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A checklist of the non-acarine arachnids (Chelicerata: Arachnida) of the Ndumo Game Reserve, Maputaland, South Africa C. R. Haddad, a. S. dippenaaR-SCHoeman and W. WesołoWska Haddad, C.R., A.S. Dippenaar-Schoeman and W. Wesołowska. 2006. A checklist of the non-acarine arachnids (Chelicerata: Arachnida) of the Ndumo Game Reserve, Maputaland, South Africa. Koedoe 49(2): 1–22. Pretoria. ISSN 0075-6458. Arachnids (Chelicerata: Arachnida) were collected in the Ndumo Game Reserve (Maputaland, South Africa) during 11 collecting trips in the period 2000–2006. Sampling was undertaken by various methods in eight broad habitat types: Acacia tortilis savanna; Acacia xanthophloea (fever tree) forests; deciduous broadleaf woodland; Ficus (wild fig tree) forests; floodplain vegetation; riparian forest; sand forest; and subtropical bush. In total, 457 species of arachnids were collected, representing six orders, 59 families and 240 determined genera. The most diverse order was the Araneae (46 families, 431 spp.), followed by the Pseudoscorpiones (6 families, 12 spp.), Scorpiones (3 families, 8 spp.), Opiliones (2 families, 3 spp.), Solifugae (1 family, 2 spp.) and Amblypygi (a single species). The most diverse families all belonged to the Araneae: Salticidae (82 spp.), Thomisidae (56 spp.) and Araneidae (38 spp.). The spider diversity is the highest recorded from any protected area in South Africa so far, and represents approximately 22 % of the country’s spider fauna. The habitat and guild associations of each species are provided. Key words: Arachnida, Araneae, conservation, diversity, habitats, Ndumo Game Reserve, South African National Survey of Arachnida (SANSA). C. R. Haddad, Department of Zoology & Entomology, University of the Free State, P.O. Box 339, Bloemfontein 9300, South Africa; A. S. Dippenaar-Schoeman, ARC – Plant Protection Research Institute, Private Bag X134, Queenswood 0121, South Africa/ Department of Zoology & Entomology, Faculty of Natural and Agricultural Sciences, University of Pretoria, Pretoria, 0002 South Africa; W. Wesołowska, Zoological Insti- tute, Wrocław University, Sienkiewicza 21, PL-50335, Wrocław, Poland. Introduction potential for invertebrate research in this part of southern Africa. The Ndumo Game Reserve (NGR) is situ- ated in the western parts of the Maputaland Relative to other parts of South Africa, few bioregion in northern KwaZulu-Natal, South surveys have been conducted on the arachnid Africa (Van Wyk & Smith 2001). Despite fauna of the KwaZulu-Natal Province, and many distribution records are limited to spe- its relatively small size it is one of South cies descriptions (Lawrence 1937a, 1937b, Africa’s most attractive reserves, forming a 1938, 1942a, 1942b, 1952). Lawrence et RAMSAR site protecting several wetland al. (1980) studied the arachnid fauna of and riparian ecosystems (Ramsar 2001). At Maputaland, with the emphasis on the fauna NGR, a combination of subtropical climate of the eastern shore of Lake Sibaya. Van der and rich habitat diversity provide conditions Merwe et al. (1996) investigated spiders in conducive for impressive species diversity. indigenous forests and pine plantations at the Although several vertebrate and plant taxa Ngome State Forest, while Dippenaar-Schoe- have been well studied (e.g. Pooley 1965; man & Wassenaar (2002, in press) studied Dixon 1966; De Moor et al. 1977), the diver- the fauna of rehabilitated coastal dune forests sity of invertebrates is poorly known (Had- near Richards Bay. Most recently, Honiball dad 2003), and consequently, there is great et al. (in prep.) examined diversity patterns IISSSSNN 00007755--66445588  KKooeeddooee 4499//22 ((22000066)) haddad.indd  2006/10/15 10:54:06 PM Fig. 1. Location of the Ndumo Game Reserve in northern KwaZulu-Natal, South