Abstract
The knowledge of spiders in the Eastern Cape province lags behind that of most other South African provinces. The Eastern Cape province is renowned for its conservation areas, as the largest part of the Albany Centre of Endemism falls within this province. This article provides a checklist for the spider fauna of the Addo Elephant National Park, one of the most prominent conservation areas of the Eastern Cape, to detail the species found in the park and determine their conservation status and level of endemicity based on their known distribution. Various collecting methods were used to sample spiders between 1974 and 2016. Forty-seven families that include 184 genera and 276 species were recorded. Thomisidae (39 spp.), Araneidae (39 spp.), Salticidae (35 spp.) and Theridiidae (25 spp.) were the most species-rich families, while 14 families were only represented by a single species.
Conservation implications: A total of 12.7% of the South African spider fauna and 32.9% of the Eastern Cape fauna are protected in the park; 26.4% are South African endemics, and of these, 3.6% are Eastern Cape endemics. Approximately, 4% of the species are possibly new to science, and 240 species are recorded from the park for the first time.
Keywords: Arachnida; conservation; endemism; biome; records.
Introduction
One of the core research areas of the South African National Survey of Arachnida (SANSA) is to determine the number of arachnid species presently conserved in protected areas, including the South African National Parks (SANParks) (Dippenaar-Schoeman & Haddad 2006; Dippenaar-Schoeman 2016). The species distribution data generated through SANParks and other surveys feeds into the conservation assessments used to compile the Red Data List of the Arachnida of South Africa (Lyle & Dippenaar-Schoeman 2015).
Compared to some of the better surveyed provinces of South Africa, for example, KwaZulu-Natal and Limpopo, the Eastern Cape has received far less attention, but harbours relatively more endemic taxa than the Limpopo province, particularly as a function of its size (Foord et al. 2011). This serves as a possible indicator of the unique spider assemblages of the region. Although the Addo Elephant National Park (AENP) is the largest national park in the Eastern Cape and the third largest national park in South Africa, no documented surveys of spiders exist for this protected area.
Work in the province has largely been limited to the Mountain Zebra National Park, the first national park for which a spider species list was published (Dippenaar-Schoeman 1988, 2006), and a study on the subsocial eresid Stegodyphus tentoriicola Purcell, 1904 (Ruch et al. 2009a, 2009b, 2012). Further work in the Eastern Cape includes aspects of the spider fauna in the Mkambati Nature Reserve (Dippenaar-Schoeman, Hamer & Haddad 2011; Hamer & Slotow 2017) and the Silaka Nature Reserve (Forbanka & Niba 2013), while surveys in the Asante Sana Nature Reserve were the focus of a doctoral study on epigaeic invertebrates (Midgley 2012).
The only records of spiders from the AENP (30 spp.) were included in the First Atlas of South African Spiders (Dippenaar-Schoeman et al. 2010), and six species have been referred to in taxonomic papers in the past (Dippenaar-Schoeman 1980; Jocqué 1984; Lyle & Haddad 2018; Wesołowska & Haddad 2013, 2018). Here, we document all spider species recorded from the AENP over the last 50 years, with efforts being concentrated over the last decade. Information is provided on the global distribution, endemicity and conservation status of each species.
Research method and design
Study area and period
The AENP is situated in the Sundays River region, between the city of Port Elizabeth and the town of Makhanda (Grahamstown) (Figure 1). Although originally proclaimed in 1931 to protect one of the four elephant populations that survived the start of the 20th century in South Africa, it is very significant in the conservation of the Albany thicket, which is predominantly found in the Eastern Cape (Johnson, Cowling & Phillipson 1999) and is the dominant biome of the region (Mucina & Rutherford 2006). The Woody Cape Nature Reserve and some portions of privately owned land were recently added to the park, bringing its total area to approximately 179 000 hectares (SANParks 2015).
|
FIGURE 1: Map of South Africa, showing the locality of the Addo Elephant National Park in the Eastern Cape province (inset) and the different sites where spiders were collected. |
|
Rainfall in the park is mainly concentrated in spring and autumn, varying between 350 mm and 900 mm per annum, depending on topography and biomes (SANParks 2015). The region seldom experiences temperatures below 0 °C, while summer extremes of 45 °C are quite common (Lombard et al. 2001).
