# Updated list of Collembola species currently recorded from South Africa

Charlene Janion-Scheepers, Louis Deharveng, Anne Bedos, Steven L. Chown

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Charlene Janion-Scheepers, Louis Deharveng, Anne Bedos, Steven L. Chown. Updated list of Collembola species currently recorded from South Africa. Zookeys, 2015, 503, pp.55-88. 10.3897/zookeys.503.8966. hal-01220558

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# Updated list of Collembola species currently recorded from South Africa

Charlene Janion-Scheepers¹,³, Louis Deharveng², Anne Bedos², Steven L. Chown³

¹ Centre for Invasion Biology, Department of Botany and Zoology, Stellenbosch University, Private Bag X1, Matieland 7602, South Africa ² Institut de Systématique, Évolution, Biodiversité ISYEB - UMR 7205 - CNRS, MNHN, UPMC, EPHE, Muséum national d'Histoire naturelle, Sorbonne Universités, 45 rue Buffon, F-75005, Paris, France ³ School of Biological Sciences, Monash University, Clayton, Victoria, Australia

Corresponding author: Charlene Janion-Scheepers (charlene.janionscheepers@monash.edu)

Academic editor: W. Weiner | Received 18 November 2014 | Accepted 5 April 2015 | Published 11 May 2015

http://zoobank.org/C0DA02DC-49B0-43C4-A012-5377CAA94629

**Citation:** Janion-Scheepers C, Deharveng L, Bedos A, Chown SL (2015) Updated list of Collembola species currently recorded from South Africa. ZooKeys 503: 55–88. doi: 10.3897/zookeys.503.8966

## Abstract

Understanding the abundance and richness of species is one of the most fundamental steps in effecting their conservation. Despite global recognition of the significance of the below-ground component of diversity for ecosystem functioning, the soil remains a poorly studied terrestrial ecosystem. In South Africa, knowledge is increasing for a variety of soil faunal groups, but many still remain poorly understood. We have started to address this gap in the knowledge of South African soil biodiversity by focusing on the Collembola in an integrated project that encompasses systematics, barcoding and ecological assessments. Here we provide an updated list of the Collembola species from South Africa. A total of 124 species from 61 genera and 17 families has been recorded, of which 75 are considered endemic, 24 widespread, and 25 introduced. This total number of species excludes the 36 species we consider to be dubious. From the published data, Collembola species richness is high compared to other African countries, but low compared to European countries. This is largely a consequence of poor sampling in the African region, as our discovery of many new species in South Africa demonstrates. Our analyses also show that much ongoing work will be required before a reasonably comprehensive and spatially explicit picture of South Africa's springtail fauna can be provided, which may well exceed 1000 species. Such work will be necessary to help South Africa meet its commitments to biodiversity conservation, especially in the context of the 2020 Aichi targets of the Convention on Biological Diversity.

## Keywords

Biodiversity, endemicity, soil fauna, introduced species, taxonomy
**Introduction**

The documentation of biodiversity is an essential first step for its conservation. A major barrier to so doing for invertebrates is a lack of taxonomic information on various groups. This taxonomic impediment and its implications for biodiversity studies have been widely discussed (Godfray 2002, Samper 2004). Despite these challenges, taxonomic knowledge continues to increase globally (Nilsson-Örtman and Nilsson 2010, Joppa et al. 2011, Platnick 2014, van Noort 2014). Nonetheless, given rapid environmental change and its effects on biodiversity (Butchart et al. 2010), it is unclear what the rate of extinction is for many groups (Pimm et al. 2010, Costello et al. 2013), complicating conservation efforts and assessments of their efficacy, thus underscoring the urgency to further document global biodiversity (Dirzo and Raven 2003, Bacher 2012).

This situation is as true for southern Africa as it is elsewhere. Knowledge of the South African fauna is increasing rapidly, especially in the case of a wide range of invertebrate groups (Foord et al. 2002, Robertson 2000, 2002, Parr et al. 2003, Dippenaar-Schoeman et al. 2006, Dippenaar-Schoeman and González Reyes 2006, Haddad and Dippenaar-Schoeman 2006, Hlavac 2007, Rousse and van Noort 2013). Nonetheless, many groups still remain relatively poorly studied, especially soil-dwelling taxa, which are essential for both above- and below-ground ecosystem functioning (Wardle et al. 2004, Hugo-Coetzee and Avenant 2011, Janion et al. 2011a). At the same time, considerable impacts on biodiversity continue to be documented as a consequence of habitat modification for agriculture and urban development, biological invasions, pollution, and climate change (Erasmus et al. 2002, Rouget et al. 2003, Biggs et al. 2008, Chown 2010, Pryke and Samways 2010, Huntley and Barnard 2012, Liu et al. 2012). In consequence, much need exists for documenting and understanding biodiversity and the processes underlying its variation across a wide range of groups, and especially the soil fauna.

Collembola are amongst the most widespread and abundant soil arthropods (Petersen and Luxton 1982, Hopkin 1997). Despite their obvious significance in soil systems, their utility as bioindicators (Lawrence 1953, Hopkin 1997, van Straalen 1998), their significance in the alien species faunas of many areas (Roques et al. 2009, Terauds et al. 2011), and the current growth in both morphological (Deharveng 2004) and molecular (Hogg and Hebert 2004, Rougerie et al. 2009) means to assess their diversity, they remain poorly known in South Africa. Indeed, by comparison with other regions of the world (Deharveng 2004), and other invertebrate taxa in the country (Scholtz and Chown 1995, Robertson 2000, Foord et al. 2011, Dippenaar-Schoeman 2014), knowledge of the group can be considered scanty.

The first attempt to collate all taxonomic information on the Collembola of South Africa was undertaken by Paclt (1959), listing *ca.* 65 species. Subsequently, an unpublished list entitled “Aquatic Collembola of South Africa” was made available online (P. Greenslade, no date), while Thibaud (2013) listed most publications until 2013. To date there are 38 publications on Collembola recorded or described from South Africa, the earliest by Börner (1908). Most notably, comprehensive descriptions were made by
Yosii (1959), Paolt (1959, 1964, 1965, 1967), Coates (1968a, 1968b, 1969), Weiner and Najt (1991, 1998, 1999), and later Barra (1994, 1995, 1997, 1999, 2001, 2002, Barra and Weiner 2009). However, little other work has been done and the current list of species for the country is clearly an underestimate, with an incomplete understanding of which species might be introduced and thus might have substantial impacts, despite the fact that such impacts have been suggested for the country (Annecke and Moran 1982, Liu et al. 2012).

To address this substantial gap in the knowledge of soil biodiversity, a collaborative project was established in 2008 (Janion et al. 2011a, Bengtsson et al. 2011, 2012). Besides large-scale sampling and systematic assessments, which have resulted in new discoveries and species descriptions (Janion et al. 2011b, Potapov et al. 2011, Janion et al. 2012, 2013), a major component of the project has comprised the compilation of all currently available information on Collembola recorded from South Africa. Here we present this compilation as an updated checklist. It will provide a starting point for understanding the diversity of this group, as has been done for other geopolitical regions (e.g. Culik and Zeppelini 2003, Abrantes et al. 2010, 2012), and will assist South Africa to meet its obligations under the Convention on Biological Diversity (see for example Aichi Target 9 on identifying invasive alien species, and Aichi Target 17 on a national biodiversity strategy, http://www.cbd.int/sp/targets).

## Methods

All publications on Collembola species described or recorded from South Africa were collated from Salmon (1964) and Thibaud (2013). The list was checked and completed using the website “Checklist of the Collembola of the World” (Bellinger et al. 2014), the bi-annual bibliographical lists issued by the Museum National d'Histoire Naturelle (Paris, France), Zoological Record, Web of Science™ (full date range of 1900 to 2014), and genus or species revisions from taxonomic journals sourced from the references identified using the original search methods. Nomenclature follows Bellinger et al. (2014), as it may have changed for certain taxonomic groups since the original description of the species. All published papers and webpages were examined and the following information was recorded when available: collection details including date, collector, province, place, nearest town, habitat type, and collection method, type locality and accession number if given. Only species with full species names were included in the species list of Table 2, thus excluding morphospecies identified to genus or to suspected species (e.g. *Seira* sp. or *Isotomurus* cf. *maculatus*). However, every record from the literature is listed in the Supplementary material (Suppl. material 1). The species were assigned a South African province from the locality recorded. From these points a species richness map was produced in ArcMap V10.2 (ESRI 2014).

