax III II* I I * IIII I Clubionidae Clubiona sp. 2 *III *I 1I1* I*1 III Deinopldae Deinopis sp. 1 1 I 1* * 1 1 II * 1** 1 III Desidae Badumna insignis III I II 1 * I I I 1 * 1 Salticidae Unidentati, Genus 16, sp. 1 III 1 * 1*1 I * II* III .-----+-+---+------+------+----+-+---+-----+--+--+--+----+-+---+-----+-- Assemblage-4 Lycosidae Genus 1, sp. 1 III III '* ** 1** 1 I** IIII**** I Salticidae Unidentati, Genus 01, sp. 1 I 1 ** I III 1** *I** 1 1* II 1* *I**I Segestriidae Ariadna sp. 4 II 1 * IIIII** I* I* II 1* 1*** I Lycosidae Lycosa sp. 12 I 1 *1 II 1 **1 I *1 1**1*** 1*1 1*****1 Trochanteriidae Genus 3, sp. 1 III III * II 1* 1* II*I ** I**** I Lycosidae Lycosa sp. 21 III II I I*I** II*I*II*II Lycosidae Lycosa sp. 07 II * ** I I * I 1** IIII*II 1* *I **** I Lycosidae Lycosa sp. 10 I 1* *** *I I*II 1* ** 1 I I I * I I* I***** I Lycosidae Lycosa sp. 14 II* ***1* II 1* 1* **1* II *1 *I 1***1** **1 Miturgidae Genus 2, sp. I II 1 1****1 1 1**** III 1* I 1***1*** *1 Oonopidae Gamasomorpha sp. 1 1 1 I * I I*I * I** IIIII 1* *I**I Miturgidae MilUrga agelenina III I II*I I1II II I* ** I Miturgidae Uliodon tarantulinus III I II*I *II*I1 II*I **** I Salticidae Lycidas sp. 3 III I I I *I I I 1**1 I 1 ** I**I -----+-+---+------+------+----+-+---+-----+--+--+--+----+-+---+-----+-- Assemblage-S Heteropodidae Holconia nigrigularis II * I II II 1 I Oonopidae Grymeus sp. 3 II *1 II 1 1 1** I Trochanteriidae Genus 1, sp. 1 I 1 * I 1 I II I * *1 Stiphidiidae Corasoides sp. 1 1 * I *1 II * II 1 1 -----+-+---+------+------+----+-+---+-----+--+--+--+----+-+---+-----+-- Assemblage-6 Heteropodidae [sopedella saundersi II II 1* 1 I * I I 1 * I I Lycosidae Lycosa forresti II 1 1 I II 1* 1 II * I I Lycosidae Lycosa sp. 23 II II I I 1* II I* I * I I Heteropodidae Neosparassus sp. 4 II 1* *1 I II II II * I I Salticidae Unidentati, Genus 05, sp. 2 II I ** 1 I 1 1 II II * I I Prodidomidae Molycriinae Genus 2, sp. 3 II I ** I 1 I 1 II 1 I * I *1 -----+-+---+------+------+----+-+---+-----+--+--+--+----+-+---+-----+-- Assemblage-7 Heteropodidae Neosparassus sp. 9 III* III* I 1 I I I Stiphidiidae Genus 1, sp. 1 1 I 1* 1I I*I*I*I 1I Zoridae Thasyraea sp. 1 III I I 1 I** IIII -----+-+---+------+------+----+-+---+-----+--+--+--+----+-+---+-----+-- 318 M.S. Harvey, A. Sampey, P.L.]. West, ].M. Waldock

Table 3 (cant.)