Africa, as indicated by the arrow of spiders in the Tembe Elephant Park and orders (excluding Acari) were collected, the surrounding areas. Consequently, large gaps primary focus of this study was on the spi- exist in the knowledge of spider diversity of ders (Araneae), which was done with the Maputaland. The question raised is whether, aim of providing a baseline for future eco- for this group of invertebrates, Maputaland logical research. This study forms part of the represents a centre of endemism, as is the SANSA projects in conservancies and the case for other biological taxa such as plants Savanna Biome. (Matthews et al. 2001; Van Wyk & Smith 200). Study area Although there are a limited number of pub- lished papers dealing specifically with spider The Ndumo Game Reserve (NGR) is situ- diversity in the undisturbed conservation ated in Maputaland, a geographical biore- areas of South Africa, surveys have steadily gion covering the northern-most parts of the increased during the last decade, largely KwaZulu-Natal Province, South Africa, and due to the initiation of the South African the southern parts of Mozambique (Fig. 1). National Survey of Arachnida (SANSA). The game reserve borders on Mozambique This initiative aims, in part, to provide bio- to the north (Usutu [Usuthu] River), and lies diversity information on arachnids in South close to South Africa’s border with Swazi- Africa’s conservancies, identify biodiversity land to the west. The NGR covers an area hotspots, and aid conservation planning. As of 10 117 ha, and falls within the Bushveld- a consequence, several papers have now Savanna ecozone of southern Africa (Grant been published (Dippenaar-Schoeman 1988, & Thomas 1998). 2006; Dippenaar-Schoeman & Gonzalez- A rich habitat variety can be found at NGR Reyes 2006; Dippenaar-Schoeman & Leroy (De Moor et al. 1977), but for the pur- 2003; Dippenaar-Schoeman et al. 989, poses of this study they are grouped into 1999, 2005; Foord et al. 2002; Whitmore et eight broad habitat types, which are briefly al. 200). described here. Classification of plants fol- The relatively poor knowledge of the arach- lows Germishuizen et al. (2006): nids of Maputaland provided the impetus to . Acacia tortilis (Forssk.) Hayne (umbrella set up a checklist for Ndumo Game Reserve. thorn) savanna (AS)—situated in the Although specimens of all of the arachnid south-western parts of the reserve. This Koedoe 49/2 (2006) 2 ISSN 0075-6458 haddad.indd 2 2006/10/15 10:54:06 PM habitat type is dominated by A. tortilis, neglecta I.Verd., Haplocoelum foliosum Albizia anthelmintica (A.Rich.) Brongn. (Hiern) Bullock, Hymenocardia ulmoides and A. petersiana (Bolle) Oliv. trees, Oliv., Phyllanthus sp. and Spirostachys and has a well-developed grass layer africana Sond. The ground is gener- (Figs 2–3), including Aristida adscen- ally devoid of any grass, and short plants sionis L., Chloris pycnothrix Trin., C. (Asparagus africanus Lam., A. falcatus L. virgata Sw., Enteropogon macrostachyus and Sansevieria hyacinthoides (L.) Druce, (Vahl) K.Schum., Hyperthelia dissoluta amongst others) occur sporadically in the (Nees ex Steud.) Clayton, Setaria sp., habitat (Fig. 5). Sporobolus fimbriatus (Trin.) Nees and 4. Subtropical bush (ST), which covers the Themeda triandra Forssk. largest area of the reserve, is a hetero- 2. Deciduous broadleaf woodland (BW) geneous habitat composed largely of —situated in the southern, higher lying Mahemene thicket and Acacia nigrescens parts of the reserve. This is a highly Oliv. woodland (De Moor et al. 977). diversified savanna habitat, with extreme Although other habitat types can be variations in plant composition and veg- included under subtropical bush, samp- etative density (Fig. 4). Common trees ling was primarily carried out in these include Acacia burkei Benth., Albizia two