The species were also divided into the following categories based on their distribution: 1) endemic if they were described from South Africa and have not been recorded elsewhere, 2) introduced if there is evidence from the literature that the species was
introduced from another place, 3) widespread if the species is also present outside of South Africa but its origin is unknown, thus not considered introduced, or 4) dubious, when the species name given in the literature is considered a misidentification based on current taxonomic knowledge or if subsequent taxonomic work suggested this is the case (see Suppl. material 1).

To make an estimate of expected species richness, we used data collected from extensive sampling in the Western Cape Province of South Africa, which has been the main focus of much work on the group. The dataset comprises a total of 217 samples we obtained using several sampling techniques (see below) in as many localities and different microhabitats as possible throughout the Western Cape, including Afromontane forest, different fynbos vegetation types (see Mucina and Rutherford 2006), intertidal habitats, caves, and disturbed areas such as gardens and agricultural areas. Leaf litter, moss, rotten wood and soil samples were taken at different sampling sites over the duration of the project (2008–2012), and occasionally sieving and pitfall traps were also used. Typically, samples were extracted using a Berlese-Tullgren approach for five to seven days, or until dry (Berlese 1905, Tullgren 1918, Hopkin 1997). In addition, active searching was done in the field. Riparian soil was washed for water-dependent species, which were collected with a fine brush on the surface of water. Fine sand such as sea sand was washed in the laboratory and animals were also collected with a brush. Vegetation such as branches from bushes, fynbos shrubs, and grasses was beaten over a tray and animals were collected by means of an aspirator. All samples are stored in 96–99% ethanol at the Centre for Invasion Biology (C.I.B) Stellenbosch, or the Museum National d'Histoire Naturelle (MNHN) Paris. As identifications and species descriptions are still ongoing, we only used confirmed morphospecies for the purpose of calculating the number of species expected for the Western Cape.

Sampled-based rarefaction curves were plotted to estimate the number of species for the Western Cape, using Chao1 and Jacknife 2 in EstimateS V8.2.0 (Colwell 2009). Jacknife 2 does not require data to be normally distributed and provides conservative, but accurate estimates (Magurran 2004). Sampling is considered adequate when the rarefaction curves and the estimators converge at the highest observed values (Longino et al. 2002).

## Results

According to the literature, a total of 160 species from 61 genera and 17 families have been recorded from South Africa (Table 1), with a relatively steady increase in descriptions since the first records in the early 1900s (Fig. 1). Of the recorded species, 36 are considered dubious, most of them misidentified records from Paclt (1959, 1967). Of the other species, 75 are endemic, 25 are thought to be alien species introduced to the country by human activity, and 24 have a widespread distribution, at least so far as current sampling indicates (Table 2). The majority of species have been recorded from
<table><caption>**Table 1.** A summary of the Collembola species recorded from South Africa based on the literature.</caption><thead><tr><th></th><th>Number of species recorded from literature</th><th>Number of species accepted from literature</th><th>Introduced</th><th>Endemic</th><th>Widespread</th></tr></thead><tbody><tr><td><strong>PODUROMORPHA</strong></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Hypogastruridae</td><td>19</td><td>11</td><td>4</td><td>5</td><td>2</td></tr><tr><td>Brachystomellidae</td><td>6</td><td>6</td><td>1</td><td>5</td><td>0</td></tr><tr><td>Neanuridae</td><td>16</td><td>15</td><td>2</td><td>10</td><td>3</td></tr><tr><td>Odontellidae</td><td>3</td><td>2</td><td>0</td><td>1</td><td>1</td></tr><tr><td>Onychiuridae</td><td>5</td><td>1</td><td>0</td><td>1</td><td>0</td></tr><tr><td>Tullbergiidae</td><td>8</td><td>7</td><td>1</td><td>3</td><td>3</td></tr><tr><td><strong>TOTAL</strong></td><td><strong>57</strong></td><td><strong>42</strong></td><td><strong>8</strong></td><td><strong>25</strong></td><td><strong>9</strong></td></tr><tr><td><strong>ENTOMOBRYOMORPHA</strong></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Isotomidae</td><td>23</td><td>19</td><td>5</td><td>8</td><td>6</td></tr><tr><td>Entomobryidae</td><td>49</td><td>36</td><td>8</td><td>25</td><td>3</td></tr><tr><td>Cyphoderidae</td><td>10</td><td>9</td><td>0</td><td>8</td><td>1</td></tr><tr><td>Paronellidae</td><td>1</td><td>1</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Tomoceridae</td><td>1</td><td>1</td><td>0</td><td>1</td><td>0</td></tr><tr><td><strong>TOTAL</strong></td><td><strong>84</strong></td><td><strong>66</strong></td><td><strong>13</strong></td><td><strong>42</strong></td><td><strong>11</strong></td></tr><tr><td><strong>NEELIPLEONA</strong></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Neelidae</td><td>1</td><td>1</td><td>0</td><td>0</td><td>1</td></tr><tr><td><strong>TOTAL</strong></td><td><strong>1</strong></td><td><strong>1</strong></td><td><strong>0</strong></td><td><strong>0</strong></td><td><strong>1</strong></td></tr><tr><td><strong>SYMPHYPLEONA</strong></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Sminthurididae</td><td>2</td><td>1</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Katiannidae</td><td>5</td><td>4</td><td>1</td><td>1</td><td>2</td></tr><tr><td>Dicyrtomidae</td><td>2</td><td>2</td><td>1</td><td>1</td><td>0</td></tr><tr><td>Bourletiellidae</td><td>7</td><td>7</td><td>1</td><td>6</td><td>0</td></tr><tr><td>Sminthuridae</td><td>2</td><td>1</td><td>1</td><td>0</td><td>0</td></tr><tr><td><strong>TOTAL</strong></td><td><strong>18</strong></td><td><strong>15</strong></td><td><strong>4</strong></td><td><strong>8</strong></td><td><strong>3</strong></td></tr><tr><td><strong>TOTAL</strong></td><td><strong>160</strong></td><td><strong>124</strong></td><td><strong>25</strong></td><td><strong>75</strong></td><td><strong>24</strong></td></tr></tbody></table>

the Western Cape (67 species), Kwazulu-Natal (46 species) and the Eastern Cape (20 species) (Fig. 2). Records from the other provinces are sparse (1–10 species), with the North West Province and Limpopo having the lowest recorded richness (three and one species, respectively). Although many authors did not indicate the habitat type where collections took place (Supplementary Material Suppl. material 1), the majority mentioned were from sites that are within the forest biome (see Mucina and Rutherford 2006 for full details of South Africa's biomes and vegetation types). However, other vegetation types mentioned include those of the grassland biome and disturbed areas such as gardens, orchards and plantations.

