Abstract
Banana orchards, like other crops, are attacked by insect pests. Although spiders are among the natural enemies of these pests, we have limited data on the different types of spider species that occur in the banana orchards in South Africa. As such, in this study, we determined the occurrence of species of ground-dwelling spiders in a banana orchard. In addition, we compared species richness, abundance and composition of spiders sampled using pitfall traps in the colder (July) and warmer (November) months. A total of 635 specimens from 14 families, 30 genera and 33 species were recorded. Although the Corinnidae was represented by two species only, one of these species, Copa flavoplumosa Simon, 1886, contributed 53% of the total abundance of spiders recorded from the banana orchard. The Lycosidae, Salticidae, Gnaphosidae and Zodariidae were the most species-rich families, while the rest of the families were represented by one or two species only. There were greater species richness and abundance of spiders sampled in November than in July. In addition, our study showed that assemblages of ground-dwelling spiders are influenced by season. Although most of the collected spider species are represented by fewer individuals, their presence suggests a greater importance of spiders in the biological control of insect pests. Furthermore, despite the low numbers of individuals and species of spiders that we collected during the colder month, variation in species composition between the warmer and colder months suggests the continuous presence of spiders in the banana orchard, which augurs well for pest control.
Conservation implications: Understanding the taxonomic diversity of spiders and their importance in pest control is likely to reduce reliance on chemicals in banana orchards. Thus, reducing both advertent and inadvertent biodiversity loss and ensuring ecosystem stability.
Keywords: agroecosystems; seasons; conservation; abundance; species richness.
Introduction
In South Africa, spiders have been studied in natural, seminatural and agricultural landscapes mostly as an initiative of the South African National Survey of Arachnida (Dippenaar-Schoeman et al. 2023). The agricultural systems in which spiders have been surveyed in South Africa include avocado, citrus, cotton, macadamia, maize, pistachio (Dippenaar-Schoeman et al. 2013) and litchi (Yekwayo et al. 2025a). Improving our knowledge of spider diversity and composition, especially in agroecosystems, is vital because spiders are natural enemies of insect pests in these systems. Despite an intensive survey of spiders in South Africa, bananas (Musa sp.) are one of the least studied crops (Simba, Yekwayo & Mwabvu 2023).
Banana is native to Southeast Asia and is a staple food in East African countries, with Uganda leading the continent in the production of bananas (Karamura et al. 1998; Viljoen et al. 2004). According to the Department of Agriculture, Land Reform and Rural Development (2020), bananas are produced in subsistence and commercial farming, with the latter having both seasonal and permanent employees; production is for local markets and enhances socioeconomic development in South Africa.
Banana production is affected by pests, the most important pest being the banana weevil, Cosmopolites sordidus Germar, 1824 (De Graaf et al. 2008). Some of the control methods for the banana weevil include cultural and chemical methods. However, in KwaZulu-Natal in South Africa, De Graaf et al. (2008) found that these methods did not reduce pest population nor improve the yield. Thus, biological control could be an alternative method, with spiders as one of the diverse predators in agroecosystems being possible candidates (Dippenaar-Schoeman et al. 2013). In India, spiders prey on a variety of insects in banana orchards; thus, spiders may reduce the population of pests because of their diverse feeding behaviours (Keswani & Vankhede 2014; Kumar 2021). In addition, banana orchards support diverse spider species because they occupy different microhabitats, including those in the ground layer and the foliage (Keswani & Vankhede 2014).
In a banana orchard in Hazyview, Mpumalanga, Simba et al. (2023) found that Corinnidae, Lycosidae, Salticidae and Oonopidae were the most abundant families, while the Salticidae, Corinnidae, Gnaphosidae and Lycosidae were most species rich. The study by Simba et al. (2023) was foundational, but the species composition of spiders in this orchard remains unknown. Thus, in the current study, we provide a checklist of species of ground-dwelling spiders that occur in a banana orchard in Hazyview in the Mpumalanga province. Additionally, we also determined the effect of season on species richness, abundance and composition of ground-dwelling spiders.