Group A BC D E F GH I JKL M NO P Q ~ BOBEEIAlooOloOOOAEEIJJJRJJDDOOOOOIRRRRIEIBBBluuuuulLLIEEIEEIEEEEIEIAAAluuuuuluE 1234311112311345243113512445142351253415124512345611211212511345121345112453111 Assemblage-g Lycosidae Lycosa sp. 15 1 ** I I I I I 1* I* I I Salticidae Unidentati, Genus 12, sp. I 1 I I I * 1 I * I 1* * I I Prodidomidae Cryptoerithus sp, 5 I I I I I I *I I * I *I Salticidae Unidentati, Genus 07, sp. I *1 I 11 11* 11 I I Stiphidiidae Genus 6. sp, 1 *1 I I I I I 11 I I Salticidae Unidentati, Genus 15, sp. 1 *1 III 11 III I Theridiidae Gmogala sp. 1 ***1 I * I I I I I 1 I I -----+-+---+------+------+----+-+---+-----+--+--+--+----+-+---+-----+-- Assemblage-9 Clubionidae Meedosp. I I 1 1 I I I I I * I I I 1** 1 Miturgidae Genus 2, sp. 3 I I 1 I I I I I ** I 1 1 I **** I Heteropodidae Neosparassus sp. 3 I I I I I I 1 I I I I 1* * 1 Salticidae Margaromma sp. 1 I I I I I I I* I I I 1 1** **1 Salticidae Unidenlati, Genus 09, sp. 1 I I 1* I 1 I I * 1 * 1 I I * I ** I Stiphidiidae Forsterina sp. 2 I I I I I I I 1 I 11 **1**** I Milurgidae Miturga sp. 4 1 I I I I I I I I I 1 I * I Salticidae Holoplarys sp. 2 1 I I I I 1 I I I I 1 1* 1 Salticidae Unidentati, Genus 14, sp. I I 1 I I I 1 1* I I I I I * I Unyphiidae Genus 3, sp. 1 I I I 1 I I I I *I I I I I Salticidae Zebraplarys fractivittata 1 1 I I I I I I I I I I *I Thomisidae Stephanopis sp. 7 I 1 I -I I I 1 1* 1 I I I ** I -----+-+---+------+------+----+-+---+-----+--+--+--+----+-+---+-----+-- Assemblage-l0 Lycosidae Genus 5, sp. 1 I 1 I I* I I I Theridiidae Gmogala sp. 2 I I 1 I* I *1 *1 Prodidomidae Cryptoerithus sp. 6 I 1 I* I 1 1* 1 Salticidae Unidentati, Genus 13, sp. 1 I I I** I I I 1 Tetragnathidae Nephila edulis I I I* I I I 1 -----+-+---+------+------+----+-+---+-----+--+--+--+----+-+---+-----+-- Assemblage-ll Araneidae Genus 2. sp. 1 1 I * 1* ** I I I I* I I Segestriidae Ariadna sp. 2 I I 1* ** 1 I I I I 1 Miturgidae Genus 2, sp. 4 I I I*** I 1** * 1 I 1 1 Stiphidiidae Genus 2, sp. 1 I I I**I I *1 I I I Uocranidae Orthobula sp. 2 11 I * 11 I 1 I I I Theridiidae Steatoda sp. 3 I I 1** I I * 1 1 1 I I Gnaphosidae Hemicloea sp. 5 1 I I *I I I I I I I Heteropodidae Neosparassus sp. 5 I 1 I *I I I 1 1 I I Heleropodidae Neosparassus sp. 7 I I I *1 1 I I I I *1 -----+-+---+------+------+----+-+---+-----+--+--+--+----+-+---+-----+-- AssembJage-12 Corinnidae Genus 2, sp. 1 I* I I I *1 Lycosidae Genus 3, sp. I I* 1* I I I Thomisidae Stephanopis sp. 4 I* I I I I Gnaphosidae Hemicloea