The sample-based species rarefaction curve for the Western Cape did not reach an asymptote (Fig. 3). The two richness estimators (Jacknife2: 348 species, Chao1 with 95% Confidence Intervals: 323, lower CI: 270, upper CI: 416) suggest that at least 6–7 times more than the number of species currently recorded from the literature will be found in the province, given the steep slope of the non-asymptotic curve.
<table><caption>**Table 2.** Collembola species recorded from South Africa, with “Current species name” as confirmed name (Bellinger et al. 2014), and “Name published in source” as name used in the original source when different from current species name. Abbreviations used: South Africa (SA), Western Cape (WC), Eastern Cape (EC), Kwazulu-Natal (KZN), Gauteng (G), Limpopo (L), Free State (FS), Northern Cape (NC), Mpumalanga (MP), North West Province (NWP), Lesotho (Les), endemic (E), introduced (I), dubious record (D) or widespread (W, species present outside of South Africa but not considered introduced). Genera endemic to South Africa are indicated by an asterisk (*). See Suppl. material 1: Table S1 for full collection and citation details.</caption><thead><tr><th>Current species name</th><th>Source</th><th>Province recorded from in SA</th><th>Status</th><th>Habitat if given in source</th><th>Name published in source if different from the current one</th><th>Comments</th></tr></thead><tbody><tr><td><strong>PODUROMORPHA</strong></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td><strong>Hypogastruridae</strong></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Acherontiella thibaudi<br>Barra, 1994</td><td>Barra 1994</td><td>KZN</td><td>W</td><td>Beach sand</td><td></td><td>South Africa and several tropical regions of East Africa and Southeast Asia (Thibaud 2010)</td></tr><tr><td>Astrogastrura lobata<br>(Yosii, 1959)</td><td>Yosii 1959</td><td>WC</td><td>E</td><td></td><td></td><td></td></tr><tr><td>Ceratophysella armata<br>(Nicolet, 1842)</td><td>Womersley 1934, Paclt 1959, 1967, Coates 1970</td><td>WC, KZN, EC, FS, G, NC</td><td>D</td><td>Damp soil, moss, litter</td><td>Hypogastrura armata Nicolet, 1842</td><td>Western palaearctic distribution.</td></tr><tr><td>Ceratophysella armata trispina<br>(Womersley, 1934)</td><td>Womersley 1934</td><td>WC</td><td>D</td><td></td><td>Hypogastrura armata var. trispina Womersley, 1934</td><td>Described from a single specimen with three anal spines, could also have been <i>Triacanthella</i> sp.</td></tr><tr><td>Ceratophysella longispina<br>(Tullberg, 1876)</td><td>Womersley 1934</td><td>NC, KZN,</td><td>D</td><td></td><td>Hypogastrura longispina Tullberg, 1876</td><td>Northern hemisphere circumpolar distribution (Fjellberg, 1998)</td></tr><tr><td>Hypogastrura manubrialis<br>(Tullberg, 1876)</td><td>Womersley 1934, Paclt 1959, 1967</td><td>NC, KZN, WC</td><td>I</td><td>Wet habitat</td><td></td><td>Distributed worldwide, considered introduced in the Southern hemisphere</td></tr><tr><td>Hypogastrura manubrialis<br>neglectus (Börner, 1901)</td><td>Womersley 1934</td><td>WC</td><td>D</td><td></td><td>Hypogastrura manubrialis var. neglectus (Börner, 1901)</td><td>Dubious: lacks two anal spines, no more information provided.</td></tr><tr><td>Hypogastrura purpureoscens<br>(Lubbock, 1868)</td><td>Womersley 1934, Paclt 1959, 1967</td><td>WC</td><td>I</td><td>Wet leaves</td><td>Hypogastrura pseudopurpureoscens Womersley, 1928 in Womersley 1934 Hypogastrura (Hypogastrura) purpureoscens (Lubbock, 1868) in Paclt 1959, 1967</td><td>The species can be considered as introduced from Northern hemisphere, as has recently been confirmed for Australia (Greenslade et al. 2014).</td></tr><tr><td>Hypogastrura sahlbergi<br>(Reuter, 1895)</td><td>Paclt 1959</td><td>WC</td><td>D</td><td>Near stream</td><td></td><td>Dubious record: holarctic distribution (Bellinger et al. 1996–2014).</td></tr><tr><td>Hypogastrura sahlbergi rosea<br>(Reuter, 1895)</td><td>Womersley 1934</td><td>WC</td><td>D</td><td>Damp rocks</td><td>Hypogastrura sahlbergi var. rosea (Reuter, 1895)</td><td>Agrees with <i>sahlbergi</i> s. str. except for colour. <i>Species inquirenda</i>.</td></tr></tbody></table>
<table><thead><tr><th>Current species name</th><th>Source</th><th>Province recorded from in SA</th><th>Status</th><th>Habitat if given in source</th><th>Name published in source if different from the current one</th><th>Comments</th></tr></thead><tbody><tr><td>Hypogastrum viatica (Tullberg, 1872)</td><td>Womersley 1934, Paclt 1959</td><td>WC</td><td>I</td><td>Littoral</td><td></td><td>Nordic countries and Arctic, considered introduced in southern hemisphere (Greenslade 2002).</td></tr><tr><td>Mesogastura libyca (Caroli, 1914)</td><td>Paclt 1959</td><td>WC</td><td>D</td><td>Forest litter</td><td>Choreutinula libyca Caroli, 1914</td><td>Probably Austrogastrura lobata (Yosii, 1959), present in the same locality.</td></tr><tr><td>Triacanbella madiba Janion, D'Haeze & Deharveng, 2012</td><td>Janion et al. 2012</td><td>WC</td><td>E</td><td>Cave guano</td><td></td><td></td></tr><tr><td>Willemia tribobata Barra, 1995</td><td>Barra 1995</td><td>KZN</td><td>E</td><td>Beach sand</td><td></td><td></td></tr><tr><td>Xenylla capensis Weiner & Naji, 1991</td><td>Weiner and Naji 1991</td><td>WC</td><td>E</td><td>Forest leaf litter</td><td></td><td></td></tr><tr><td>Xenylla maritima Tullberg, 1869</td><td>Paclt 1959, 1967</td><td>WC, EC, KZN, NWP</td><td>I</td><td>Wet habitat, forest</td><td></td><td>Cosmopolitan distribution (Fjellberg 1998), probably introduced in the Southern hemisphere</td></tr><tr><td>Xenylla rhodesiensis Womersley, 1929</td><td>Coates 1970</td><td>MP</td><td>E</td><td>Wet habitat</td><td></td><td>Only recorded from Europe, while the collection locality in South Africa is very isolated and at a high altitude</td></tr><tr><td>Xenylla schillei Bömer, 1903</td><td>Paclt 1959</td><td>Les</td><td>D</td><td>At stream</td><td></td><td>Pan-tropical distribution (Deharveng et al. 2011)</td></tr><tr><td>Xenylla yucatana Mills, 1938</td><td>Barra 1995</td><td>KZN</td><td>W</td><td>Forest soil</td><td></td><td></td></tr><tr><td><strong>Brachystomellidae</strong></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Brachystomella africana Yosii, 1959</td><td>Yosii 1959</td><td>WC</td><td>E</td><td></td><td>Brachystomella parvula africana Yosii, 1959</td><td></td></tr><tr><td>Brachystomella coatesi Weiner & Naji, 1991</td><td>Weiner and Naji 1991</td><td>WC</td><td>E</td><td>Forest leaf litter</td><td></td><td></td></tr><tr><td>Brachystomella georgenis Weiner & Naji, 1991</td><td>Weiner and Naji 1991</td><td>WC</td><td>E</td><td>Forest leaf litter</td><td></td><td></td></tr><tr><td>Brachystomella parvula (Schäffer, 1896)</td><td>Womersley 1934, Paclt 1959, 1967, Coates 1970</td><td>MP WC, KZN, EC, FS</td><td>I</td><td>Wet litter</td><td></td><td>Cosmopolitan distribution (Fjellberg 1998)</td></tr><tr><td>Probrachystomellides nicolai Weiner & Naji, 1991*</td><td>Weiner and Naji 1991</td><td>WC</td><td>E</td><td>Forest leaf litter</td><td></td><td></td></tr><tr><td>Setanodosa capitata (Womersley, 1934)</td><td>Womersley 1934</td><td>WC</td><td>E</td><td></td><td>Brachystomella capitata Womersley, 1934</td><td></td></tr><tr><td><strong>Neanuridae</strong></td><td></td><td></td><td></td><td></td><td></td><td></td></tr></tbody></table>
