Dippenaar SOVENGA.qxd A checklist of spiders from Sovenga Hill, an inselberg in the Savanna Biome, Limpopo Province, South Africa (Arachnida: Araneae) M.A. MODIBA, S.M. DIPPENAAR and A.S. DIPPENAAR-SCHOEMAN Modiba, M.A., S.M. Dippenaar and A.S. Dippenaar-Schoeman. A checklist of spiders from Sovenga Hill, an inselberg in the Savanna Biome, Limpopo Province, South Africa (Arachnida: Araneae). Koedoe 48(2): 109–115. Pretoria. ISSN 0075-6458. The South African National Survey of Arachnida (SANSA) was initiated to make an inventory of the arachnid fauna of South Africa. Various projects are underway to pre- pare inventories of the spider fauna of the different floral biomes and provinces of South Africa. During April and May 2004 five different collecting methods were used to sam- ple spiders from four slopes on Sovenga Hill, an inselberg situated in the Savanna Biome, near Polokwane, in the Limpopo Province of South Africa. A total of 793 spec- imens represented by 29 families, 62 genera and 76 species were recorded over the two- month period. The Thomisidae was the most abundant (n = 167) representing 21.1 % of all spiders sampled, followed by the Gnaphosidae (n = 101) with 12.7 % and the Lycosi- dae (n = 77) with 9.7 %. The most abundant species was a thomisid Tmarus comellini Garcia-Neto (n = 82), representing 10.3 % of the total, followed by a clubionid Clu- biona godfreyi Lessert (n = 66) with 8.3 %. The Thomisidae was the most species-rich family with 12 species, followed by the Gnaphosidae with 11 species and the Araneidae with 10 species. Of the species collected 83.9 % were wandering spiders and 16.1 % web builders. This is the first quantitative survey of the Savanna Biome in the Polok- wane area. Keywords: Araneae, checklist, diversity, Limpopo Province, Savanna Biome, South Africa, spiders. M.A. Modiba, S.M. Dippenaar, School of Molecular and Life Sciences, University of Limpopo, Private Bag X1106, Sovenga, 0727, South Africa; A.S. Dippenaar-Schoeman, ARC-Plant Protection Research Institute, Private Bag X134, Queenswood, 0121, South Africa/Department of Zoology and Entomology, University of Pretoria, South Africa ISSN 0075-6458 109 Koedoe 48/2 (2005) Introduction Although spiders constitute an abundant and successful group of invertebrates in South Africa they are poorly sampled and little is known about their diversity within most eco- regions (Dippenaar-Schoeman 2002a). This lack of knowledge undermines meaningful conservation (De Wet & Schoonbee 1991). Additionally, the critical lack of professional taxonomic expertise within the country pre- sents significant problems for the under- standing of invertebrate biodiversity, which leads to an under-appreciation and estimation of the actual species pool (Gibbons 1999; Dippenaar-Schoeman 2002a). In South Africa, more than 2000 spider species have been recorded but only about 30 % of the families have been revised (Dippenaar- Schoeman 2002a). The South African National Survey of Arach- nida (SANSA) was initiated in 1997 to address this lack of baseline information by conducting surveys and biodiversity assess- ments of the arachnid fauna of South Africa (Dippenaar-Schoeman & Craemer 2000). One such a project of SANSA is to determine spider diversity in the Savanna Biome of South Africa, with the primary aim to gather information on areas still poorly sampled (Dippenaar-Schoeman & Leroy 2003; Van den Berg et al. 2003). Research on the spider fauna of the Savanna Biome in South Africa is presently restricted to surveys of the Roodeplaat Dam Nature Reserve (Dippenaar-Schoeman et al. 