Methodology
Spiders were surveyed in a banana orchard in an Agricultural Research Council (ARC) farm (−25.112700, 31.083100) in Hazyview, Mpumalanga, South Africa. The orchard consisted of banana plants that ranged from 15 to 20 years in age. These bananas were planted in blocks with the smallest being approximately 8000 m2. Separating these blocks of bananas were dirt roads (± 8 m wide). The surface layer in these blocks of bananas was characterised by a layer of plant litter that ranged in depth from 1.7 cm to 13.5 cm. The banana plants were watered every second week during the dry winter season, while no irrigation took place during the wet summer season. Two herbicides, glyphosate and paraquat, were used to control weeds within the orchard.
A total of five sampling stations, one per block, were established with each being in the middle of the block. Each sampling station was a 10 m × 10 m quadrat. Sampling in these five stations was conducted during the dry-cold winter season in July 2020 and during the wet-warm summer season in November 2020. For the purposes of the checklist, this study included data from 10 stations that Simba et al. (2023) collected in July 2020 only from the same ARC farm. Therefore, a total of 15 stations were sampled in July. Pitfall traps with 50% ethylene glycol solution as a preservative were used to collect spiders. Within each station, there were four 10-m long line transects that were parallel to each other; each of those line transects had six pitfalls. These pitfall traps were active for a period of 21 days in each month; within that period, they were emptied on every 7th day. The specimens from the 24 traps per week (72 traps per month) were combined to form a sample for each station. Spiders were extracted from the samples and identified by the first and last authors. Voucher specimens of spiders are housed in the National Collection of Arachnida at ARC in Pretoria, South Africa.
Although we had a low sample size (five stations that were sampled in July and November), we statistically compared species richness, abundance and composition between the months. To determine the normality of species richness and abundance datasets, we used the Shapiro–Wilk test, and both datasets were normally distributed. As a result, linear models in R version 4.5.1 (R Foundation for Statistical Computing) were used to compare spider species richness and abundance between winter and summer seasons. In PRIMER 7, we tested the homogeneity of multivariate dispersions using the permutational analysis of multivariate dispersions (PERMDISP), and species composition of spiders was compared using the permutational multivariate analysis of variance (PERMANOVA).
Ethical considerations
This article followed all ethical standards for research without direct contact with human or animal subjects. Before 2020, the University of Mpumalanga did not require ethical clearance for studies conducted on arthropods.
Results and discussion
A total of 635 individuals (all adults) in 14 families, 30 genera and 33 species were collected from the study site (Table 1). More than half (n = 337, 53.07 %) of all the individuals were represented by a single species, Copa flavoplumosa Simon, 1886 (Corinnidae), as shown in Table 1. One other corinnid species, Copuetta magna Haddad, 2013, represented by a single individual, was also sampled (Table 2). The greater abundance of C. flavoplumosa is linked to this generalist species having a wide distribution across all South African provinces (Dippenaar-Schoeman et al. 2023; Haddad et al. 2023). In addition to C. flavoplumosa having been recorded in most South African biomes, this species has also been recorded in crops, such as avocado, citrus, cotton, macadamia, maize, pistachio (Dippenaar-Schoeman 2023; Dippenaar-Schoeman et al. 2010, 2023) and litchi (Yekwayo et al. 2025a).
| TABLE 1: Abundance in families of spiders and their taxonomic diversity in a banana orchard in Hazyview in Mpumalanga, South Africa. |
| TABLE 2: List of species and individuals collected from the banana orchard in Hazyview in Mpumalanga, South Africa. |
After the Corinnidae, the highest abundance of spiders was recorded in the Lycosidae, Salticidae, Oonopidae and Zodariidae. The Ctenidae and Gnaphosidae were represented by 14 individuals each, while there were nine individuals in the Linyphiidae. The other six families had fewer than five individuals each. The Lycosidae and Salticidae had five genera and six species each, while there were four genera and species of the Gnaphosidae and Zodariidae that were recorded in the banana orchard. The remaining 10 families of spiders recorded in the banana orchard either had one or two species.