<table><thead><tr><th>Current species name</th><th>Source</th><th>Province recorded from in SA</th><th>Status</th><th>Habitat if given in source</th><th>Name published in source if different from the current one</th><th>Comments</th></tr></thead><tbody><tr><td><i>Aethiopella capensis</i> (Womersley, 1934)</td><td>Womersley 1934, Pacht 1959</td><td>WC, KZN</td><td>E</td><td>Stony stream</td><td><i>Ceratrimeria flavoantennatus</i> var. <i>capensis</i> Womersley, 1934</td><td>Described and previously only known from Ethiopia (Massoud 1967)</td></tr><tr><td><i>Aethiopella handachini</i> (Denis, 1924)</td><td>Pacht 1959</td><td>Les, WC</td><td>D</td><td>Under stone, litter</td><td></td><td></td></tr><tr><td><i>Anurida maritima</i> (Cúérin-Méneville, 1836)</td><td>Womersley 1934, Pacht 1959, Yosii 1959, Lawrence 1953</td><td>WC, KZN</td><td>W</td><td>Littoral</td><td></td><td>Cosmopolitan distribution (Fjellberg 1998)</td></tr><tr><td><i>Ecitonura barni</i> Janion, Bedos & Deharveng, 2011</td><td>Janion et al. 2011b</td><td>WC</td><td>E</td><td>Forest leaf litter</td><td></td><td></td></tr><tr><td><i>Ecitonura coatesi</i> Barra, 1994</td><td>Barra 1994</td><td>KZN</td><td>E</td><td>Litter on dunes</td><td></td><td></td></tr><tr><td><i>Ecitonura monochaeta</i> Janion, Bedos & Deharveng, 2011</td><td>Janion et al. 2011b</td><td>WC</td><td>E</td><td>Forest leaf litter</td><td></td><td></td></tr><tr><td><i>Ecitonura natalensis</i> (Womersley, 1934)</td><td>Womersley 1934, Pacht 1959</td><td>KZN, WC, EC</td><td>E</td><td>Litter</td><td><i>Achorutes natalensis</i> Womersley, 1934<br><i>Neanura natalensis</i> Womersley, 1934)</td><td></td></tr><tr><td><i>Ecitonura oribiensis</i> (Coates, 1968)</td><td>Coates 1968</td><td>KZN</td><td>E</td><td>Soil, litter</td><td><i>Neanura oribiensis</i> Coates, 1968</td><td></td></tr><tr><td><i>Friesea claviseta</i> Axelson, 1900</td><td>Womersley 1934</td><td>KZN, WC</td><td>I</td><td>Litter</td><td></td><td>Cosmopolitan, possibly introduced in the southern hemisphere</td></tr><tr><td><i>Friesea versabilis</i> Barra, 1995</td><td>Barra 1995</td><td>KZN</td><td>W</td><td>Under vegetation</td><td></td><td>Recorded from South Africa and Madagascar (Thibaud 2008)</td></tr><tr><td><i>Naja Africa riebi</i> (Barra, 1994)*</td><td>Barra 1994</td><td>KZN</td><td>E</td><td>Dune litter</td><td><i>Stachorutes riebi</i> Barra, 1994</td><td></td></tr><tr><td><i>Neanura muscorum</i> (Templeton, 1835)</td><td>Coates 1968a</td><td>EC</td><td>I</td><td>Litter</td><td></td><td>Sub-cosmopolitan, introduced in the southern hemisphere. All other species of the genus are in Europe.</td></tr><tr><td><i>Pseudachorutella africana</i> Weiner & Najt, 1991</td><td>Weiner and Najt 1991</td><td>WC</td><td>E</td><td>Forest leaf litter</td><td></td><td></td></tr><tr><td><i>Pseudachorutes altaudi</i> (Delamare Deboutteville, 1946)</td><td>Pacht 1959</td><td>KZN</td><td>W</td><td>Forest leaf litter</td><td><i>Ceratrimeria altaudi</i> Delamare Deboutteville, 1946</td><td>Described and only known so far from Eastern Africa (Massoud 1967).</td></tr><tr><td><i>Pseudachorutes univesicatus</i> Weiner & Najt, 1991</td><td>Weiner and Najt 1991</td><td>WC</td><td>E</td><td>Forest leaf litter</td><td></td><td></td></tr><tr><td><i>Virronura joanna</i> (Coates, 1968)</td><td>Coates 1968a</td><td>NWNP</td><td>E</td><td>Soil</td><td><i>Neanura joanna</i> Coates, 1968</td><td></td></tr><tr><td><i>Odontellidae</i></td><td></td><td></td><td></td><td></td><td></td><td></td></tr></tbody></table>
<table><thead><tr><th>Current species name</th><th>Source</th><th>Province recorded from in SA</th><th>Status</th><th>Habitat if given in source</th><th>Name published in source if different from the current one</th><th>Comments</th></tr></thead><tbody><tr><td>Odontella sylvatica Weiner &amp; Nait, 1991</td><td>Weiner and Nait, 1991</td><td>WC</td><td>E</td><td>Forest leaf litter</td><td></td><td></td></tr><tr><td>Odontellina deharvengi Barra, 1995</td><td>Barra 1995</td><td>KZN</td><td>W</td><td>Soil</td><td></td><td>Recorded from South Africa and Madagascar (Thibaud 2008)</td></tr><tr><td>Superodontella empodialis (Stach, 1934)</td><td>Pact 1959</td><td>KZN</td><td>D</td><td></td><td>Odontella empodialis Stach, 1934</td><td>Dubious identification, European distribution</td></tr><tr><td colspan="7"><strong>Onychiuridae</strong></td></tr><tr><td>Deuteraphorura inermis (Tullberg, 1869)</td><td>Womersley 1934, Pact 1959</td><td>WC</td><td>D</td><td>Under stones</td><td>Onychiurus fimetarius (Linné, Lubbock) (sic) in Womersley 1934<br>Onychiurus peudinermis Börner, Börner 1903 in Pact 1959</td><td>Given the confusion around the species <i>fimetarius</i>, <i>inermis</i> and <i>pseudinermis</i>, and the age of the specimen slides, the identification given by authors (following Bellinger et al. 1996-2014) is uncertain. The Schött description is insufficient to recognize the species.</td></tr><tr><td>Orthonychiurus camerunensis (Schött, 1926)</td><td>Pact 1967</td><td>G</td><td>D</td><td>Soil</td><td>Onychiurus saasveldensis Weiner &amp; Nait, 1926</td><td></td></tr><tr><td>Orthonychiurus saasveldensis (Weiner &amp; Nait, 1991)</td><td>Weiner and Nait, 1991</td><td>WC</td><td>E</td><td>Forest, on bark</td><td>Onychiurus armatus</td><td>A holarctic distribution. Southern records of <i>Protaiphorura</i> are usually <i>Thalassaphorura</i> species, or possible introductions.</td></tr><tr><td>Protaiphorura armata (Tullberg, 1869)</td><td>Lawrence 1953</td><td>?</td><td>D</td><td></td><td>Onychiurus armatus</td><td>A species inquirenda after Yosii (1977), only recorded so far from Japan.</td></tr><tr><td>Protaiphorura matsumotoi (Kinoshita, 1923)</td><td>Pact 1959</td><td>FS</td><td>D</td><td>Soil</td><td>Onychiurus matsumotoi Kinoshita, 1923</td><td></td></tr><tr><td colspan="7"><strong>Tullbergiidae</strong></td></tr><tr><td>Delamarephorura capensis Janion, Weiner &amp; Deharveng, 2013</td><td>Janion et al. 2013</td><td>WC</td><td>E</td><td>Soil</td><td></td><td></td></tr><tr><td>Delamarephorura szepykii Barra &amp; Weiner, 2009</td><td>Barra and Weiner 2009</td><td>EC</td><td>E</td><td>Dry grassland</td><td></td><td></td></tr><tr><td>Fisuaraphorura miscellanea Barra, 1995</td><td>Barra 1995</td><td>KZN</td><td>E</td><td>Soil</td><td></td><td></td></tr><tr><td>Mesaphorura krausbaueri (Börner, 1901)</td><td>Womersley 1934, Pact 1959</td><td>WC, EC, FS</td><td>D</td><td>Soil, under stones</td><td>Tullbergia krausbaueri Börner, 1901</td><td>Dubious identification, most <i>Mesaphorura</i> have been identified as <i>M. krausbaueri</i> before the split of this species by Rusek (1971). Older records are not reliable (Fjellberg 1998).</td></tr></tbody></table>