Sampling methods and identification
The first spider survey in the AENP took place in 1974, when the members of the Agricultural Research Council (ARC) – Plant Protection Research Institute (PPRI) (now ARC – Plant Health and Protection) sampled there. In 2009, a second survey was undertaken by the members of the ARC-PPRI. Between these events, there was occasional collecting by various museum researchers and members of the public who deposited specimens in several museums in South Africa and internationally. In 2010, more regular sampling started in the park after a formal research agreement was established between SANSA and SANParks, to stimulate more intensive spider sampling in the national parks. As a result, more regular sampling was undertaken in five different parts of the park (Figure 2), which included the semi-arid Sundays River succulent thicket around the Darlington Dam; the fynbos and Afromontane forests of the Zuurberg Mountains; the subtropical thickets in the Kabouga area just north of Kirkwood; the Alexandria coastal forest and dune thickets of the Woody Cape area; and the Albany thickets of the main camp in the Sundays River Valley (Figure 2).
|
FIGURE 2: Habitat types in the Addo Elephant National Park: (a) Darlington Dam (Succulent Karoo thicket); (b) Main Camp (Albany thickets); (c) and (d) Kabouga (subtropical thicket); (e) and (f) Woody Cape (coastal forest and dune thicket); (g) and (h) Zuurberg (fynbos and Afromontane forest). |
|
Conventional sampling techniques, as described by Dippenaar-Schoeman et al. (2015), were used. This included sweepnet sampling, beating, leaf-litter sifting and active searches during the day and night. Sampling did not follow a standardised protocol but was undertaken sporadically and ad hoc, depending on locality and time constraints. Voucher specimens were predominantly deposited in the National Collection of Arachnida (NCA) at the ARC–Plant Health and Protection in Pretoria. Taxonomic constraints, juveniles and undescribed species necessitated the use of morphospecies in some instances (Appendix 1).
Endemicity value and conservation status
A detailed explanation of the indices to assess the endemicity value and conservation status of each species can be found at the end of Appendix 1, but it distinguishes between species only known from the type locality (6), the Eastern Cape (5), two adjoining provinces (4), South Africa (3), Southern Africa (2), Afrotropical Region (1) and finally areas outside the Afrotropical Region (0). The conservation status was derived from a recent National Red List assessment of spiders in South Africa.
Ethical considerations
This research followed all ethical standards for research without direct contact with human or animal subjects.
Results and discussion
Numbers present
In total, 276 species in 184 genera and 47 families were recorded (Table 1; Appendix 1). Seven species are probably new and 16 species are undetermined, because of the lack of resolved taxonomy of their genera (Appendix 1). Possible new species belong to large families that require comprehensive revision to determine the taxonomic status and identity of these taxa.
TABLE 1: Spider diversity of the Addo Elephant National Park, with the total number of families, genera and species sampled. |
Family diversity
Of the 47 spider families collected from AENP (Table 1; Appendix 1), the Thomisidae and Araneidae with 39 spp. are the most species-rich, followed by the Salticidae (35 spp.) and Theridiidae (25 spp.). For comparison, at the Mkambati Nature Reserve, the Theridiidae and Araneidae were the most species-rich families (Dippenaar-Schoeman et al. 2011), and at the Mountain Zebra National Park, it was the Thomisidae (Dippenaar-Schoeman 1988, 2006).