1989); Makalali Game Reserve (Whitmore et al. 2001; Whitmore et al. 2002), the spiders of the Western Soutpansberg mountains (Foord et al. 2002), a survey of the spiders of the Kruger National Park (Dippenaar-Schoeman & Leroy 2003) and the spiders of the Spring- bok Flats (Van den Berg et al. 2003). There are numerous collecting methods used for sampling arachnids (Eardley & Dippe- naar-Schoeman 1996), but there are biases associated with some methods frequently resulting in under-sampling of sites when only one method is used or one habitat is sampled (New 1999). Scientists have for years recognised that standard methods are essential to be able to compare biodiversity assessments (Coddington et al. 1996). Spi- ders are able to occupy nearly every terres- trial habitat (Dippenaar-Schoeman & Jocqué 1997). It is therefore essential to sample the fauna separately at different layers during biodiversity surveys. Areas showing low diversity may simply be a reflection of inad- equate sampling (Whitmore et al. 2002), therefore sampling protocols are very impor- tant. Unfortunately, many of the surveys undertaken in South Africa have used restricted sampling techniques, sampling only one habitat type. In this study, a survey was undertaken to determine the spider diversity of Sovenga Hill, an inselberg, situated near Polokwane, Limpopo Province. During a two-month period this inselberg was extensively sur- veyed using five collecting methods. Twen- ty-four sites were sampled and samples were taken from both the ground, litter, grass and tree layers. The collecting methods were evaluated for surveys to be continued in the Polokwane area. This is the first of a series of papers on spider diversity of the Savanna Biome in the Polokwane area. Material and methods Sovenga Hill (23º53'S, 29º44'E), an inselberg, was sampled over a two-month period from 1 April to 31 May 2004. This unique landform consists of dome- shaped granite rocks and is situated on the campus of the University of Limpopo near Polokwane in the Limpopo Province of South Africa. It covers an area of approximately 6.3 ha and is surrounded by indige- nous flora consisting mostly of Euphorbia cooperi N.E.Br. ex A. Berger var. cooperi, Heteropyxis natal- ensis Harv. and Croton gratissimus Burch. var. gratissimus. This is a closed woodland plant com- munity classified as Mamabolo Mountain Bushveld of which small, outlying, isolated communities are found on the Pietersburg Plateau (S.A. vegmap pro- ject, in prep. - P. Winter pers. comm.). Six sampling sites of 4 m x 4 m were selected on each of the southern, eastern, northern and western slopes, resulting in a total of 24 sites sampled. The first two sites were at an altitude of between 1318 m–1335 m, the second two sites between 1333 m–1354 m and the third two sites between 1347 m–1374 m. Sampling was conducted twice a week during April and May 2004. During April sampling was conduct- ed on all the sites while no sampling was done on the sites of the northern slope during May due to van- dalism of the sampling sites. The following methods were used during sampling: Sweep netting (S) — a sweeping net was used to sweep through the grass and herb layer to dis- lodge specimens into the net. Walking in a straight line, 20 sweeps were taken to form one sample. The contents from the net were emptied into a marked plastic bag and spiders and other organisms were separated from the vegetation in the laboratory. A total of 101 sweepnet samples were taken. This method could only be used at sites where grass and herb layers were present. Active searching (A) — active lifting of stones and searching for spiders was done at all sites for 10 minutes per sampling session (eight session dur- ing April and nine in May). Tree beating (T) — a stick was used to knock spiders from trees while a tray was used to catch all the specimens and plant material knocked off. A total of 20 beats formed one sample. The con- tents from the tray were placed in a marked plas- tic bag and spiders and other organisms were separated from the vegetation in the laboratory. A total of 354 samples were collected during the sampling period using this method. Different types of trees present on a site were sampled. Pitfall trapping (P) — small plastic containers (10.5 cm diameter) were planted into the ground, with the upper rim level with the ground surface. Wooden planks of about 30 cm long were placed in three directions towards the trap to increase its effective catch area. A funnel was placed over the opening of the trap to prevent Koedoe 48/2 (2005) 110 ISSN 0075-6458 spiders from escaping and a small container with 70 % ethanol was placed inside to immo- bilise and preserve the captured specimens. There were 20 pitfall traps per slope, randomly distributed. The traps were kept open for the whole period and were emptied twice a week. Leaf litter sifting (L) — a leaf litter sample was col- lected in a wooden box (53 cm x 20 cm x 18.5 cm) with a chicken wired base and speci- mens were sieved from the litter. One sample consisted of enough litter to fill the wooden box. Two samples of litter were collected from sites where leaf litter was found. Captured specimens were preserved in 70 % ethanol and were sorted and counted in the laboratory. All spiders were identified to species level by the third author, where possible. Some specimens could not be identified to species owing to the unresolved tax- onomy of certain families in Africa (for instance the Lycosidae) and the immature stages of some speci- mens collected. Voucher specimens were deposited in the National Collection of Arachnida (NCA) at the Plant Protection Research Institute in Pretoria, an institute of the Agricultural Research Council. Results and discussion During the two-month sampling period a total of 793 spiders, represented by 29 fam- ilies, 62 genera and 76 species were collect- ed (Table 1). The few published surveys of spiders from herbaceous and ground layers in Africa indicate that the relative abun- dance of species differs between different areas and different plant biomes (Dippenaar- Schoeman et al. 1989; Russell-Smith 1981). Three surveys of spiders are known from the Limpopo Province. A study at Makalali Pri- vate Game Reserve was conducted over 11 months and resulted in a total of 4832 spiders collected, representing 38 fam- ilies (Whitmore et al. 2002). Another study, conducted over a two-year period on the Springbok Flats, resulted in a total of 3139 specimens caught, representing 35 families (Van den Berg et al. 2003). Spo- radic collection over a five-year period at Lajuma in the western Soutpansberg result- ed in 127 species belonging to 46 families (Foord et al. 2002). From the above it is clear that although the current study was only conducted for two months, on an iso- ISSN 0075-6458 111 Koedoe 48/2 (2005) Table 1 Spider families collected at Sovenga Hill indicating the number of species and percentage of the total number of specimens collected Family Genera Species Specimens % Agelenidae 1 1 6 0.8 Amaurobiidae 1 1 4 0.5 Araneidae 8 10 57 7.2 Corinnidae 3 3 42 5.3 Clubionidae 1 1 66 8.3 Caponiidae 1 1 9 1.1 Ctenidae 1 1 7 0.9 