<table><thead><tr><th>Current species name</th><th>Source</th><th>Province recorded from in SA</th><th>Status</th><th>Habitat if given in source</th><th>Name published in source if different from the current one</th><th>Comments</th></tr></thead><tbody><tr><td>Mesaphorura yosii (Rusck, 1967)</td><td>Barra 1995</td><td>KZN</td><td>W</td><td></td><td></td><td>Cosmopolitan distribution</td></tr><tr><td>Paratullergia callipygos (Börner, 1902)</td><td>Womersley 1934</td><td>WC</td><td>I</td><td></td><td>Tullbergia callipygos Börner, 1902</td><td>Holarctic distribution</td></tr><tr><td>Tullbergia meridionalis Cassagnau & Rapoport, 1962</td><td>Barra 1995</td><td>KZN</td><td>W</td><td>Dune sand</td><td></td><td>Described from Argentina and later recorded from South Africa.</td></tr><tr><td>Tullbergia kilimanjarica (Delamare Deboutteville, 1953)</td><td>Pact 1959, 1967, Coates 1970</td><td>WC, KZN, MP</td><td>W</td><td>Forest leaf litter, garden soil</td><td>Mesaphorura kilimanjarica Delamare Deboutteville, 1953</td><td>Described from Tanzania and later recorded from South Africa.</td></tr><tr><td colspan="7"><strong>ENTOMOBYMORPHA</strong></td></tr><tr><td>Isotomidae</td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Archisotoma sabudosa Barra, 1997</td><td>Barra 1997</td><td>KZN</td><td>E</td><td>Littoral dune sand</td><td></td><td></td></tr><tr><td>Arlea tridens Barra, 1997</td><td>Barra 1997</td><td>KZN</td><td>E</td><td>Dune litter</td><td></td><td></td></tr><tr><td>Ballistura seboiti (Dalla Torre, 1895)</td><td>Womersley 1934, Yosii 1959, Pact 1959, 1967</td><td>WC, EC</td><td>I</td><td>Vegetation, rain pools</td><td>Proisotoma schoetti (Dalla Torre, 1895) in Womersley 1934 and Pact 1959, 1967</td><td>Cosmopolitan distribution</td></tr><tr><td>Clavisotoma africana (Womersley, 1934)</td><td>Womersley 1934, Pact 1959</td><td>WC</td><td>E</td><td>Wet leaves, rain pools</td><td>Proisotoma africana (Womersley, 1934)</td><td>Cosmopolitan distribution</td></tr><tr><td>Folsonides americanus Denis, 1931</td><td>Pact 1959, Barra 1997</td><td>KZN</td><td>W</td><td>From dry leaves</td><td></td><td>Cosmopolitan distribution</td></tr><tr><td>Folsonina onychiirina Denis, 1931</td><td>Barra 1997</td><td>KZN</td><td>W</td><td></td><td></td><td>Pantropical distribution</td></tr><tr><td>Hemisotoma thermophila (Axelson, 1900)</td><td>Womersley 1934, Pact 1959, Coates 1970</td><td>KZN, WC</td><td>W</td><td>Under rotting leaves</td><td>Isotoma bituberudata Wahlgren, 1906 in Womersley 1934 and in Pact 1959 Isotoma thermophila (Axelson, 1900) in Coates 1970</td><td>Cosmopolitan distribution. Isotoma bituberudata is proposed as a synonym of either Hemisotoma thermophila or H. orientalis (Stach, 1947) in Potapov (2001). We provisionally consider it as a synonym of H. thermophila, the most widespread species of the genus Hemisotoma.</td></tr><tr><td>Isotoma fuititina Scherbakov, 1899</td><td>Pact 1959</td><td>KZN</td><td>D</td><td></td><td>Sorensia fuititina (Scherbakov, 1899)</td><td>The species is described without PAO, but body pigment is present; as such, it does not fit any known genus (Potapov 2001). Species inquirenda.</td></tr></tbody></table>
<table><thead><tr><th>Current species name</th><th>Source</th><th>Province recorded from in SA</th><th>Status</th><th>Habitat if given in source</th><th>Name published in source if different from the current one</th><th>Comments</th></tr></thead><tbody><tr><td><i>Isotoma maturetanica</i><br> Handschin, 1926</td><td>Womersley 1934</td><td>WC</td><td>D</td><td></td><td></td><td><i>Species inquirenda</i>, Stach (1947) considered this Algerian species as a possible member of the genus <i>Isotomurus</i>, but the description is too brief to support such a statement. See also <i>Isotomurus palustris</i>.</td></tr><tr><td><i>Isotomiella soduana</i><br> Barra, 1997</td><td>Barra 1997</td><td>KZN</td><td>E</td><td>Litter and humus on sand dunes</td><td></td><td></td></tr><tr><td><i>Isotomodes productus</i><br> (Axelson, 1906)</td><td>Womersley 1934</td><td>WC</td><td>I</td><td>Under stones</td><td></td><td>Subcosmopolitan, records from southern hemisphere scattered.</td></tr><tr><td><i>Isotomurus balteatus</i><br> (Reuter, 1876)</td><td>Womersley 1934</td><td>WC</td><td>D</td><td></td><td><i>Isotomurus palustris</i> var. <i>balteata</i> (Reuter, 1876).</td><td><i>I. balteatus</i> is a species of Europe recognizable by its transversal stripes on tergites. We have seen such a colour pattern in South African Isotomidae of an undetermined genus which is not <i>Isotomurus</i>. The record of this species for South Africa is therefore dubious.</td></tr><tr><td><i>Isotomurus palustris</i><br> (Müller, 1776)</td><td>Womersley 1934, Paclt 1959, 1967</td><td>WC, EC, G, KZN</td><td>I</td><td></td><td></td><td>Specimens of <i>Isotoma maturetanica</i><br> Handschin, 1926 recorded in Womersley 1934 were re-identified as <i>I. palustris</i> by Paclt (1959). This change is probably wrong, as Paclt states that specimens lack bothriotrichia.</td></tr><tr><td><i>Isotomurus tricuspis</i><br> Börner, 1906</td><td>Paclt 1959, 1967</td><td>WC</td><td>D</td><td>Damp moss</td><td></td><td>Specimens of <i>Isotomurus palustris</i> var. <i>balteata</i> cited by Womersley (1934) are considered as <i>I. tricuspis</i> by Paclt (1959), based on similar pattern. of transversal stripes on tergites. However, these South African forms need to be examined morphologically to confirm their congeneric status with <i>I. tricuspis</i> from Java.</td></tr><tr><td><i>Micranurophorus musci</i><br> Bernard, 1977</td><td>Barra 1997</td><td>KZN</td><td>W</td><td>Humid sand 20 cm under pioneer vegetation</td><td></td><td>Subcosmopolitan interstitial species.</td></tr></tbody></table>
<table><thead><tr><th>Current species name</th><th>Source</th><th>Province recorded from in SA</th><th>Status</th><th>Habitat if given in source</th><th>Name published in source if different from the current one</th><th>Comments</th></tr></thead><tbody><tr><td><i>Mucrosomia caeca</i> (Wahlgren, 1906)</td><td>Paclt 1959</td><td>KZN, WC</td><td>W</td><td>From wet debris</td><td><i>Cryptopygus caecus</i> Wahlgren, 1906</td><td>Current name after Potapov (2001).</td></tr><tr><td><i>Parisotoma masopi</i> (Womersley, 1934)</td><td>Paclt 1959</td><td>FS</td><td>E</td><td>From soil containing organic material</td><td><i>Iotoma notabilis</i> ssp. masopi Womersley, 1934</td><td></td></tr><tr><td><i>Parisotoma notabilis</i> (Schäffer, 1896)</td><td>Paclt 1959, 1967</td><td>WC</td><td>I</td><td>Wet leaves, leaf litter,</td><td><i>Iotoma notabilis</i> Schäffer, 1896 in Paclt 1959, 1967</td><td></td></tr><tr><td><i>Parisotoma obsurocellata</i> Potapov, Janion &amp; Deharveng, 2011</td><td>Potapov et al. 2011</td><td>WC</td><td>E</td><td>Litter under plants, coastal</td><td></td><td></td></tr><tr><td><i>Parisotoma sexetosa</i> Potapov, Janion &amp; Deharveng, 2011</td><td>Potapov et al. 2011</td><td>WC</td><td>E</td><td>Forest leaf litter</td><td></td><td></td></tr><tr><td><i>Pauropygus caussaneli</i> (Thibaud, 1996)</td><td>Barra 1997</td><td>KZN</td><td>W</td><td>Littoral sand</td><td><i>Cryptopygus riebi</i> Barra, 1997</td><td>Synonymy after Potapov, Gao and Deharveng 2013. On the coasts of Indian and Atlantic Oceans</td></tr><tr><td><i>Proisotoma davidi</i> Barra, 2001</td><td>Barra 2001</td><td>EC.