The higher abundance and species richness of the lycosids (wolf spiders) and salticids (jumping spiders) can be attributed to the pitfall trapping method, which predominantly captures surface-active species. The wolf spiders (ground-dwellers) and jumping spiders (some are ground-dwellers) are active hunters (Dippenaar-Schoeman 2023); this increases their likelihood of falling into pitfall traps. Among the jumping spiders collected, Stenaelurillus guttiger Simon, 1901, was the most abundant (n = 70), which can be explained by their habitat preference; in this case, the deeper layer of plant litter and higher moisture from irrigation. Stenaelurillus guttiger forages in leaf litter among its prey items are collembolans (Pekár et al. 2020), which are abundant in moist habitats. The second most abundant jumping spider species was Hispo georgius Peckham & Peckham, 1892 (Table 2). This was unexpected given that in the past, H. georgius was mostly recorded on foliage (Wesołowska & Haddad 2009). Hyllus argyrotoxus Simon, 1902; Thyene natalii Peckham & Peckham, 1903 and Thyene ogdeni Peckham & Peckham, 1903 had an individual each, while there were two individuals of Natta horizontalis Karsch, 1879. The low catches in these jumping spiders may be attributed to the fact that these species mostly occupy the foliage rather than the ground layer (Dippenaar-Schoeman 2023; Wesołowska & Haddad 2009).
Cydrela schoemanae Jocqué, 1991 (n = 29) and Diores lesserti Lawrence, 1952 (n = 7) were the most abundant Zodariidae that we collected from the banana orchard. The abundance of the Zodariidae species and Gamasomorpha australis Hewitt, 1915 (n = 39) in the Oonopidae could be due to their activity near pitfall traps as ground-dwellers (Dippenaar-Schoeman 2023; Dippenaar-Schoeman et al. 2024). Furthermore, gnaphosids are ground dwellers that have been captured using pitfall traps (Dippenaar-Schoeman 2023; Foord et al. 2019). However, in our study, we recorded four species of gnaphosids, which were represented by fewer individuals.
The banana orchard may not have provided a suitable habitat for Oxyopes falconeri Lessert, 1915, and Oxyopes russoi Caporiacco, 1940 because, according to Dippenaar-Schoeman et al. (2020a), these species are abundant in grasslands. The ground layer of the banana orchard lacked grasses; instead, it was covered by a deep layer of plant litter. The banana litter may explain the occurrence of the nine individuals of Agyneta habra Locket, 1968, that we sampled, as this species can make its sheet-webs on the litter layer (Dippenaar-Schoeman 2023). Additionally, A. habra appears not to be restricted to a particular vegetation type as this species has been recorded in a variety of agroecosystems and in all South African provinces (Dippenaar-Schoeman 2023; Dippenaar-Schoeman et al. 2023). Furthermore, the low numbers of some spiders (e.g. Anyphops pococki Lawrence, 1940, Scytodes sp. and Theuma sp.) may be because these species are usually found underneath rocks, stones and the bark of trees (Dippenaar-Schoeman 2023; Dippenaar-Schoeman et al. 2020b).
We recorded significantly greater species richness (t = 3.92, p = 0.004) and abundance of spiders (t = 4.45, p = 0.002) in November than in July. Although PERMDISP showed no differences (F = 4.96, p = 0.11) in the dispersion of stations between seasons, PERMANOVA revealed significant variations (Pseudo-F = 4.85, p = 0.009) in species composition of spiders between the dry-cold winter and wet-warmer months. These results are in line with previous studies on the response of spiders to seasonal changes in macadamia and litchi orchards in Mpumalanga (Yekwayo et al. 2025a, 2025b).