habitats. Dominant trees include var- adianthifolia (Schumach.) W.Wight, A. ious Acacia spp. (including A. burkei, A. petersiana, Combretum molle R.Br. ex karroo Hayne and A. nigrescens), Albiz- G.Don, Dichrostachys cinerea (L.) Wight ia species (including A. anthelmintica, & Arn., Grewia occidentalis L., Sclero- A. petersiana and A. versicolor Welw. carya birrea (A.Rich.) Hochst., Strych- ex Oliv.), Apodytes dimidiata E.Mey. nos madagascariensis Poir., Terminalia ex Arn., Balanites maughamii Sprague, sericea Burch. ex DC., Trichilia emetica Euclea species (E. crispa (Thunb.) Gürke, Vahl and Ziziphus mucronata Willd. Vari- E. daphnoides Hiern and E. divinorum ous short shrubs, including Chionanthus Hiern), Euphorbia ingens E.Mey. ex peglerae (C.H.Wright) Stearn, Maytenus Boiss., Ptaeroxylon obliquum (Thunb.) sp., Olinia sp. and Teclea natalensis Radlk., Trema orientalis (L.) Blume and (Sond.) Engl., dominate the lower strata Vepris lanceolata (Lam.) G.Don. Shrubs of the habitat. Common grasses include and shorter plants include Carissa bispi- Aristida adscensionis, Cymbopogon nosa, Croton menyharthii Pax, Euphor- nardus (L.) Rendle, Cynodon dactylon, bia grandicornis Goebel ex N.E.Br., Dactyloctenium spp., Digitaria erian- Hypericum sp., Gardenia spp., Grewia tha Steud., various Eragrostis spp. (E. bicolor Juss., Sansevieria hyacinthoides lehmanniana Nees, E. rigidior Pilg. and and Tarchonanthus camphoratus L. The E. superba Peyr.), Heteropogon contor- grass and herb layer is highly variable, tus (L.) Roem. & Schult., Hyperthelia in some areas consisting primarily of dissoluta (Nees ex Steud.) Clayton, Pani- the creepers Cissus rotundifolia (Forssk.) cum maximum Jacq., P. repens L. and Vahl and Crassula sp. (Acacia nigrescens Setaria species. woodland), while other areas, including 3. Sand forest (SF)—situated in the south Mahemane thicket, consist primarily of east of the reserve. Sand forest is more short grasses such as Chloris spp., Dac- typical of the Tembe Elephant Park (Mat- tyloctenium spp., Eragrostis spp. and thews et al. 2001) and is a habitat unique Setaria verticillata (L.) P.Beauv., with to Maputaland. Only a small patch is Cissus rotundifolia creepers found near found in Ndumo at the bottom of Ndumo the base of trees (Figs 6–7). hill along the southern boundary fence. 5. Acacia xanthophloea Benth. (fever tree) Common trees and shrubs include Art- forests (AX)—situated around the mar- abotrys monteiroae Oliv., Carissa bispi- gins of the freshwater pans Banzi, Hotwe, nosa (L.) Desf. ex Brenan, Commiophora Nyamiti [Nyamithi] and Shokwe, in rela- ISSN 0075-6458 3 Koedoe 49/2 (2006) haddad.indd 3 2006/10/15 10:54:07 PM 2 3 4 5 6 7 Figs. 2–7. Savanna and associated habitats in Ndumo Game Reserve: 2–3) Acacia tortilis savanna (AS) domi- nated by Acacia tortilis (2) and Albizia spp. (3) trees; 4) Deciduous broadleaf woodland (BW); 5) Sand forest (SF); 6–7) Subtropical bush (SB) consisting of Mahemane thicket (6) and Acacia nigrescens woodland (7). tively localised areas. This habitat type 6. Floodplain (FP)—situated inland from is strongly dominated by Acacia xantho- riparian forests along the Pongola phloea trees, with shorter shrubs (e.g. [Phongolo] and Usutu Rivers. This habi- Carissa and Grewia spp.). Weeds such tat is dominated by the reed species as Solanum spp. occur sporadically (Figs Phragmites australis (Cav.) Steud. and P. 8–9). Cynodon dactylon (L.) Pers. is mauritianus Kunth, and various shorter the dominant grass species, although the grasses and sedges, including Cynodon taller Cyperus sp. and Leptochloa fusca dactylon, Cyperus fastigiatus Rottb. and (L.) Kunth are also present in places. In Echinochloa pyramidalis (Lam.) Hitchc. some areas (e.g. some sites at Banzi Pan) & Chase (Figs 10–11). Trees and shrubs grasses were entirely absent. occur sporadically along the