Deinopidae 1 1 2 0.3 Eresidae 1 1 8 1.0 Gnaphosidae 5 11 101 12.7 Hahniidae 1 1 6 0.8 Hersiliidae 1 1 3 0.4 Linyphiidae 1 1 7 0.9 Lycosidae 2 2 77 9.7 Miturgidae 1 1 7 0.9 Nemesiidae 1 1 11 1.4 Oxyopidae 1 1 18 2.3 Palpimanidae 1 1 4 0.5 Philodromidae 2 5 51 6.4 Pholcidae 1 1 2 0.3 Pisauridae 1 1 9 1.1 Salticidae 5 5 54 6.8 Scytodidae 1 1 16 2.0 Segestriidae 1 1 3 0.4 Selenopidae 1 1 12 1.5 Theridiidae 6 6 21 2.6 Thomisidae 9 12 167 21.1 Uloboridae 1 1 3 0.4 Zodariidae 2 2 20 2.5 TOTAL 62 76 793 100 lated hill, a good representation of families (29) were caught. However, because of the short duration of the present study, the results may not reflect the actual abundance and species richness of Sovenga Hill, but nevertheless provide an indication of the minimum abundance and richness. The family composition of the spider fauna as a whole is shown in Tables 1 and 2. The Thomisidae was the most abundant family (n = 167) representing 21.1 % of all spiders sampled, followed by the Gnaphosidae (n = 101) representing 12.7 %, the Lycosi- dae (n = 77) representing 9.7 % and Clu- Koedoe 48/2 (2005) 112 ISSN 0075-6458 Table 2 Checklist of the spiders of Sovenga Hill, Polokwane, South Africa. (A: active search; B: beating; L: litter sifting; P: pittrapping; S: sweepnetting; W: wanderer; WD: web dweller) Family Species Nr Method Guild Habitat Agelenidae Agelena sp. (immature) 6 A WD funnel-web Amaurobiidae Chresiona sp. (immature) 4 P;L WD retreat-web Araneidae Argiope sp. (immature) 1 B WD orb-web Argiope australis (Walckenaer, 1805) 3 B WD orb-web Caerostris sexcuspidata (Fabricius, 1793) 5 B WD orb-web Cyphalonotus larvatus (Simon, 1881) 5 B;L WD orb-web Cyrtophora citricola (Forskal, 1775) 1 B WD orb-web Nemoscolus sp. (immature) 1 B WD orb-web Neoscona blondeli (Simon, 1885) 20 P;B;L WD orb-web Neoscona subfusca (C.L. Koch, 1837) 19 B;L;A WD orb-web Prasonica sp. (immature) 1 B WD orb-web Singa sp. (immature) 1 L WD orb-web Caponiidae Caponia sp. (immature) 9 P;A W soil Clubionidae Clubiona godfreyi Lessert, 1921 66 P;B;L W soil Corinnidae Thysanina sp. 12 P;L W soil Cetonana sp. (immature) 13 P;L W soil Copa flavoplumosa Simon, 1885 17 B;L W soil Ctenidae Ctenus sp. (immature) 7 L;B W soil Deinopidae Menneus camelus Pocock, 1902 2 B WD throw-web Eresidae Gandanameno sp. (immature) 8 P;B;L WD retreat-web Gnaphosidae Asemesthes sp. (immature) 2 L;B W soil Camillina sp. (immature) 6 L;P;B W soil Drassodinae (undetermined) 2 B W soil Setaphis browni (Tucker, 1923) 12 L;A;P W soil Zelotes lightfooti (Purcell, 1907) 21 L;P W soil Zelotes oneili (Purcell, 1907) 13 P;L W soil Zelotes reduncus (Purcell, 1907) 12 B;A;L;P W soil Zelotes ungulus Tucker, 1923 17 L;P W soil Zelotes sp. A 3 P W soil Zelotes sp. B 3 B;P W soil Hahniidae Hahnia tabulicola Simon, 1898 6 L WD sheet-web Hersiliidae Hersilia sericea Pocock, 1898 3 B W tree trunk Linyphiidae sp. (undetermined) 7 P WD sheet-web Lycosidae sp. (undetermined) 69 P W soil Pardosa sp. 8 P W soil Miturgidae Cheiracanthium sp. (immature) 7 B W leaves Nemesiidae Lepthercus sp. 11 P;B W burrow soil Oxyopidae Oxyopes schenkeli Lessert, 1927 18 P;B W grass Palpimanidae Palpimanus transvaalicus Simon, 1893. 