Conclusion
Although some species were sampled in low numbers in the banana orchard, their presence remains important in the biological control of insect pests; their low numbers may be explained by the sampling method used. As a result, we recommend additional sampling that will target both the ground and vegetation layers in the banana orchard.
Acknowledgements
We are grateful to the Agricultural Research Council, Hazyview, for allowing us to conduct the study. Additionally, we thank our field assistants (Ntombikayise Ndwandwe, Sizwe Sambo, Simphiwe Sibeko, Sithabile Ndwandwe and Simphiwe Sibitane).
This article is based on research previously presented in abstract form at the National Global Change Conference, held at the University of Mpumalanga on 01–04 December 2025. The abstract has since been developed into a full article, which has been expanded and revised for journal publication. This republication is done with permission from the conference organisers.
The author reported that they received funding from the National Research Foundation, which may be affected by the research reported in the enclosed publication. The author has disclosed those interests fully and has implemented an approved plan for managing any potential conflicts arising from their involvement. The terms of these funding arrangements have been reviewed and approved by the affiliated university in accordance with its policy on objectivity in research.
Competing interests
The authors declare that they have no financial or personal relationship(s) that may have inappropriately influenced them in writing this article.
CRediT authorship contribution
Inam Yekwayo: Conceptualisation, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Validation, Visualisation, Writing – original draft. Tarombera Mwabvu: Conceptualisation, Investigation, Methodology, Writing – review & editing. Vhuawelo Simba: Investigation, Methodology, Writing – review & editing. Anna S. Dippenaar-Schoeman: Data curation, Methodology, Writing – original draft. All authors reviewed the article, contributed to the discussion of results, approved the final version for submission and publication and take responsibility for the integrity of its findings.
Funding information
The study was funded by the National Research Foundation (Competitive Support for Unrated Researchers – Grant number: 134117).
Data availability
The data analysed in this study are included within the manuscript, and the voucher specimens are housed in the National Collection of Arachnida at the Agricultural Research Council in Pretoria.
Disclaimer
The views and opinions expressed in this article are those of the authors and are the product of professional research. It does not necessarily reflect the official policy or position of any affiliated institution, funder, agency or that of the publisher. The authors are responsible for this article’s results, findings and content.
References
De Graaf, J., Govender, P., Schoeman, A.S. & Viljoen, A., 2008, ‘Efficacy of cultural control measures against the banana weevil, Cosmopolites sordidus (Germar), in South Africa’, Journal of Applied Entomology 132(1), 36–44. https://doi.org/10.1111/j.1439-0418.2007.01252.x
Department of Agriculture, Land Reform and Rural Development, 2020, A profile of the South African banana market value chain, South African Government, Arcadia.
Dippenaar-Schoeman, A.S., 2023, Field guide to the spiders of South Africa, Struik Nature, Cape Town.
Dippenaar-Schoeman, A.S., Haddad, C.R., Foord, S.H. & Lotz, L.N., 2020a, The Oxyopidae of South Africa. Version 1: 1–57, pp. 1–57, South African National Survey of Arachnida Photo Identification Guide, Irene.
Dippenaar-Schoeman, A.S., Haddad, C.R., Foord, S.H. & Lotz, L.N., 2020b, The Selenopidae of South Africa. Version 1: 1–23, pp. 1–82, South African National Survey of Arachnida Photo Identification Guide, Irene.
Dippenaar-Schoeman, A.S., Haddad, C.R., Foord, S.H., Lyle, R., Lotz, L., Helberg, L. et al., 2010, South African National Survey of Arachnida technical report 2010 version 1: First Atlas of the spiders of South Africa (Arachnida: Araneae), South African National Survey of Arachnida, Irene.