floodplain, Koedoe 49/2 (2006) 4 ISSN 0075-6458 haddad.indd 4 2006/10/15 10:54:10 PM 8 9 10 11 12 13 Figs. 8–13. Floodplain and associated habitats in Ndumo Game Reserve: 8–9) Fever tree or Acacia xantho- phloea forests at Shokwe pan (8) and Banzi pan (9). Note the differences in understory plant growth; 10–11) Floodplain vegetation (FP); 12) Wild fig or Ficus forest (FF) at Shokwe pan; 13) Riparian forest (RF) at the Pongola River. and include Acacia xanthophloea, Ficus fig trees (particularly F. sycomorus and sycomorus L., Grewia caffra Meisn., F. sur Forssk.), and is characterised by a Gymnosporia senegalensis (Lam.) Loes. near absence of shrubs and grasses, and and Kigelia africana (Lam.) Benth., and presence of a very well developed leaf lit- are often covered near their trunks by ter layer (Fig. 12). Other trees that occur Monanthotaxis caffra (Sond.) Verdc. sporadically in the habitat include A. creepers. xanthophloea, Breonadia salicina (Vahl) 7. Ficus (wild fig tree) forests (FF)—situ- Hepper & J.R.I.Wood, K. africana and ated around the margins of the freshwater Trichilia emetica. pans Shokwe and Banzi, and along riv- 8. Riparian forest (RF)—situated along the ers. This habitat is dominated by wild banks of the Usutu and Pongola rivers ISSN 0075-6458 5 Koedoe 49/2 (2006) haddad.indd 5 2006/10/15 10:54:12 PM ( F i g . 1 3 ) . T h is habitat is dominated by tall and by hand (grass); web-builders were collected by trees including B. salicina, F. sur, F. syco- hand and sweep-nets; spiders were collected on or morus, Harpephyllum caffrum Bernh., under bark with a pooter, using a modified net, or by hand. Night collecting was done by hand with the Olea capensis L. and Rauvolfia caffra aid of a flashlight. Spiders attracted to light traps at Sond., with M. caffra creepers frequently night were also collected. Specimens were preserved growing between trees. Shrubs present in 70 % ethanol for each site sampled. include Antidesma venosum E.Mey. ex The second author identified most of the mate- Tul., Grewia caffra, Tarenna pavettoi- rial to species level, while the third author identified des (Harv.) Sim. and Tricalysia species. the Salticidae. Various taxa were identified by the A dense leaf litter layer is present, and specialists listed in the Acknowledgments. Voucher grasses (e.g. Cynodon dactylon, Hemar- specimens have been deposited in the following thria altissima (Poir.) Stapf & C.E.Hubb. institutions: Royal Museum of Central Africa, Ter- and Ischaemum spp.) only occur very vuren, Belgium (Linyphiidae, Salticidae and Zodari- idae); National Museum, Bloemfontein, South Africa sporadically. (Miturgidae, Sicariidae and Opiliones); Western Aus- Sampling was generally limited to winter, tralian Museum, Perth, Australia (Pseudoscorpiones); when game counts (wildlife censuses) were Zoological Institute and Museum Alexander Koenig, Bonn, Germany (Pholcidae); American Museum of conducted, and mid-summer. Arachnids were Natural History, New York, U.S.A. (Scorpiones); and collected during the following periods: July the National Collection of Arachnida, ARC—Plant 2000 (2 weeks); November–December 2000 Protection Research Institute, Pretoria, South Africa (4 weeks); January (1 week) and July 2002 (all other arachnids). (2 weeks); June–July 2003 (2 weeks); July 2004 (2 weeks); February (1 week) and June 2005 (2 weeks), and January 2006 (2 weeks), Guilds observed April (1 week) and June 2006 (2 weeks). Two main guilds can be distinguished among arach- nids, namely wanderers and web-builders. The wan- derers can be further divided into plant wanderers (PW) and ground wanderers (GW). Plant wanderers Methods were separated and placed in a guild based on the Sampling was conducted primarily on an ad hoc vegetation type that they were most commonly found basis, and was not quantitative. Consequently, sam- on, namely