Thomisidae: Thomisids are mainly plant dwellers and are easily dispersed by wind. Consequently, most species have a wide distribution. Only five species were previously reported from AENP in the Spider Atlas (Dippenaar-Schoeman et al. 2010); therefore, 34 spp. are newly recorded from the park. Twenty-one species have a wide distribution throughout Africa, while four species are also known from Europe and nine occur throughout southern Africa. Four species are South African endemics, with two – Holopelus almiae Dippenaar-Schoeman, 1986 and Mystaria lindaicapensis Lewis & Dippenaar-Schoeman, 2014 (Figure 3l) – being near endemics of the Eastern Cape. Both species are also known from the Western Cape. The species listed as Monaeses sp. 2 is new to science, and the same species was also recorded at the Mkambati Nature Reserve (Dippenaar-Schoeman et al. 2011). Diaea dorsata (Fabricius, 1777) (Figure 3k) is known from Europe to Japan (World Spider Catalog 2019) and is reported from South Africa for the first time.
|
FIGURE 3: Selected spiders of the Addo Elephant National Park: (a) Araneus detrimentosus (O. Pickard-Cambridge, 1889) (Araneidae); (b) Undescribed Cyclosa sp. (Araneidae); (c) Ideocaira transversa Simon, 1903 (Araneidae); (d) Zygiella sp. (Araneidae); (e) Ancylotrypa cornuta Purcell, 1904 (Cyrtaucheniidae); (f) Allothele australis (Purcell, 1903) (Dipluridae); (g) Moggridgea rupicoloides Hewitt, 1914 (Migidae); (h) Hermacha grahami (Hewitt, 1915) (Nemesiidae); (i) New genus (Philodromidae); (j) Harpactirella magna Purcell, 1903 (Theraphosidae); (k) Diaea dorsata (Fabricius, 1777) (Thomisidae); (l) Mystaria lindaicapensis Lewis & Dippenaar-Schoeman, 2014 (Thomisidae). |
|
Araneidae: The Araneidae are web builders and produce typical orb webs. Of the 39 species recorded, only four were previously known from the park (Dippenaar-Schoeman et al. 2010). Twenty of the species have a wide African distribution, three species are endemic to southern Africa, and only three are South African endemics that include Ideocaira transversa Simon, 1903 (Figure 3c), Nemoscolus elongatus Lawrence, 1947 and Ursa turbinata Simon, 1895. Three genera, Acanthepeira, Larinioides and Zygiella (Figure 3d), represented by undetermined species, are newly recorded for South Africa, as well as the species Araneus detrimentosus (O. Pickard-Cambridge, 1889) (Figure 3a) and Chorizopesoides orientalis (Simon, 1909). Species of six genera could not be determined (e.g. Figure 3b) and might represent new records.
Salticidae: The Salticidae are free-living spiders found on vegetation and the soil surface. Only Cyrba nigrimana Simon, 1900 (Dippenaar-Schoeman et al. 2010), Heliophanus (Heliophanus) gramineus Wesołowska & Haddad, 2013, Massagris mirifica Peckham & Peckham, 1903 (Wesołowska & Haddad 2018) and Pseudicius maculatus Haddad & Wesołowska, 2011 (Wesołowska & Haddad 2013) were previously recorded from the park. Eighteen species are widely known throughout Africa, and one, Menemerus bivittatus (Dufour, 1831), has a pantropical distribution. Six species are endemic to southern Africa, and 10 species are South African endemics; five species are near endemic to the Eastern Cape. Only one Tanzania species is possibly new to science.
Theridiidae: The theridiids construct different types of webs known as cobwebs or gumfoot webs. No theridiids were previously reported from the park. Unfortunately, the taxonomy of most theridiid genera in Africa is still unresolved. Seven species (28%) could not be identified to species level. This corresponds with the Mkambati Nature Reserve where 67% of the species were unresolved (Dippenaar-Schoeman et al. 2011). Eight species have a wide global distribution and five species are South African endemics, with only one species (Phoroncidia capensis [Simon, 1895]) being endemic to the Eastern Cape. Some of the undetermined species might be new to science.