4 P W soil Philodromidae Philodromus sp. (immature) 10 B;A,S W grass Philodromus brachycephalus Lawrence, 1952 15 B,S W grass Philodromus grosi Lessert, 1943 1 B,S W grass Philodromus partitus Lessert, 1919 3 S W grass Suemus punctatus Lawrence, 1938 22 B W grass Pholcidae Smeringopus sp. (immature) 2 B WD space-web Pisauridae Chiasmopes sp. (immature) 9 S;A;B WD sheet-web Salticidae Heliophanus sp. (immature) 5 B W leaves Natta sp. 18 L W soil Portia sp. 4 L W litter Thyene ogdeni Peckham & Peckham, 1903 24 B W leaves bionidae (n = 66) representing 8.3 %. The rest of the families were low in abundance and none exceeded 7 % of the total. These percentages differ from other studies done in the Savanna Biome, possibly because of the difference in vegetation types. During the study on the Makalali Game Reserve the two vegetation types sampled were mixed lowveld bush and mopane bushveld, and 32 % of the specimens caught belonged to the Araneidae, followed by the Salticidae (18 %) and Thomisidae (10 %) (Whitmore et al. 2002). The study of Roodeplaat Dam Nature Reserve was done in an open savan- na and Tetragnathidae with 29.3 % was the most abundant family, followed by the Araneidae with 22.7 % and Salticidae with 21.4 % (Dippenaar-Schoeman et al. 1989). On the Springbok Flats, an open savanna but more disturbed area, Lycosidae was the dominant family (30 %), followed by Gnaphosidae (21 %) (Van den Berg et al. 2003). The Thomisidae was the most species-rich family with 12 species followed by the Gnaphosidae with 11 species and the Aranei- dae with 10 species. Twenty families were represented by a single species (Table 1). The three most abundant species was a thomisid Tmarus comellini Garcia-Neto (n = 82), representing 10.3 % of the total, ISSN 0075-6458 113 Koedoe 48/2 (2005) Scytodidae Scytodes sp. 16 L;P W soil Segestriidae Ariadna sp. (immature) 3 P WD soil Selenopidae Anyphops sp. 12 B W tree Theridiidae Chorizopella tragardhi 1 B WD g u m f o o t - Lawrence, 1947 web Dipoena sp. 4 B WD gumfoot- web Enoplognatha sp. 4 B WD gumfoot- web Episinus bilineatus Simon, 1894 1 B WD gumfoot- web Euryopis sp. 1 B WD gumfoot- web Theridion sp. 10 B WD gumfoot- web Thomisidae Heriaeus fimbriatus Lawrence, 1942 10 S,P W grass Misumenops rubrodecoratus Millot, 1941 2 S W grass Monaeses austrinus Simon, 1910 3 B W tree Oxytate argenteooculata (Simon, 1886) 4 B W tree Runcinia flavida (Simon, 1881) 1 S W grass Stiphropus bisigillatus Lawrence, 1952 5 P W soil Thomisus granulatus Karsch, 1880 2 S W grass Thomisus scrupeus (Simon, 1886) 2 S W grass Thomisus sp. (immature) 2 S W grass Tmarus africanus Lessert, 1919 25 S,P W grass Tmarus comellini Garcia-Neto, 1989 82 S,P W grass Xysticus lucifugus Lawrence, 1937 29 B;P W soil Uloboridae Miagrammopes brevicaudus O.P.-Cambridge, 1882 3 B;P WD vegetation Zodariidae Cydrela sp. (immature) 3 P W soil Diores sp. (immature) 17 B;P W soil TOTAL 793 Table 2 (continued) followed by the clubionid Clubiona godfreyi Lessert (n = 66), representing 8.3 % and an undetermined lycosid (n = 69), representing 8.7 %. Neither T. comellini nor C. godfreyi have before been collected in high numbers during surveys. Tmarus comellini is known from Zaire and South Africa and has previ- ously been collected from the grass and tree layer. It has a fairly wide distribution throughout South Africa and is known from the Eastern Cape, KwaZulu-Natal, Gauteng and Limpopo Province. Clubiona godfreyi is a species first recorded in Uganda. The first record from South Africa is from Lajuma, Soutpansberg. Species dominance also differs from area to area. A quantitative survey of the herbaceous layer of coastal dune forest at Richards Bay, South Africa showed the pisaurid Charminus atomarius (Lawrence) to be the most abun- dant species, representing 18 % of the total, followed by the salticid Thyene ogdeni (Peckham & Peckham) with 12 % and the araneid Caerostris sexcuspidata (Fabricius) with 11 % (Dippenaar-Schoeman & Wasse- naar in prep.). Most spiders live in