</td><td>E</td><td>Grassland soil</td><td></td><td></td></tr><tr><td><i>Proisotoma minuta</i> (Tullberg, 1871)</td><td>Paclt 1959, 1967</td><td>WC, KZN, FS, EC.</td><td>I</td><td>Litter</td><td></td><td>Cosmopolitan species.</td></tr><tr><td><b>Entomobryidae</b></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td><i>Caphrya marshalli</i> Barra, 1999*</td><td>Barra 1999</td><td>EC</td><td>E</td><td>Grassland</td><td></td><td></td></tr><tr><td><i>Caphrya themeda</i> Barra, 1999*</td><td>Barra 1999</td><td>EC</td><td>E</td><td>Grassland</td><td></td><td></td></tr><tr><td><i>Cocobrya caeca</i> (Schött, 1896)</td><td>Goto 1953</td><td>WC</td><td>D</td><td>In cave</td><td><i>Sinella caeca</i> (Schött, 1896)</td><td>C. <i>coeca</i> is restricted to northern America according to Chen and Christiansen (1997), and unlikely to have been introduced in South African caves. The South African species might be the cosmopolitan <i>C. tenebricosa</i> (Folsom, 1902) (Zhang et al. 2009)</td></tr><tr><td><i>Cocobrya haefti</i> (Schäffer, 1896)</td><td>Paclt 1959</td><td>WC</td><td>D</td><td>In cave</td><td></td><td>Extra-European records are dubious (Jordana 2012). The Paclt specimens, from the same locality as the Goto (1953) specimens, may rather belong to the cosmopolitan species <i>C. tenebricosa</i> (Zhang et al. 2009)</td></tr></tbody></table>
<table><thead><tr><th>Current species name</th><th>Source</th><th>Province recorded from in SA</th><th>Status</th><th>Habitat if given in source</th><th>Name published in source if different from the current one</th><th>Comments</th></tr></thead><tbody><tr><td>Entomobrya atrocincta Schött, 1897</td><td>Paclt 1967</td><td>WC</td><td>I?</td><td>Litter</td><td></td><td>The large distribution of the species makes it difficult to determine from which region it may have been introduced. In addition, most colour patterns described in the literature do not fit the original and clear description of Schött (1897).</td></tr><tr><td>Entomobrya decemfasciata (Packard, 1873)</td><td>Womersley 1934</td><td>WC</td><td>D</td><td></td><td></td><td>Contrary to the claim of Womersley, E. decemfasciata does not occur in "most temperate parts of the world, including Europe". Reliable records are restricted to North America. The colour pattern given by Womersley is different from that given by Christiansen and Bellinger (1998) for specimens of the USA.</td></tr><tr><td>Entomobrya lanuginosa (Nicolet, 1842)</td><td>Womersley 1934</td><td>WC</td><td>I?</td><td></td><td>Entomobrya nivalis Linnaeus, 1758 f. immaculata Schäffer, 1896</td><td>The cited form is tentatively reported to E. lanuginosa. In that case it would be an introduced species.</td></tr><tr><td>Entomobrya minima Brown, 1926</td><td>Brown 1926</td><td>KZN</td><td>E</td><td>Under stone</td><td></td><td></td></tr><tr><td>Entomobrya multifasciata (Tullberg, 1871)</td><td>Paclt 1967</td><td>WC, NC, G</td><td>I</td><td>Litter, next to stream</td><td></td><td>Widespread in the bolarctic region.</td></tr><tr><td>Entomobrya nicoleti (Lubbock, 1876)</td><td>Womersley 1934</td><td>WC</td><td>I?</td><td></td><td>Entomobrya nivalis f. maculata Schäffer, 1896</td><td>The cited form is tentatively reported to E. nicolai. In that case it would be an introduced species.</td></tr><tr><td>Entomobrya nivalis (Linnaeus, 1758)</td><td>Paclt 1959, 1967, Coates 1970</td><td>WC, EC, FS, KZN</td><td>I</td><td>Litter, rainwater pool</td><td></td><td>Cosmopolitan distribution, but most reliable records are in the bolarctic region.</td></tr><tr><td>Lepidocytus cyanens Tullberg, 1871</td><td>Paclt 1959</td><td>KZN, EC</td><td>I?</td><td>Dry leaves, damp soil</td><td></td><td>Cosmopolitan distribution, but considered introduced in southern hemisphere where other related species are absent.</td></tr><tr><td>Lepidocytus ferrugineus (Schött, 1893)</td><td>Paclt 1959</td><td>KZN</td><td>D</td><td>Dry leaves</td><td></td><td>Described from Africa, the species needs a modern redescription to be recognizable.</td></tr><tr><td>Lepidocytus lanuginosus (Gmelin, 1788)</td><td>Womersley 1934, Paclt 1967</td><td>WC</td><td>D</td><td>Litter</td><td></td><td>Records of this species from the southern hemisphere need to be checked.</td></tr></tbody></table>
<table><thead><tr><th>Current species name</th><th>Source</th><th>Province recorded from in SA</th><th>Status</th><th>Habitat if given in source</th><th>Name published in source if different from the current one</th><th>Comments</th></tr></thead><tbody><tr><td>Lepidokrugeria meyerae Coates, 1969*</td><td>Coates 1969</td><td>MP</td><td>E</td><td>Dead leaves</td><td></td><td>Assigned to the genus <i>Entomobrya</i> by Paclt (1959), today considered as an <i>Orchesella</i> (Christiansen and Bellinger 1998). All reliable records are from the USA.</td></tr><tr><td>Orchesella hexfasciata (Harvey, 1896)</td><td>Paclt 1959</td><td>FS, G</td><td>D</td><td>Litter</td><td><i>Entomobrya hexfasciata</i> Harvey, 1896</td><td>Cosmopolitan distribution, but most reliable records are in the holarctic region.</td></tr><tr><td>Pseudosinella alba (Packard, 1873)</td><td>Paclt 1959</td><td>WC, EC</td><td>I</td><td>Litter</td><td></td><td></td></tr><tr><td>Pseudosinella bigutata Barra 1997</td><td>Barra 1997</td><td>KZN</td><td>E</td><td>Sand forest litter</td><td></td><td>All reliable records of this species are from Western Europe (Gisin and Da Gama 1972), following major taxonomic changes in species delimitations introduced in the 60'</td></tr><tr><td>Pseudosinella immaculata (Lie- Pettersen, 1897)</td><td>Paclt 1959</td><td>KZN</td><td>D</td><td></td><td></td><td>Subcosmopolitan distribution, but most tropical and southern hemisphere records need confirmation.</td></tr><tr><td>Pseudosinella octopuntiata Börner, 1901</td><td>Paclt 1959</td><td>WC, FS</td><td>I?</td><td>Wet litter</td><td></td><td></td></tr><tr><td>Seira addoenis Coates, 1968</td><td>Coates 1968</td><td>EC</td><td>E</td><td>Soil and vegetation</td><td></td><td></td></tr><tr><td>Seira annuda Coates, 1968</td><td>Coates 1968, 1970</td><td>EC, WC</td><td>E</td><td>Shore vegetation</td><td></td><td></td></tr><tr><td>Seira annulicornis (Börner, 1903)</td><td>Yosii 1959, Coates 1968, 1970</td><td>WC, MP G, FS, KZN,</td><td>W</td><td></td><td><i>Seira (Lepidocryptinus annulicornis</i> (Börner, 1903) in Yosii 1959<br><i>Lepidocryptus annulipes</i>, mispelling for <i>Lepidocryptinus annulipes</i> Handschin, 1929)</td><td>African distribution</td></tr><tr><td>Seira annulipes (Handschin, 1929)</td><td>Womersley 1934</td><td>KZN, WC</td><td>W</td><td>On vegetation</td><td><i>Lepidocryptinus flavovirens</i> var. <i>annulosa</i> Wahlgren, 1906</td><td>African distribution. Redescription needed on modern standards.</td></tr><tr><td>Seira annulosa (Wahlgren, 1906)</td><td>Womersley 1934</td><td>WC</td><td>D</td><td>Shore vegetation</td><td><i>Lepidocryptinus</i> cooperi var. barnardi Wahlgren, 1934</td><td>Species previously known from Sudan; morphological features given by Wahlgren and Womersley do not allow reliable identification.</td></tr><tr><td>Seira barnardi (Womersley, 1934)</td><td>Womersley 1934, Yosii 1959, Paclt 1959, 1967, Coates 1968, 1970</td><td>WC, NWP</td><td>E</td><td>Wet leaves</td><td><i>Seira (Lepidocryptinus) barnardi</i> (Womersley, 1933) (sic)</td><td></td></tr></tbody></table>