Species endemicity and conservation status
Of the 276 species sampled, 17 spp. (6.2%) are data deficient (DD) and lack taxonomic or distribution data, while 31 spp. (11.2%) were not evaluated (NE) (Table 2; Appendix 1). The majority of the species (227 spp., 82.2%) sampled are listed as least concern (LC). Ninety-seven of the species (35.1%) are found throughout Africa, while 50 species (18.1%) are endemic to southern Africa. One species, Mystaria lindaicapensis, is considered to be vulnerable, because of its restricted distribution in the Eastern and Western Cape provinces (Table 3).
TABLE 2: Conservation status and endemicity of the spider species sampled at the Addo Elephant National Park. |
TABLE 3: Species of special concern recorded from the Addo Elephant National Park. |
Of the species identified to species level, only 73 spp. (26.4%) are South African endemics (Table 2; Appendix 1). There are presently no endemic species known only from the AENP, but 10 species from eight families are Eastern Cape endemics (ECE) and a further species is near endemic. These species are of special concern because of their small distribution ranges (Table 3), and need more sampling and/or taxonomic study to improve knowledge of their distributions. The 10 ECE are listed as data deficient, of which only Ancylotrypa cornuta Purcell, 1904 (Figure 3e) is known from both sexes. The VU near-endemic species, M. lindaicapensis, is only known from three south coast localities, including AENP. The ECE species include three trapdoor spider species (A. cornuta, Figure 3e; Hermacha grahami [Hewitt, 1915], Figure 2e; Moggridgea rupicoloides Hewitt, 1914, Figure 3g) and two baboon spider species (Harpactira curvipes Pocock, 1897; Harpactirella magna Purcell, 1903, Figure 3j).
Twenty-five of the AENP spiders have a wide global distribution (Table 2; Appendix 1), and most of them have been recorded from several continents (World Spider Catalog 2019). Some of these species listed from AENP have not been recorded from South Africa before, including Araneus detrimentosus, Chorizopesoides orientalis and Diaea dorsata.
Conclusion
A total of 240 species have been newly recorded from the AENP, bringing the total number of species known from the park to 276. All these new data have been made available for the Spider Red Listing project. The AENP spider fauna represents 12.7% of South African species and 32.9% of Eastern Cape species, of which 10 species are ECE that are of special concern and need additional collecting.
Acknowledgements
This survey forms part of the South African National Survey of Arachnida (SANSA) inventories of the South African National Parks. The Agricultural Research Council (ARC) and the South African National Biodiversity Institute (SANBI) Threatened Species Programme are thanked for generously funding SANSA Phase II. The authors would like to thank the park manager and officials of the Addo Elephant National Park for their friendliness and assistance, and all the people who sampled in the park and donated their specimens. The authors are grateful to Dr W. Wiese, husband of the second author, who assisted with the fieldwork. The authors also received financial support from the University of Pretoria. The authors also thank the staff of the Arachnology Unit of the Biosystematics Programme, ARC – Plant Health and Protection, for their assistance with processing and databasing the material collected.
Competing interests
The authors declare that they have no financial or personal relationships that may have inappropriately influenced them in writing this article.
Authors’ contributions
All the authors are team members of SANSA and contributed towards planning this national survey. L.W. participated in fieldwork, while the rest of the authors assisted in the identification of specimens and the preparation and editing of the manuscript.
Funding information
A.S.D.-S. S.H.F. and C.R.H. are grateful for the financial support from the National Research Foundation of South Africa.
Data availability statement
All specimens sampled in this study were deposited in the National Collection of Arachnida (NCA) at the ARC – Plant Health and Protection in Pretoria, South Africa. The specimen data are available from the NCA and SANSA databases on request.
Disclaimer
The views and opinions expressed in this article are those of the authors and do not necessarily reflect the official policy or position of any affiliated agency of the authors.
References
Benoit, P.L.G., 1968, ‘Les Selenopidae africains au Nord du 17e parallèle Sud et reclassement des espèces africaines de la famille (Araneae)’, Revue de Zoologie et de Botanique Africaines 77(1–2), 113–141.
Dippenaar-Schoeman, A.S., 1980, ‘The crab-spiders of southern Africa; Araneae: Thomisidae). 1. The genus Runcinia Simon, 1875’, Journal of the Entomological Society of South Africa 43(2), 303–326.