plant wanderer on foliage (PWF), plant pling intensity in each habitat varied considerably. wanderer on grass (PWG) and plant wanderer on For example, sand forest was relatively poorly sam- bark (PWB). pled, while deciduous broadleaf woodland and sub- The web-building spiders can also be subdivided into tropical bush were very well sampled. different guilds based on the structure of the webs The sampling methods for each arachnid guild can be that they build: sheet-web builders (SWB), space- summarised as follows: ground wanderers were sam- web builders (SPWB), orb-web builders (OWB), pled using pitfall traps, rock and log turning, and leaf modified orb-web builders (MOWB), funnel-web litter sifting; plant wanderers were collected using builders (FWB), retreat-web builders (RWB) and beating sheets and by hand (trees), and sweep-nets gum-foot-web builders (GWB). Table 1 Order composition of the non-acarine arachnids in the Ndumo Game Reserve, Maputaland Order Common Name Families Genera Species Amblypygi Tailless whip-scorpions    Araneae Spiders 46 222 431 Opiliones Harvestmen 2 2 3 Pseudoscorpiones False scorpions 6 8 2 Scorpiones Scorpions 3 5 8 Solifugae Sun spiders  2 2 Total 59 240 457 Koedoe 49/2 (2006) 6 ISSN 0075-6458 haddad.indd 6 2006/10/15 10:54:12 PM Table 2 Proportional diversity of the non-acarine arachnid orders in Ndumo Game Reserve relative to total South African (Dippenaar-Schoeman & Haddad, unpubl.) and global (Harvey 2002) diversity of each order Order NGR South Africa Global Species % Species % Species % Araneae 431 94.52 2000 78.03 36000 74.70 Pseudoscorpiones 2 2.63 135 5.27 3239 6.72 Scorpiones 8 .75 93 3.63 279 2.65 Opiliones 3 0.66 79 6.98 6000 2.45 Solifugae 2 0.44 50 5.85 087 2.26 Amblypygi  0.22 3 0.2 136 0.28 Others 0 0.00 3 0.2 45 0.94 457 ~00.00 2563 ~00.00 48 92 ~00.00 Specimens were identified in the laboratory and the other arachnid orders (excluding Acari) placed within a particular guild. Additional notes are under-represented relative to the global were made of the habitats in which each species was arachnid diversity of each order (Table 2; collected, and information on the habits of some spe- Harvey 2002; Dippenaar-Schoeman & Had- cies was gathered to provide a fuller picture of the dad unpubl.). The under-representation of ecology of the spiders within the reserve. some of these orders (Solifugae and Scor- piones) could be attributed to their generally higher diversity in more arid environments, Results and Discussion and lower diversity in subtropical environ- ments, in southern Africa (Lawrence 1955, Diversity 1963; Prendini 2005). In total, 457 species representing six orders, The most diverse families collected were 59 families and 240 determined genera the Salticidae (82 spp.), followed by the of arachnids were collected (Appendix 1, Thomisidae (51 spp.) and Araneidae (38 Table 1). The greatest diversity was found spp.). Eighteen families were represented in the Araneae (46 families, 431 spp.), fol- by a single species only. The total spider lowed by the Pseudoscorpiones (6 families, diversity (431 spp.) represents the highest 12 spp.), Scorpiones (3 families, 8 spp.), number of species recorded from a single Opiliones (2 families, 3 spp.) and Solifugae conservancy surveyed in South Africa thus (1 family, 2 spp.). The Amblypygi was only far. Previous surveys have yielded between represented by a single species. Compared 75 and 268 species (Table 3). It must be to the high diversity of spiders collected, noted that the current study took place over Table 3 Currently known total species diversity of spiders (Araneae) recorded from South African conservancies Conservancy Families Species Reference Karoo National Park 38 6 Dippenaar-Schoeman et al. (999) Kruger National Park 40 52 Dippenaar-Schoeman & Leroy (2003) Makalali