sp. 2 1** I 1 I I Salticidae Unidentati, Genus 18, sp. I I*I I I I Theridiidae Genus 3, sp. I I* 1 * I 1 I -----+-+---+------+------+----+-+---+-----+--+--+--+----+-+---+-----+-- Assemblage·13 Cyatholipidae Matilda sp.2 1 I I I I * 1 I I I I I Thomisidae Tharpyna sp. 3 1 I I I I * I I I I I I Salticidae Simaethula sp. 1 I I I I 1 ** I I I I I I Lycosidae Lycosa sp. 25 11 I 111*111 I11 Desidae Genus 1, sp. 1 * I I I I I 1 ****1 I I I I 1* I Lycosidae Lycosa sp. 22 * 1 1 I I I 1*** * I I I * I I I I 1 Prodidomidae Molycriinae Genus 2, sp. 2 1 1 I I 1 I **** I I I- I I 1 I 1 Salticidae Grayenulla sp. 2 I I I I I I ***** I I* I I I 1 1 1 Lycosidae Lycosu sp. 01 I I *1 11***1*****1 I I *1 I I I *1 Filistatidae Wandella sp. 3 1 I * I 1 I I * * 1 I I I 1 I I 1* Heteropodidae Neosparassus sp. 2 * I 1 * I I 1 * I * I I 1 I I 1 I I -----+-+---+------+------+----+-+---+-----+--+--+--+----+-+---+-----+-- Assemblage-14 Corinnidae Genus 4, sp. 1 *1 I I I *1 * 1 1**1* 1 1 I I I Diclynidae Genus 2, sp. 1 *1 *I I I *I I 1* 1* 1* I I *1 I ,Oonopidae Grymeus sp. 4 I* * I I I *1 I I I * I I I 1 I Filistatidae Wandella sp. 2 **1* I I 1 I *1 I 1 *I *I I I I Lycosidae Lycosa sp. 06 **1 I I I I 1 I 1** 1 I I I *1 Thomisidae Tharpyna sp. I **1 I I1** 1 I I I *I I I I I Cyatholipidae Matildasp.1 I I I 1 * I I I I ** I I I 1 I Filistatidae Wandella sp. 1 I I I I I I I I ** I** I I I I Araneomorph Spiders 319

Table 3 (cant.)

Group A BeD EFGHI J K L M N 0 P Q BBBKKINIMMMIBBBBNPP IGGGMGGMMWWWWWIMMMMIKIBBBlccccclEEIKKI PP 1NNNN IN INNN IZZ zzzlcp BOBEEIAIDDDIOOOOAEEIJJJRJJDDOOOOOIRRRRIEIBBBluuuuulLLIEEIEEIEEEEIEIAAAluuuuulUE 1234311112311345243113512445142351253415124512345611211212511345121345112453111 Salticidae Unidentati, Genus 02, sp. 2 1 1 *1 1 1 1I1 1II1 1I1 1 Salticidae Simaelha sp. 1 1 1* 1 I I 1 1 1 1 I 1 I 1 I 1 1 Stiphidiidae Genus 5, sp. 1 I 1* 1 1 1 1 II 1 1 1 1 1 1 1 I -----+-+---+------+------+----+-+---+-----+--+--+--+----+-+---+-----+-- Assemblage-23 Salticidae Unidentati, Genus 08, sp. 1 I I 1 * III 1 1 Thomisidae Slephanopis sp. 1 I 1 1 *1 1 1 1 1 Tetragnathidae Genus I, sp. 1 1 1 * 1 *1 1 I I 1 -----+-+---+------+------+----+-+---+-----+--+--+--+----+-+---+-----+-- Assemblage-24 Corinnidae Genus 6, sp. 1 *1 1 1 * 1 1 Gallieniellidae Genus I, sp. 1 *1 1 1 II Pholcidae Trichocyc/us sp. 2 *1 1 I I 1 Salticidae Lycidas sp. 4 *1 II 1 1 Salticidae Zenodorus sp. 1 *1 II II -----+-+---+------+------+----+-+---+-----+--+--+--+----+-+---+-----+-- Assemblage-25 