Dippenaar-Schoeman, A.S., 1988, ‘Annotated check list of the spiders (Araneae) of the Mountain Zebra National Park’, Koedoe 31(1), 151–160. https://doi.org/10.4102/koedoe.v31i1.492
Dippenaar-Schoeman, A.S., 2006, ‘New records of 43 spider species from the Mountain Zebra National Park, South Africa (Arachnida: Araneae)’, Koedoe 49(2), 23–28. https://doi.org/10.4102/koedoe.v49i2.113
Dippenaar-Schoeman, A.S., 2016, ‘SANSA: Protected areas: National parks’, SANSA News 25, 5–6.
Dippenaar-Schoeman, A.S. & Haddad, C.R., 2006, ‘What is the South African National Survey of Arachnida (SANSA) all about?’, SANSA News 1, 1–3.
Dippenaar-Schoeman, A.S., Haddad, C.R., Foord, S.H., Lyle, R., Lotz, L.N. & Marais, P., 2015, ‘South African National Survey of Arachnida (SANSA): Review of current knowledge, constraints and future needs for documenting spider diversity (Arachnida: Araneae)’, Transactions of the Royal Society of South Africa 70(3), 245–277. https://doi.org/10.1080/0035919X.2015.1088486
Dippenaar-Schoeman, A.S., Haddad, C.R., Foord, S.H., Lyle, R., Helberg, L. & Mathebula, S. et al., 2010, First atlas of the spiders of South Africa (Arachnida: Araneae), ARC – Plant Protection Research Institute, Pretoria.
Dippenaar-Schoeman, A.S., Hamer, M. & Haddad, C.R., 2011, ‘Spiders (Arachnida: Araneae) of the vegetation layer of the Mkambati Nature Reserve, Eastern Cape, South Africa’, Koedoe 53(1), 1058. https://doi.org/10.4102/koedoe.v53i1.1058
Foord, S.H., Dippenaar-Schoeman, A.S. & Haddad, C.R., 2011, ‘The faunistic diversity of spiders (Arachnida, Araneae) of the Savanna Biome in South Africa’, Transactions of the Royal Society of South Africa 66(3), 170–201. https://doi.org/10.1080/0035919X.2011.639406
Forbanka, D.N. & Niba, A.S., 2013, ‘Distribution and diversity of epigaiec invertebrate assemblages in Silaka Nature Reserve, Eastern Cape, South Africa’, African Entomology 21(1), 58–69. https://doi.org/10.4001/003.021.0111
Griswold, C.E., 1987, ‘The African members of the trap-door spider family Migidae (Araneae: Mygalomorphae) 1: The genus Moggridgea O.P.-Cambridge, 1875’, Annals of the Natal Museum 28(1), 1–118.
Hamer, M.L. & Slotow, R., 2017, ‘A conservation assessment of the terrestrial invertebrate fauna of Mkambati Nature Reserve in the Pondoland Centre of Endemism’, Koedoe 59(1), a1428. https://doi.org/10.4102/koedoe.v59i1.1428
Huber, B.A., 2003, ‘Southern African pholcid spiders: Revision and cladistic analysis of Quamtana gen. nov. and Spermophora Hentz (Araneae: Pholcidae), with notes on male-female covariation’, Zoological Journal of the Linnean Society 139(4), 477–527. https://doi.org/10.1046/j.0024-4082.2003.00082.x
Jocqué, R., 1984, ‘Linyphiidae (Araneae) from South Africa. Part I: The collection of the Plant Protection Research Institute, Pretoria’, Journal of the Entomological Society of South Africa 47(1), 121–146.