<table><thead><tr><th>Current species name</th><th>Source</th><th>Province recorded from in SA</th><th>Status</th><th>Habitat if given in source</th><th>Name published in source if different from the current one</th><th>Comments</th></tr></thead><tbody><tr><td>Seira capensis (Womersley, 1934)</td><td>Womersley 1934, Yosii 1959, Coates 1968</td><td>WC, EC</td><td>E</td><td>On vegetation</td><td>Lepidocryptinus capensis (Womersley, 1934) Seina (Seina) capensis (Womersley, 1934) in Yosii 1959</td><td></td></tr><tr><td>Seira damerella Coates, 1968</td><td>Coates 1968, 1970</td><td>L, MP</td><td>E</td><td>Litter</td><td>Seina (Lepidocryptinus) dayi Yosii, 1959</td><td></td></tr><tr><td>Seira dayi Yosii, 1959</td><td>Yosii 1959, Coates 1968</td><td>WC</td><td>E</td><td></td><td></td><td>Also recorded from Mozambique by Coates (1968) and from Yemen by Barra (2004)</td></tr><tr><td>Seira elana Coates, 1968</td><td>Coates 1968, 1970</td><td>MP</td><td>W</td><td>From dry vegetation</td><td>Lepidocryptinus flavovirens Börner, 1903 in Womersley 1934; author should be (Börner, 1903) Seina (Seina) flavovirens (Börner, 1903) in Yosii 1959</td><td>May correspond to several whitish species of Seina.</td></tr><tr><td>Seira flavovirens (Börner, 1903)</td><td>Womersley 1934, Yosii 1959, Coates 1968</td><td>WC</td><td>D</td><td></td><td>Pseudosira grisea (Womersley, 1934)</td><td>Possibly a synonym of Seina flavovirens according to Yosii (1959)</td></tr><tr><td>Seira grisea (Womersley, 1934)</td><td>Womersley 1934, Coates 1968</td><td>WC</td><td>E</td><td>From vegetation</td><td>Pseudosira grisea var. annulata (Womersley, 1934)</td><td>The taxonomic value of this form is uncertain. This variety might be synonym of S. flavovirens after Yosii (1959).</td></tr><tr><td>Seira grisea annulata (Womersley, 1934)</td><td>Womersley 1934</td><td>WC</td><td>D</td><td></td><td>Lepidocryptinus incertus Handschin, 1926</td><td>The species has a characteristic colouration, but is only known from the Mediterranean region where it is uncommon, so unlikely to have been introduced to South Africa.</td></tr><tr><td>Seira incerta (Handschin, 1926)</td><td>Womersley 1934</td><td>WC</td><td>D</td><td>Estuary</td><td>Pseudosira (Mesina) laeta Börner, 1908</td><td></td></tr><tr><td>Seira laeta (Börner, 1908)</td><td>Börner 1908</td><td>NC</td><td>E</td><td></td><td></td><td></td></tr><tr><td>Seira lindei Coates, 1968</td><td>Coates 1968</td><td>EC, WC</td><td>E</td><td>Wet litter</td><td></td><td></td></tr><tr><td>Seira marphila Coates, 1968</td><td>Coates 1968</td><td>EC, WC</td><td>E</td><td>Litter</td><td></td><td></td></tr><tr><td>Seira matheusi Coates, 1968</td><td>Coates 1968, 1970</td><td>EC, WC</td><td>E</td><td>From vegetation</td><td></td><td></td></tr><tr><td>Seira metala Coates, 1968</td><td>Coates 1968</td><td>WC</td><td>E</td><td>Litter</td><td></td><td></td></tr><tr><td>Seira metarsisva Coates, 1968</td><td>Coates 1968</td><td>FS, NC</td><td>E</td><td>From grass</td><td>Dianantimum munroi Paclt, 1959</td><td>Transferred to Seina by Salmon (1964)</td></tr><tr><td>Seira munroi (Paclt, 1959)</td><td>Paclt 1959</td><td>NC</td><td>E</td><td>In ants' nest</td><td>Seina (Seina) nagatai Yosii, 1959</td><td></td></tr><tr><td>Seira nagatai Yosii, 1959</td><td>Yosii 1959</td><td>WC</td><td>E</td><td></td><td>Pseudosira nyassica var. pallens Börner, 1908</td><td></td></tr><tr><td>Seira pallens (Börner, 1908)</td><td>Börner 1908</td><td>NC</td><td>E</td><td></td><td></td><td></td></tr></tbody></table>
<table><thead><tr><th>Current species name</th><th>Source</th><th>Province recorded from in SA</th><th>Status</th><th>Habitat if given in source</th><th>Name published in source if different from the current one</th><th>Comments</th></tr></thead><tbody><tr><td>Seira pseudocoerulea (Denis, 1924)</td><td>Womersley 1934, Yosii 1959</td><td>WC</td><td>D</td><td>Estuary</td><td>Lepidocytinus pseudocoeruleus (Denis, 1924) in Womersley 1934</td><td>African species. A study of the chaetotaxy of Ethiopian specimens would be however necessary to confirm identification (Yosii 1959).</td></tr><tr><td>Seira rowani Yosii, 1959</td><td>Yosii 1959, Coates 1968, 1970</td><td>WC</td><td>E</td><td>On vegetation</td><td>Seira (Afroseira) rowani Yosii, 1959</td><td></td></tr><tr><td>Seira rylei Coates, 1968</td><td>Coates 1968</td><td>WC</td><td>E</td><td>On vegetation</td><td></td><td>The numerous records of this species by Pack are all dubious, and concern various endemic species of the genus. S. squamoornata is today considered to be limited to the Palaearctic region.</td></tr><tr><td>Seira squamoornata (Scherbakov, 1898)</td><td>Padt 1959, 1967</td><td>KZN, WC, FS, G, NC</td><td>D</td><td>Soil and vegetation</td><td></td><td></td></tr><tr><td>Seira tsikana Coates, 1968</td><td>Coates 1968, 1970</td><td>WC</td><td>E</td><td>Forest leaf litter</td><td></td><td></td></tr><tr><td>Seira vaneedeni Coates, 1968</td><td>Coates 1968</td><td>KZN</td><td>E</td><td>From shrub and grass</td><td></td><td></td></tr><tr><td colspan="7"><strong>Cyphoderidae</strong></td></tr><tr><td>Calobatinus rhadinopus (Börner, 1913)</td><td>Börner 1913, Packt 1967</td><td>KZN, G</td><td>E</td><td>Termite nest</td><td>Calobatella rhadinopus Börner, 1913</td><td></td></tr><tr><td>Cyphoda colura (Börner, 1908)</td><td>Börner 1908</td><td>NC</td><td>E</td><td>Termite nest</td><td>Cyphoderus colurus Börner, 1908</td><td></td></tr><tr><td>Cyphoda limboxiphia (Börner, 1913)</td><td>Börner 1913, Packt 1967</td><td>KZN, G</td><td>E?</td><td>Termite nest</td><td>Cyphoderus limboxiphitus Börner, 1913</td><td></td></tr><tr><td>Cyphoda natalensis (Börner, 1913)</td><td>Börner 1913, Womersley 1934</td><td>KZN, WC</td><td>E</td><td>Termite nest</td><td>Cyphoderus natalensis Börner, 1913</td><td></td></tr><tr><td>Cyphoderus assimilis (Börner, 1906)</td><td>Padt 1959</td><td>KZN</td><td>W</td><td>Ant nest</td><td></td><td>Cosmopolitan distribution</td></tr><tr><td>Cyphoderus bidenticulatus Parona, 1888</td><td>Börner 1913</td><td>KZN</td><td>E</td><td>Termite nest</td><td></td><td></td></tr><tr><td>Cyphoderus omoensis Delamare Deboutteville, 1945</td><td>Padt 1959, Womersley 1934</td><td>WC</td><td>D</td><td>In cave</td><td>Cyphoderus arcuatus var. aethiopicus Hanschin, 1929 in Womersley 1934, Cyphoderus arcuatus var. squamidives in Silvestri 1918</td><td>Wrong identification of Womersley after Packt (1959)</td></tr><tr><td>Cyphoderus squamidives Silvestri, 1918</td><td>Silvestri 1918, Packt 1959, 1967</td><td>KZN, WC, G</td><td>E?</td><td>Termite nest</td><td></td><td></td></tr><tr><td>Cyphoderus trinervoidis Packt, 1965</td><td>Padt 1965</td><td>G</td><td>E</td><td>Termite nest</td><td></td><td></td></tr><tr><td>Pendocyphoderus wasmanni Börner, 1913</td><td>Börner 1913, Packt 1967</td><td>KZN, G</td><td>E</td><td>Termite nest</td><td></td><td></td></tr></tbody></table>