Game Reserve 38 268 Whitmore et al. (200) Mountain Zebra National Park 34 75 Dippenaar-Schoeman (988, 2006) Ndumo Game Reserve 46 431 Present study Roodeplaat Dam Nature Reserve 27 0 Dippenaar-Schoeman et al. (989) Soutpansberg West Conservancy 46 27 Foord et al. (2002) Swartberg Nature Reserve 45 86 Dippenaar-Schoeman et al. (2005) ISSN 0075-6458 7 Koedoe 49/2 (2006) haddad.indd 7 2006/10/15 10:54:13 PM 90 80 70 60 es eci50 p of s er 40 b m u N30 20 10 0 Salticidae Thomisidae Araneidae Theridiidae Gnaphosidae Corinnidae Oxyopidae Lycosidae Pisauridae Zodariidae Sparassidae Tetragnathidae Clubionidae Philodromidae Miturgidae Eresidae Nephilidae Oonopidae Prodidomidae Uloboridae Dictynidae Idiopidae Theraphosidae Deinopidae Mimetidae Pholcidae Scytodidae Trochanteriidae Cyrtaucheniidae Ctenidae Linyphiidae Palpimanidae Segestriidae Selenopidae Web-building singletons Wandering singletons Fig. 14. Species diversity of spider families collected in Ndumo Game Reserve ranked from highest to lowest. Black bars indicate wandering spiders and grey bars indicate web-builders. a period of seven years (11 intensive sam- Common taxa by stratum pling trips). Furthermore, a wide variety of Since this study was not conducted quanti- sampling methods were used in most strata of tatively there is no specific data available on the habitats present, while many of the other the abundance of particular species in each surveys were restrictive in terms of sampling habitat. However, based on observations and intensity (often ad hoc sampling) or meth- collecting frequency during this study the ods. The survey in Roodeplaat Dam Nature following patterns emerged for each stratum Reserve only involved the field and tree layer and guild: and did not include ground-dwelling arach- Ground wanderers: A large number of spe- nids (Dippenaar-Schoeman et al. 989). cies (Appendix 1) are wandering arachnids on the ground surface. The community com- Guilds position varied greatly between different habitats. Open savanna habitats (AS, BW) The vast majority of the arachnids collected were dominated by various gnaphosid genera were wanderers (75.9 %), with web-builders (Asemesthes, Zelotes, Setaphis and Camilli- comprising 24.1 % of the species. Amongst na), lycosids (Geolycosa sp. and Evippomma spiders only, wanderers contributed a much squamulatum (Simon)), corinnids (Merenius higher proportion of the species (Fig. 14), alberti Lessert, Copa flavoplumosa Simon with two families (Salticidae and Thomis- and Castianeira sp.), salticids (Mexcala ele- idae) contributing more than 32 % of the gans Peckham & Peckham, Stenaelurillus total spider species. Two of the diverse web- and Phlegra spp.) and zodariids (Caesetius building families, Araneidae and Theridiidae, bevisi (Hewitt), Capheris sp., Palfuria sp. contributed 15.5 % of the spider species, and Psammorygma sp.), while Uroplectes combined. olivaceus Pocock (Buthidae), Opistacan- Koedoe 49/2 (2006) 8 ISSN 0075-6458 haddad.indd 8 2006/10/15 10:54:13 PM thus spp. (Liochelidae), Brachionopus sp. sampled was AX, with a survey having (Theraphosidae), Scytodes caffra Lawrence been conducted specifically on the arach- (Scytodidae), Loxoscelis spinulosa Purcell nids associated with Acacia xanthophloea (Sicariidae) and Caponia chelifera Lessert bark at five pan and floodplain sites in the (Caponiidae) were most common under logs reserve (Haddad unpubl.). The most common and rocks. associations with this tree species are Ceto- nana spp. (Corinnidae), Pseudicius spp. and The fauna of floodplain habitats (AX, FF, Heliophanus spp. (Salticidae), various gna- FP) was strongly dominated by wolf spiders phosids, miturgids (Cheiracanthium spp. and (Lycosidae), gnaphosids (particularly Zelotes Cheiramiona spp.), Hersilia sericea Pocock spp.), corinnids (particularly M. alberti) and (Hersiliidae), Platyoides