Earthworm Species Occurrence in Agroecosystems in the Midlands, Kwazulu-Natal, South Africa
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African Invertebrates 62(2): 411–425 (2021) doi: 10.3897/AfrInvertebr.62.67875 RESEARCH ARTICLE https://africaninvertebrates.pensoft.net Earthworm species occurrence in agroecosystems in the Midlands, KwaZulu-Natal, South Africa Thembeka C. Nxele1,3, Tarombera Mwabvu2,3, Inam Yekwayo4 1 KwaZulu-Natal Museum, Private Bag 9070, Pietermaritzburg, 3200, South Africa 2 School of Biology & Environmental Sciences, University of Mpumalanga, Private Bag X11283, Mbombela, 1200, South Africa 3 School of Life Sciences, University of KwaZulu-Natal, Private Bag X54001, Westville campus, Durban, 4001, South Africa 4 Department of Biological and Environmental Sciences, Walter Sisulu University, Private Bag X1, Mthatha, 5127, South Africa Corresponding author: Thembeka C. Nxele ([email protected]) Academic editor: Burgert Muller | Received 26 April 2021 | Accepted 2 August 2021 | Published 12 August 2021 http://zoobank.org/A87C1D0F-0D1A-4EBD-AAE4-01E8D7850053 Citation: Nxele TC, Mwabvu T, Yekwayo I (2021) Earthworm species occurrence in agroecosystems in the Midlands, KwaZulu-Natal, South Africa. African Invertebrates 62(2): 411–425. https://doi.org/10.3897/AfrInvertebr.62.67875 Abstract Little is known about the species composition of earthworms in agroecosystems in South Africa even though earthworms provide soil ecosystem services and are useful biological indicators of changes in the habitats. Given the land use and management impact biodiversity, the aim of this study was to docu- ment earthworm species that occur under cultivated land in the KwaZulu-Natal Midlands. A survey of nine farms that practise conservation agriculture was carried out between 2018 and 2020. Twelve earthworm species belonging to four introduced families: Acanthodrilidae (Dichogaster bolaui), Rhin- odrilidae (Pontoscolex corenthrurus), Lumbricidae (Aporrectodea caliginosa, Aporrectodea rosea, Aporrecto- dea trapezoides, Lumbricus rubellus, Octolasion cyaneum, Octolasion lacteum), Megascolecidae (Amynthas aeruginosus, Amynthas corticis, Amynthas gracilis, Amynthas rodericensis) and juveniles from an indigenous family Tritogeniidae were recorded from cultivated fields. The type of crop (habitat) affected both species richness and abundance of earthworms significantly. However, post hoc results showed differences in spe- cies richness between the soya and the maize only, with greater species richness in the maize. Our results demonstrate that habitat type has a major influence on communities of earthworms in agroecosystems. Keywords agroecosystems, diversity, earthworms, indigenous Copyright Thembeka C. Nxele et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 412 Thembeka C. Nxele et al. / African Invertebrates 62(2): 411–425 (2021) Introduction Terrestrial ecosystems benefit immensely from soil organisms (Nielsen et al. 2015). Knowledge on soil fauna has increased and more attention has been given to these taxa in recent years (Brown et al. 2018) and earthworms’ contribution to soil ecosystem and soil fertility has been documented. Earthworms contribute to ecosystem services by converting organic matter into rich humus in the form of casts. Earthworms improve soil fertility and quality, influence soil formation, improve soil nutrient availability, sta- bilise the soil, increase soil porosity, improve water infiltration and increase the overall health of the soil (Lavelle et al. 2006; Jouquet et al. 2006; Brown et al. 2018). Food production relies on healthy soils and there is an urgent need to understand biodiversity and biophysical regulations of soil fertility better (Plisko and Nxele 2015). Therefore, access to accurate taxonomic information of soil organisms is essential. Unfortunately, taxonomists trained to identify soil fauna are in decline (Brown et al. 2018). The adoption of environmentally friendly and sustainable agriculture is there- fore long overdue because of rapid increase in human population, climate change and deteriorating soils (Kassam et al. 2009; Delgado and Gantzer 2015). Earthworm populations tend to do better in no-till systems (Bartz et al. 2013; San- tos et al. 2018), hence earthworms are widely used as soil health indicators (Nadolny et al. 2020). A recent review in Brazilian no-tillage agriculture highlighted that the no-till system promotes earthworm populations (Demetrio et al. 2019). However, according to Santos et al (2018), although work has been done in South America on no-till sys- tems, knowledge of earthworm diversity in agroecosystems is still limited. In South Africa, few studies have documented earthworm species in agroecosys- tems. The