a defined environment with limitations set by both physical condi- tions and biological factors (Foelix 1982) and species can be grouped into guilds based on available information on their habitat preferences and predatory methods. A guild is a group of species that potentially compete for jointly exploited limited resources (Polis & McCormick 1986). For the present study two main guilds were recognised, namely wandering spiders (W) and web builders (WB), with further subdivisions based on microhabitat and web structure (Dippenaar- Schoeman et al. 2005). Twelve of the col- lected families (16.1 %) are web dwellers. Three families namely Araneidae, Deinopi- dae and Uloboridae, construct orb-webs or adapted orb-webs while the Amaurobiidae, Eresidae and Segestriidae build retreat-webs usually low in vegetation, the Hahniidae, Linyphiidae and Pisauridae construct sheet- webs, the Pholcidae space-webs, the Theridi- idae gumfoot-webs and the Agelenidae fun- nel-webs. Seventeen of the collected families (83.9 %) are wanderers. Of the wanderers 55.6 % (370) are soil dwellers while 44.4 % (295) were collected from plants. The lack of grass on parts of Sovenga Hill may have con- tributed to the lower number of plant dwellers collected. Only one mygalomorph species, a member of the Nemesiidae (wish- bone trapdoor spiders) was collected. This is the first record of a Lepthercus species from an area outside the Eastern Cape (Dippenaar- Schoeman 2002b). Studies indicated that the physical structure of the habitat could have a distinct effect on the composition of the spi- der community (Wise 1993). Vegetation not only provides the necessary support for anchoring webs but it also increases the availability of retreat space and modifies the microclimate, which could have an effect on the spiders as well as their prey. Therefore, the structure of the vegetation is expected to influence the diversity of spiders found in a habitat (Whitmore et. al. 2002). From a sur- vey conducted at Swartberg Nature Reserve in the Succulent Karoo Biome, 76.5 % of the spiders collected over a 10-year period were wanderers and 23.5 % were web dwellers. Of the wanderers 56.7 % were associated with the ground layer (Dippenaar-Schoeman et al. 2005). Since the present study was conducted for only a short period of time (two months), numbers were too low to be able to make any meaningful comparisons amongst spiders collected from different slopes or different altitudes. Conclusion In South Africa there is a lack of baseline information on most invertebrate taxa. This causes a serious problem in conservation planning. With the ecology and diversity of the spider fauna of the Savanna Biome still poorly known, this study is a contribution to our knowledge of the geographical distribu- tion of species. As such, it represents new distribution records for all the species recorded and specimens of some unde- scribed species are now available for sys- Koedoe 48/2 (2005) 114 ISSN 0075-6458 tematic research. Since the present study was conducted over a relatively short period totaling only 17 sampling sessions over two months, the results may not reflect the actu- al abundance and species richness of Soven- ga Hill. However, it indicates the importance to sample more than one habitat type in order to obtain a clearer picture of species richness and abundance. Acknowledgements We thank the following people for assistance during the study: Mrs. A. van den Berg (ARC-Plant Protec- tion Research Institute), Mr P. Winter (South African National Biodiversity Institute) and from the Univer- sity of Limpopo (Turfloop Campus) Mr. P. Schoe- man, Mrs. H. du Plessis, Job Matlebjane and Mrs. R. Olwagen. Additionally we acknowledge both Stel- lenbosch University and the Stellenbosch Universi- ty/DEAT/USAID CBP-CCR for sponsoring the study and the provision of a student bursary. References CODDINGTON, J.A., L.H. YOUNG & F.A. COYLE. 