Araneidae Eriophara sp. 1 I I 1 1 I 1 1* 1 1* I Miturgidae Genus 2, sp. 5 I 1 1 II I I 1 1 1 Prodidomidae Prodidomus sp. 3 1 1 I * 1 1 1 1 I I I Salticidae Maralus vesper/ilio 1 1 1 * 1 1 1* 1 1 1 1 Araneidae Eriophora sp. 3 1 1 1 * 1 I 1 II II Araneidae Genus 1, sp. 1 1*I I * II 1 1 1 II * -----+-+---+------+------+----+-+---+-----+--+--+--+----+-+---+-----+-- Assemblage-26 Salticidae Clynolis sp. 1 11 I 11 II Thomisidae Slephanopis sp. 3 11 * I II 1 1 -----+-+---+------+------+----+-+---+-----+--+--+--+----+-+---+-----+-- Assemblage.27 Clubionidae Fissarena sp. 1 II 1 1 1 I 1 1 Lamponidae Lampona cylindrala II I I 1 * 1 1 1 * Salticidae Hypoblemum sp. 1 1 1 *1 I I 1 * 1 1 Hersiliidae Tamopsis occidenlalis I 1 1 1 I I I I Oonopidae Orcheslina sp. 1 II I 1 I 1 I 1 Thomisidae Slephanopis sp. 6 II I 1 I 1 I 1 -----+-+---+------+------+----+-+---+-----+--+--+--+----+-+---+-----+-- AssembJage-28 Araneidae Dolophones sp.5 I 1 1 1 1* Theridiidae Genus 2, sp. 1 II I 1 1* Desidae Genus 5, sp. 1 1 1 I 1 1 1** Theridiidae Enoplognalha sp. 1 1 1 I 1 1 1** Salticidae Margaromma sp. 2 1 1 I 1 1 1 * Araneidae Heurodes sp. 1 1*1 1 1 1 1 Salticidae Unidentati, Genus 06, sp. 1 1*1 1 I 1 I Araneidae Larinia sp. 1 II *1 1 I I Dictynidae Genus I, sp. 1 1* I *1 II I -----+-+---+------+------+----+-+---+-----+--+--+--+----+-+---+-----+-- Assemblage-29 Clubionidae CheiracanlhlUm sp. 1 1*1 t I 1 1 t 1 1 Prodidomidae Prodidominae Genus 1, sp. 1 * 1*1 t I 1 1* I 1 1 Dictynidae Genus 3, sp. 1 * 1 1 I * 1 1 1 I 1 1 * Salticidae Opislhoncus sp. 2 I 1 1 I 1 1 *I 1 1 * Salticidae Unidentati, Genus 03, sp. 1 * 1* I 1 I *1 1 I 1 1** Theridiidae Slealoda sp. 2 II 1* 1 1 I *1 *1 1 I Gallieniellidae Genus I, sp. 2 * 1 I 1 1 1 I 1 1 1 1* Salticidae Grayenulla sp. 3 * 1 1 1 1 1 I 1 I 1 1 I Selenopidae Selenops sp. 1 * 1 1 I 1 1 1 1 I I 1 I Linyphiidae Genus 2, sp. 1 * *1 1 1 1 I 1 1 1 1 I I Salticidae Cylaea sp.1 *** I 1 1 II 1 1 1 I I I -----+-+---+------+------+----+-+---+-----+--+--+--+----+-+---+-----+-- Assemblage.30 Araneidae Genus I, sp. 2 I I 1 1 1 * ** I 1 Salticidae Lycidas sp. 5 II I 1 1 ** I 1 Salticidae Zebraplatys sp. 3 1*1 II I * 1 I Corinnidae Genus 5, sp. 1 1 1 II I * 1 1 Cyatholipidae Malildasp.3 I 1 1 1 I * 1 1 Liocranidae Orthobula sp. 3 II 1 1 1 * 1 I Sallicidae Unidentati, Genus 10, sp. 1 1 1 1 1 1 *II Theridiidae Trigonobolhrys sp. 1 1 1 II 1 * I 1 320 M.S. Harvey, A. Sampey, P.L.J. West, J.M. Waldock

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