Johnson, C.F., Cowling, R.M. & Phillipson, P.B., 1999, ‘The flora of the Addo Elephant National Park, South Africa: Are threatened species vulnerable to elephant damage?’, Biodiversity & Conservation 8(11), 1447–1456. https://doi.org/10.1023/A:1008980120379
Lewis, A.S.H. & Dippenaar-Schoeman, A.S., 2014, ‘Revision of the spider genus Mystaria Simon, 1895 (Araneae: Thomisidae) and the description of a new genus from the Afrotropical region’, Zootaxa 3873(2), 101–144. https://doi.org/10.11646/zootaxa.3873.2.1
Lombard, A.T., Johnson, C.F., Cowling, R.M. & Pressey, R.L., 2001, ‘Protecting plants from elephants: Botanical reserve scenarios within the Addo Elephant National Park, South Africa’, Biological Conservation 102(2), 191–203. https://doi.org/10.1016/S0006-3207(01)00056-8
Lyle, R. & Dippenaar-Schoeman, A.S., 2015, ‘Red listing of South African spiders’, SANSA Newsletter 23, 1.
Lyle, R. & Haddad, C.R., 2018, ‘Jocquestus, a new genus of trachelid sac spiders from the Afrotropical region (Arachnida: Araneae)’, Zootaxa 4471(2), 309–333. https://doi.org/10.11646/zootaxa.4471.2.4
Midgley, J.M., 2012, ‘Quantifying the structure of epigaeic invertebrate communities along an altitudinal gradient on the Great Escarpment, South Africa’, Unpublished PhD thesis, Rhodes University, Grahamstown.
Mucina, L. & Rutherford, M.C., 2006, The vegetation of South Africa, Lesotho and Swaziland, Strelitzia 19, South African National Biodiversity Institute, Pretoria.
Ruch, J., Heinrich, L., Bilde, T. & Schneider, J.M., 2009a, ‘Relatedness facilitates cooperation in the subsocial spider, Stegodyphus tentoriicola’, BMC Evolutionary Biology 9(257), 1–7. https://doi.org/10.1186/1471-2148-9-257
Ruch, J., Heinrich, L., Bilde, T. & Schneider, J.M., 2009b, ‘The evolution of social inbreeding mating systems in spiders: Limited male mating dispersal and lack of pre-copulatory inbreeding avoidance in a subsocial predecessor’, Biological Journal of the Linnean Society 98(4), 851–859. https://doi.org/10.1111/j.1095-8312.2009.01322.x
Ruch, J., Heinrich, L., Bilde, T. & Schneider, J.M., 2012, ‘Site selection and foraging in the eresid spider Stegodyphus tentoriicola’, Journal of Insect Behaviour 25(1), 1–11. https://doi.org/10.1007/s10905-011-9273-9
South African National Parks (SANParks), 2015, Addo Elephant National Park Management Plan for the period 2015–2025, South African National Parks, Pretoria.
Wesołowska, W. & Haddad, C.R., 2013, ‘New data on the jumping spiders of South Africa (Araneae: Salticidae)’, African Invertebrates 54(1), 177–240. https://doi.org/10.5733/afin.054.0111
Wesołowska, W. & Haddad, C.R., 2018, ‘Further additions to the jumping spider fauna of South Africa (Araneae: Salticidae)’, Annales Zoologici, Warszawa 68(4), 879–908. https://doi.org/10.3161/00034541ANZ2018.68.4.011
Appendix 1
TABLE 1-A1: Spiders of the Addo Elephant National Park, listing their endemicity, conservation status, distribution and areas sampled in the park. |
TABLE 1-A1 (Continues): Spiders of the Addo Elephant National Park, listing their endemicity, conservation status, distribution and areas sampled in the park. |
TABLE 1-A1 (Continues): Spiders of the Addo Elephant National Park, listing their endemicity, conservation status, distribution and areas sampled in the park. |
TABLE 1-A1 (Continues): Spiders of the Addo Elephant National Park, listing their endemicity, conservation status, distribution and areas sampled in the park. |
TABLE 1-A1 (Continues): Spiders of the Addo Elephant National Park, listing their endemicity, conservation status, distribution and areas sampled in the park. |
TABLE 1-A1 (Continues): Spiders of the Addo Elephant National Park, listing their endemicity, conservation status, distribution and areas sampled in the park. |
|