1996. Estimating spider species richness in a southern Appalachian Cove hardwood forest. Journal of Arachnology 24: 111–128. DE WET, J.I. & H.J. SCHOONBEE. 1991. The occur- rence and conservation status of Ceratogyrus bechuanicus and C. brachycephalus in the Transvaal, South Africa. Koedoe 34: 69–75. DIPPENAAR-SCHOEMAN, A.S. 2002a. Status of South African Arachnida fauna. Proceedings of the symposium on the Status of South African species organized by the Endangered Wildlife Trust (EWT) of South Africa. Rosebank. 4-7 Sep- tember. DIPPENAAR-SCHOEMAN, A.S. 2002b. Baboon and trapdoor spiders of Southern Africa: an identifi- cation manual. Pretoria: Agricultural Research Council. (Plant Protection Research Institute Handbook no. 13). DIPPENAAR-SCHOEMAN, A. S. & C. CRAEMER. 2000. The South African National Survey of Arachni- da. Plant Protection News 56: 11–12. DIPPENAAR-SCHOEMAN, A.S. & R. JOCQUÉ. 1997. African spiders, an identification manual. Pre- toria: Agricultural Research Council. (Plant Pro- tection Research Institute Handbook no. 9). DIPPENAAR-SCHOEMAN, A.S. & A. LEROY. 2003. A checklist of the spiders of the Kruger National Park, South Africa (Arachnida: Araneae). Koe- doe 46: 91–100. DIPPENAAR-SCHOEMAN, A.S., A.M. VAN DEN BERG, & A. VAN DEN BERG. 1989. Species composi- tion and relative seasonal abundance of spiders from the field and tree layers of the Roodeplaat Dam Nature Reserve. Koedoe 32: 51–60. DIPPENAAR-SCHOEMAN, A.S., A.E. VAN DER WALT, M. DE JAGER, E. LE ROUX & A. VAN DEN BERG. 2005. The spiders of the Swartberg Nature Reserve in South Africa (Arachnida: Araneae). Koedoe 48: 77–86. EARDLEY, C.D. & A.S. DIPPENAAR-SCHOEMAN. 1996. Collection and preparation of material. I. Col- lecting methods. In: UYS, V.M. & R.P. URBAN (eds.). How to collect and preserve insects and arachnids. Pretoria: Agricultural Research Council. (Plant Protection Research Institute Handbook no. 7). FOELIX, R.F. 1982. Biology of spiders. Cambridge, Massachusetts: Harvard University Press. FOORD, S.H., A.S. DIPPENAAR-SCHOEMAN & M. VAN DER MERWE. 2002. A checklist of the spider fauna of the Western Soutpansberg, South Africa (Araneae). Koedoe 43: 35–43. GIBBONS, M.J. 1999. The taxonomic richness of South Africa's marine fauna: a crisis at hand. South African Journal of Science 95: 8–12. NEW, T.R. 1999. Untangling the web: spiders and the challenges of invertebrate conservation. Journal of Insect Conservation 3: 251–256. POLIS, G.A. & S.J. MCCORMICK. 1986. Scorpions, spiders and solfugids: predation and competition among distantly related taxa. Oecologia 71: 111–116. RUSSELL-SMITH, A. 1981. Seasonal activity and diversity of ground-living spiders in two African savanna habitats. Bulletin of the British Arach- nological Society 5: 145–154. VAN DEN BERG, A., A.S. DIPPENAAR-SCHOEMAN, E. UECKERMANN, M. VAN JAARSVELD & E. VAN DER WALT. 2003. Diversity of the Arachnida fauna of a Savanna Biome in the Limpopo Province. Proceedings of the 4th Congress of the Southern African Society for Systematic Biology. Pretoria. 9 -11 July 2003. WHITMORE, C., T.E. CROUCH, R. SLOTOW & A.S. DIPPENAAR-SCHOEMAN. 2001. Checklist of spi- ders (Araneae) from Makalali Private Game Reserve, Northern Province, South Africa: including a new family record. Durban Museum Noviates 26: 10–19. WHITMORE, C., R. SLOTOW, T.E. CROUCH & A.S. DIPPENAAR-SCHOEMAN. 