Dippenaar-Schoeman, A.S., Haddad, C.R., Lotz, L.N., Booysen, R., Steenkamp, R.C. & Foord, S.H., 2023, ‘Checklist of the spiders (Araneae) of South Africa’, African Invertebrates 64(3), 221–289. https://doi.org/10.3897/AfrInvertebr.64.111047
Dippenaar-Schoeman, A.S., Jocqué, R., Haddad, C.R., Foord, S.H. & Lotz, L.N., 2024, The Zodariidae of South Africa. Part 1 (A-D). Version 1, pp. 1–91, South African National Survey of Arachnida Photo Identification Guide, Irene.
Dippenaar-Schoeman, A.S., Van den Berg, A.M., Haddad, C.R. & Lyle, R., 2013, ‘Current knowledge of spiders in South African agroecosystems (Arachnida, Araneae)’, Transactions of the Royal Society of South Africa 68(1), 57–74. https://doi.org/10.1080/0035919X.2012.755136
Foord, S., Dippenaar-Schoeman, A.S, Haddad, C.R., Schoeman, C., Hahn, N. & Lyle, R., 2019, ‘Spider checklist for the Blouberg, in the Vhembe Biosphere Reserve, South Africa’, Bothalia 49(1), a2455. https://doi.org/10.4102/abc.v49i1.2455
Haddad, C.R., Dippenaar-Schoeman, A.S., Foord, S.H. & Lotz, L.N., 2023, The Corinnidae of South Africa. Version 1, pp. 1–72, South African National Survey of Arachnida Photo Identification Guide, Irene.
Karamura, E., Frison, E., Karamura, D.A. & Sharrock, S., 1998, ‘Banana production systems in Eastern and Southern Africa’, in C. Picq, E. Fouré & E.A. Frison (eds.), Bananas and food security, pp. 401–412, INIBAP, Montpellier.
Keswani, S. & Vankhede, G., 2014, ‘Diversity, population and habitat used by spiders in banana agro-ecosystem’, Indian Journal of Arachnology 3(1), 12–27.
Kumar, A., 2021, ‘Diversity and abundance of spider species in the banana agroecosystem of Etawah District: A taxonomic and ecological assessment’, Journal of Agriculture and Veterinary Science 14(1), 61–67.
Pekár, S., Dusšátková, L.P., Michálek, O. & Haddad, C.R., 2020, ‘Coexistence of two termite-eating specialists (Araneae)’, Ecological Entomology 45(6), 1307–1317. https://doi.org/10.1111/een.12914
Simba, V., Yekwayo, I. & Mwabvu, T., 2023, ‘Commercial banana and macadamia plantations in a savanna matrix support high levels of arthropod diversity’, African Entomology 31, e14047. https://doi.org/10.17159/2254-8854/2023/a14047
Viljoen, A., Kunert, K., Kiggundu, A., Escalant, J.V. & Bornman, C.H., 2004, ‘Biotechnology for sustainable banana and plantain production in Africa: The South African contribution’, South African Journal of Botany 70(1), 67–74. https://doi.org/10.1016/S0254-6299(15)30308-2
Wesołowska, W. & Haddad, C.R., 2009, ‘Jumping spiders (Araneae: Salticidae) of the Ndumo Game Reserve, Maputaland, South Africa’, African Invertebrates 50(1), 13–103. https://doi.org/10.5733/afin.050.0102
Yekwayo, I., Mwabvu, T. & Dippenaar-Schoeman, A.S., 2025a, ‘Survey of epigeic spiders (Arachnida: Araneae) in a litchi orchard in Mpumalanga, South Africa’, Koedoe 67(1), a1851. https://doi.org/10.4102/koedoe.v67i1.1851
Yekwayo, I., Mwabvu, T., Simba, V. & Dippenaar-Schoeman, A.S., 2025b, ‘Seasonal variation and species richness of epigeic spiders in macadamia orchards, South Africa’, African Journal of Ecology 63(6), e70086. https://doi.org/10.1111/aje.70086
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