spp. (Trochanteri- salticids (Thyenula spp. and Evarcha spp.). idae) and Clubiona spp. (Clubionidae). In Metabiantes maximus Lawrence (Biantidae) the Ficus forest (FF) and floodplain (FP) and Scytodes spp. (Scytodidae) were com- a similar fauna was found, but densities mon under logs in these habitats. were generally lower, probably due to the The remaining two forest types studied dif- more variable bark structure. Pseudoscorpi- fered considerably in their faunas. Riparian ons, particularly atemnids (e.g. Titanatemnus forest was dominated by Zelotes spp., Mere- natalensis Beier), and buthid scorpions (par- nius alberti, Copa flavoplumosa, Thyenula ticularly Uroplectes spp. and Pseudolycus spp., and various lycosids and oonopids. Spi- pegleri (Purcell)) were also very common der densities also appear to be much higher under the bark of trees in these habitats. than in sand forest. The latter habitat was The arachnids associated with tree bark in dominated by lycosid spiders, with oonopids RF and SF differed considerably. In RF, the and corinnids in lower densities. fauna contained elements of Ficus forest, Subtropical bush had relatively similar fau- which can be explained by the presence of nas. Mahemane thicket areas were domi- Ficus spp. in this habitat. In SF, jumping nated by salticids (Stenaelurillus spp. and spiders (particularly Holcolaetis zuluensis Phlegra spp., and Mexcala elegans), lyco- Lawrence and Pseudicius spp.) were most sids, oonopids and gnaphosids of the genera common, with lesser numbers of Hersilia Asemesthes, Camillina, Setaphis and Zelotes, sericea and Clubiona spp. present. while Acacia nigrescens woodland areas were dominated by lycosids, salticids and Trees in AS, BW and ST generally had a liochelid scorpions. coarser bark structure compared to the afore- mentioned habitats. Wandering spiders were Ground web-builders: Web-builders near the largely limited to H. sericea, Clubiona spp., ground surface were generally uncommon, Philodromus spp. (Philodromidae) and vari- and were largely limited to amaurobiids, ous salticids. Euprosthenops vuattouxi Blandin (Pisauri- dae), Xeviosa amica Griswold (Phyxelidi- Web-builders associated with bark: As for dae), Ariadna spp. (Segestriidae) and Hip- wanderers, AX also had a high diversity and pasa australis Lawrence (Lycosidae), which abundance of web-builders compared to other were found in contrasting habitats (Appen- habitats. The most widespread and common dix 1). The latter species was the only species species associated with bark in the eight that was widespread in NGR, occurring in all habitats include various theridiids (Theridion of the habitats sampled. Linyphiid and hah- spp., Euryopis sp. and Argyrodes spp.), ara- niid sheet-web builders were most common neids (particularly Neoscona spp.), and on in ST leaf litter. occasion Nephilengys cruentata (Fabricius) (Nephilidae). Wanderers associated with bark: Due to the variable plant composition of the dif- Wanderers on grass: Grasses were particu- ferent habitats, arachnids associated with larly widespread in AS and BW, where a bark varied considerably. The habitat best large proportion of sampling was undertak- ISSN 0075-6458 9 Koedoe 49/2 (2006) haddad.indd 9 2006/10/15 10:54:13 PM e n . C o m m o n w anderers in these two habitats while being less frequently encountered in include various salticids (particularly Thyene, the other habitats. Viciria and Heliophanus spp.), Philodromus Web-builders on foliage: The web-building and Tibellus spp. (Philodromidae), Oxyopes arachnids showed greater habitat variation spp. (Oxyopidae) and Thomisus, Runcinia than the wanderers, which could be related and Synema spp. (Thomisidae). to spacing between trees and shrubs in which In the two floodplain habitats (AX and FP) webs can be constructed. The savanna and the