2002. Diversity of spi- ders (Araneae) in a Savanna Reserve, Northern Province, South Africa. Journal of Arachnology 30: 344–356. WISE, D.H. 1993. Spiders in ecological webs. Cam- bridge: University Press. ISSN 0075-6458 115 Koedoe 48/2 (2005) << /ASCII85EncodePages false /AllowTransparency false /AutoPositionEPSFiles true /AutoRotatePages /None /Binding /Left /CalGrayProfile (Dot Gain 20%) /CalRGBProfile (sRGB IEC61966-2.1) /CalCMYKProfile (U.S. Web Coated \050SWOP\051 v2) /sRGBProfile (sRGB IEC61966-2.1) /CannotEmbedFontPolicy /Error /CompatibilityLevel 1.4 /CompressObjects /Tags /CompressPages true /ConvertImagesToIndexed true /PassThroughJPEGImages true /CreateJDFFile false /CreateJobTicket false /DefaultRenderingIntent /Default /DetectBlends true /ColorConversionStrategy /LeaveColorUnchanged /DoThumbnails false /EmbedAllFonts true /EmbedJobOptions true /DSCReportingLevel 0 /EmitDSCWarnings false /EndPage -1 /ImageMemory 1048576 /LockDistillerParams false /MaxSubsetPct 100 /Optimize true /OPM 1 /ParseDSCComments true /ParseDSCCommentsForDocInfo true /PreserveCopyPage true /PreserveEPSInfo true /PreserveHalftoneInfo false /PreserveOPIComments false /PreserveOverprintSettings true /StartPage 1 /SubsetFonts true /TransferFunctionInfo /Apply /UCRandBGInfo /Preserve /UsePrologue false /ColorSettingsFile () /AlwaysEmbed [ true ] /NeverEmbed [ true ] /AntiAliasColorImages false /DownsampleColorImages true /ColorImageDownsampleType /Bicubic /ColorImageResolution 300 /ColorImageDepth -1 /ColorImageDownsampleThreshold 1.50000 /EncodeColorImages true /ColorImageFilter /DCTEncode /AutoFilterColorImages true /ColorImageAutoFilterStrategy /JPEG /ColorACSImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /ColorImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /JPEG2000ColorACSImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /JPEG2000ColorImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /AntiAliasGrayImages false /DownsampleGrayImages true /GrayImageDownsampleType /Bicubic /GrayImageResolution 300 /GrayImageDepth -1 /GrayImageDownsampleThreshold 1.50000 /EncodeGrayImages true /GrayImageFilter /DCTEncode /AutoFilterGrayImages true /GrayImageAutoFilterStrategy /JPEG /GrayACSImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /GrayImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /JPEG2000GrayACSImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /JPEG2000GrayImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /AntiAliasMonoImages false /DownsampleMonoImages true /MonoImageDownsampleType /Bicubic /MonoImageResolution 1200 /MonoImageDepth -1 /MonoImageDownsampleThreshold 1.50000 /EncodeMonoImages true /MonoImageFilter /CCITTFaxEncode /MonoImageDict << /K -1 >> /AllowPSXObjects false /PDFX1aCheck false /PDFX3Check false /PDFXCompliantPDFOnly false /PDFXNoTrimBoxError true /PDFXTrimBoxToMediaBoxOffset [ 0.00000 0.00000 0.00000 0.00000 ] /PDFXSetBleedBoxToMediaBox true /PDFXBleedBoxToTrimBoxOffset [ 0.00000 0.00000 0.00000 0.00000 ] /PDFXOutputIntentProfile () /PDFXOutputCondition () /PDFXRegistryName (http://www.color.org) /PDFXTrapped /Unknown /Description << /ENU (Use these settings to create PDF documents with higher image resolution for high quality pre-press printing. The PDF documents can be opened with Acrobat and Reader 5.0 and later. These settings require font embedding.) /JPN /FRA /DEU /PTB /DAN /NLD /ESP /SUO /ITA /NOR /SVE >> >> setdistillerparams << /HWResolution [2400 2400] /PageSize [612.000 792.000] >> setpagedevice