studies that looked at the occurrence of earthworms in agricultural eco- systems reported that peregrine species were dominant (Visser and Reinecke 1977; Reinecke and Visser 1980; Dlamini et al. 2001; Haynes et al. 2003; Simonsen et al. 2010). Tillage is known to affect endogenic and anecic earthworm diversity and abundance negatively (Reinecke and Visser 1980) unlike no-tillage agriculture (Pei- gne et al. 2009; Hutcheon et al. 2001). According to Dlamini et al. (2001) and Haynes et al. (2003), earthworms in agroecosystems have not been studied adequate- ly in South Africa. After Nxele (2015) recorded indigenous earthworms in sugar-cane fields that had been under no-till for more than twenty years, we hypothesised that more indigenous species occur under no-till agriculture. As such, a study to docu- ment earthworm diversity under cultivated fields was initiated in 2018 in minimum tillage or no-till agroecosystems. Material and methods Study sites Nine farms in KwaZulu-Natal Midlands (Fig. 1) were sampled for earthworms. Each farm had either sugar-cane, maize, soya, ryegrass pasture, mixed species pasture or a Earthworm species occurrence in agroecosystems 413 Figure 1. Sampling sites in KZN Midlands. Table 1. Location, crop type and age of no-till agriculture and numbers of species of earthworms that were recorded in the KZN Midlands. Area Location Crop sampled Age (years) Number of species Loskop 28°54'33.2"S, 29°33'35.5"E Maize, soya > 20 5 Bergville 28°38'28.8"S, 29°16'37.8"E Maize, soya > 20 3 Karkloof 29°22'50.1"S, 30°17'34.5"E Maize, ryegrass pasture > 20 6 Karkloof 29°20'29.3"S, 30°13'03.6"E Maize > 20 2 Balgowan 29°25'27.0"S, 30°01'29.5"E Maize, ryegrass pasture < 10 3 Lidgeton 29°26'06.2"S, 30°05'10.0"E Maize, mixed species pasture < 20 4 Dalton 29°13'47.2"S, 30°40'54.7"E Sugarcane < 20 4 Nottingham Road 29°25'08.9"S, 30°00'27.7"E Mixed species pasture < 10 3 Nottingham Road 29°26'21.5"S, 29°59'03.0"E ryegrass pasture < 10 2 mixture of crops (Table 1). All the farms practise conservation agriculture; however, the farms have been under no-tillage for a different number of years. Earthworm sampling Earthworms were collected quantitatively and qualitatively. The quantitative method follows that of Nxele et al. (2015) and Bartz et al. (2014) with slight modification on plot sizes. Sampling was carried out in one hectare with nine sampling points; adjacent sampling points were 30 m apart. Earthworms were collected by digging out 50 cm x 50 cm × 20 cm soil monoliths. The soil was hand-sorted for earthworms in large plastic trays (50 cm × 40 cm × 5 cm). The holes were filled back with the soil after removing the specimens from the soil. Active searching under stones and logs was part of qualita- tive sampling. Specimens that were collected were washed and narcotised using 20% ethanol solution. Some specimens were preserved in absolute ethanol for DNA analy- sis. The remaining specimens were fixed in 4% formalin for at least 24 hours before 414 Thembeka C. Nxele et al. / African Invertebrates 62(2): 411–425 (2021) being preserved in 75% ethanol. Studies of the internal anatomy were conducted after dorsal dissections. The KZN museum database and the following literature: Plisko 2001, 2010; Ljungstom 1972; Michaelsen 1899, 1908, 1913; Reynold and Reinecke 1976; Zicsi and Reinecke 1992; Visser and Reinecke 1977 and Pickford 1937 were used to obtain information on distribution. All new material is deposited in the Oli- gochaeta collection in the KwaZulu-Natal Museum (NMSA). Data analyses Data analysis was per crop type regardless of which farm it came from. Species richness and abundance datasets were analysed in R using the generalised linear mixed models (GLMMs) because data were not normally distributed. The lme4 package (Bates et al. 2015) was used when calculating GLMMs. The Poisson distribution was the best fit for the species richness dataset, while the negative binomial distribution was the best fit for the species abundance dataset (Bolker et al. 2009). In the models, the type of crop (maize, pasture, soya and sugar-cane) was the fixed factor, while the random factor was the farm. The multcomp package (Hothorn et al. 2008) was used to determine the similarities and/or differences between pairs of crops. Abbreviations KZN KwaZulu-Natal; NW North West; EC Eastern Cape; GP Gauteng; WC Western Cape; MP Mpumalanga; NC Northern Cape; FS Free State. LP Limpopo, Checklist Family Acanthodrilidae Claus, 1880 sensu Csuzdi (2010) Subfamily Benhamiinae Michaelsen, 1897 sensu Csuzdi (1996) Genus Dichogaster Beddard, 1888 Subgenus Dichogaster (Diplothecodrilus) Csuzdi, 1996 Dichogaster bolaui (Michaelsen, 1891) Type locality. Bergedorf, Germany RSA distribution (Old material). LP: Entabeni State