grass fauna was dominated by immature woodland habitats (AS, BW and ST) had lycosids, Heliophanus spp., Oxyopes spp., higher densities of large and colourful ara- Charminus spp. and Afropisaura rothiformis neids (Gasteracantha milvoides Butler, Cyr- (Simon) (Pisauridae), and Thomisus spe- tophora citricola (Forskål), Caerostris and cies. The grass fauna of ST was dominated Isoxya spp.) and the nephilid Nephila sen- by Charminus and Oxyopes spp., with other egalensis (Walckenaer) than the other habi- common taxa including Synema, Thomisus, tats in the reserve. In contrast, riverine and Thyene and Hyllus species. Grasses were floodplain habitats had high densities of G. largely or entirely absent from FS, RF and milvoides, N. fenestrata Thorell and N. inau- SF, and consequently little data could be gen- rata (Walckenaer). The SF fauna comprised erated for these habitats. largely of small araneid spiders, particularly Araniella and Neoscona. Theridiid, dictynid Web-builders in grass: The AS and BW and araneid spiders were common on small grasslands were a rich source of web-build- shrubs and trees at AX, FF, FP and RF, while ing spiders, particularly for the families Ara- araneid, theridiid and uloborid spiders were neidae (Neoscona, Hypsosinga, Isoxya and common in AS, BW and ST. Singa spp.), Dictynidae (Archaeodictyna and Dictyna spp.) and Theridiidae (Latrodectus geometricus C. L. Koch and Theridion spp.). Conservation implications Most of these species construct a retreat or The high species diversity of arachnids, and web in the florescences of various grasses. spiders in particular, as a single taxon group In the floodplain habitats AX and FP, web- suggests that other invertebrate taxa may builders primarily included various Tet- similarly display high levels of diversity ragnatha spp. (Tetragnathidae), Neoscona, in Ndumo Game Reserve. Approximately Argiope and Araniella spp. (Araneidae) and 18 % and 22 % of the total South African small theridiids. Web-builders in ST grasses non-acarine arachnid, and spider diversity, were generally uncommon due to the short respectively (Table 2), is currently conserved grasses present, and consisted mainly of in this reserve. Thus, further study on the immature araneids and theridiids. invertebrates is needed to discover the extent of the richness of the Maputaland fauna. The Wanderers on foliage: All of the habitats high diversity of a single taxon should serve have a well-developed tree layer and there as strong supportive evidence of the need to was generally a large degree of overlap in the create awareness of the plight of invertebrate faunas between habitats. Generally, the most conservation, and increase the area under common wandering spiders throughout NGR conservation in this part of South Africa. include many of the Thomisidae (particularly Thomisus, Monaeses, Thomisops and Synema Species of notable conservation importance spp., and Misumenops rubrodecoratus Mil- include the four baboon spiders (Theraphosi- lot), Salticidae (especially Hyllus, Evarcha, dae) collected. Ceratogyrus bechuanicus Pur- Heliophanus, Thyene and Pseudicius spp.), cell is already considered to be a threatened Oxyopidae (Oxyopes and Hamataliwa spp.), species due to the pet trade (De Wet & Dip- Clubiona spp. and Miturgidae (especially penaar-Schoeman 1991). The Brachionopus Cheiramiona spp.). In particular, Philodromi- sp. is unlikely to be seriously threatened, and dae were very common on thorn trees in AS, was encountered very frequently under logs Koedoe 49/2 (2006) 0 ISSN 0075-6458 haddad.indd 0 2006/10/15 10:54:13 PM

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logical research and Tarchonanthus camphoratus L. The .. Guilds. The vast majority of the arachnids collected were wanderers (75.9 %), with
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