<table><thead><tr><th>Current species name</th><th>Source</th><th>Province recorded from in SA</th><th>Status</th><th>Habitat if given in source</th><th>Name published in source if different from the current one</th><th>Comments</th></tr></thead><tbody><tr><td><strong>Paronellidae</strong></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Dicranocentruga nigromaculata (Schött, 1903)</td><td>Paclt 1959</td><td>KZN</td><td>W</td><td></td><td>Paronella nigromaculata Schött, 1903 in Paclt 1959</td><td>African species. The generic name Dicranocentruga Wray, 1953 was reactivated by Mitra (2002)</td></tr><tr><td><strong>Tomoceridae</strong></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Neophorella dubia Womersley, 1934*</td><td>Womersley 1934</td><td>WC</td><td>E</td><td></td><td></td><td></td></tr><tr><td><strong>NEELIPLEONA</strong></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td><strong>Neelidae</strong></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Megalothorax minimus (Willem, 1900)</td><td>Paclt 1967</td><td>WC</td><td>W</td><td>Damp soil, moss</td><td></td><td>Cosmopolitan species, currently in course of splitting. South Africa specimens will have to be re-examined.</td></tr><tr><td><strong>SYMPHYPLEONA</strong></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td><strong>Sminthuridae</strong></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Denisella serrosota (Börner, 1908)</td><td>Börner 1908, Paclt 1959</td><td>NC</td><td>W</td><td></td><td>Sminthurides (Stenacidia) serrosota in Börner 1908; Sminthurides (Denisiella) serrosota in Paclt 1959</td><td>African species</td></tr><tr><td><strong>Sphaeridia minimia</strong> (Schött, 1893)</td><td>Paclt 1959, 1967</td><td>FS, WC</td><td>D</td><td>From soil</td><td>Sminthurides (Sphaeridia) minimus (Schött, 1893)</td><td>S. minimia is distributed in western Africa. It is very similar, if not identical, to the cosmopolitan species S. pumilis Krausbauer, 1898. Bretfield (1999) considers that the Paclt specimens may belong to S. pumilis, but that those from Cameroon may represent distinct species. A revision of these tropical Sphaeridia is clearly needed.</td></tr><tr><td><strong>Katiannidae</strong></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Katianna kerguelensis Denis, 1947</td><td>Paclt 1959</td><td>KZN</td><td>D</td><td></td><td></td><td>The South African records of this sub-Antarctic species need confirmation.</td></tr><tr><td>Sminthurinus mine (Börner, 1907)</td><td>Womersley 1931, Paclt 1959, Paclt 1967</td><td>WC</td><td>W</td><td>Beneath vegetation</td><td>Sminthurinus terrestris Womersley, 1931</td><td>Paclt (1959) mentions differences between the two species, that are nevertheless synonymized by Greenslade (1994). Widely distributed in the southern hemisphere and in tropical Asia.</td></tr></tbody></table>
<table><thead><tr><th>Current species name</th><th>Source</th><th>Province recorded from in SA</th><th>Status</th><th>Habitat if given in source</th><th>Name published in source if different from the current one</th><th>Comments</th></tr></thead><tbody><tr><td><i>Sminthurinus niger</i> (Lubbock, 1873)</td><td>Womersley 1931, Pactl 1959</td><td>WC</td><td>I</td><td>Under loose bark</td><td></td><td>Mostly holarctic. Tropical and Australian records may be the result of introductions. The synonymy of <i>S. pallidus</i> Womersley 1931 with <i>S. terrestris</i> proposed by Pactl (1959) is based on unsufficient ground and not accepted here.</td></tr><tr><td><i>Sminthurinus pallidus</i> Womersley, 1931</td><td>Womersley 1931, Pactl 1959</td><td>WC</td><td>E</td><td>Beneath vegetation</td><td><i>Sminthurinus terrestris</i> Womersley, 1931 in Pactl 1959</td><td></td></tr><tr><td><i>Stenognathulus stenognathus</i> (Börner, 1907)</td><td>Pactl 1959</td><td>WC, KZN</td><td>W</td><td>Litter</td><td><i>Sminthurinus stenognathus</i> (Börner, 1907)</td><td>Africa and Argentina.</td></tr><tr><td><i>Dicytomidae</i></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td><i>Dicytomina africana</i> Womersley, 1931</td><td>Womersley 1931</td><td>WC</td><td>E</td><td>On vegetation</td><td><i>Dicytomina minuta</i> form <i>africana</i> Womersley, 1931</td><td>The validity of this form needs confirmation.</td></tr><tr><td><i>Dicytomina minuta</i> (O. Fabricius, 1783)</td><td>Pactl 1959, 1967</td><td>WC</td><td>I</td><td>At stream, on vegetation</td><td></td><td>Northern hemisphere, probably introduced in southern regions. Pactl considered <i>Dicytroma minuta</i> <i>f. africana</i> as identical with <i>D. minuta</i>.</td></tr><tr><td><i>Bourletiidae</i></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td><i>Bourletiella arvalis</i> (Fitch, 1863)</td><td>Pactl 1959</td><td>WC</td><td>I</td><td>Lucerne pasture</td><td><i>Bourletiella (Bourletiella) arvalis</i> (Fitch, 1863)</td><td>Northern hemisphere, with local occurrence in southern hemisphere where it has been probably introduced.</td></tr><tr><td><i>Pronastriopes barnardi</i> (Womersley, 1931)</td><td>Womersley 1931, Pactl 1959</td><td>WC</td><td>E</td><td>Amongst grass</td><td><i>Deuterosminthurus marmonatus</i> var. <i>barnardi</i> Womersley, 1931</td><td>A colour form of <i>P. marmonatus</i>. Pactl (1959) synonymized this form with <i>R. schulzei</i> on insufficient evidence.</td></tr><tr><td><i>Pronastriopes marmonatus</i> (Womersley, 1931)</td><td>Womersley 1931, Pactl 1959</td><td>WC</td><td>E</td><td>Rainwater pools</td><td><i>Deuterosminthurus marmonatus</i> Womersley, 1931</td><td>Pactl (1959) synonymized this species with <i>R. schulzei</i> on insufficient evidence. Generic assignation after Betsch (1980).</td></tr><tr><td><i>Pronastriopes schulzei</i> (Börner, 1908)</td><td>Börner 1908</td><td>WC, G, NC</td><td>E</td><td>Among vegetation, wet habitat</td><td><i>Bourletiella schulzei</i> in Börner, 1908</td><td>Pactl (1959) proposes to synonymize <i>P. marmonatus</i>, <i>P. barnardi</i> and <i>P. schulzei</i>, with <i>R. lineata</i> on weak morphological evidence as all these species are too briefly described. The same author considers in 1967 that his previous citation of <i>schulzei</i> (in Pactl 1959) as <i>Rostriopes lineatus</i> (here <i>R. lineata</i>).</td></tr></tbody></table>
<table><thead><tr><th>Current species name</th><th>Source</th><th>Province recorded from in SA</th><th>Status</th><th>Habitat if given in source</th><th>Name published in source if different from the current one</th><th>Comments</th></tr></thead><tbody><tr><td>Pronastriopes webbi Paecl, 1964</td><td>Paecl 1964, Coates 1970</td><td>KZN, MP, EC</td><td>E</td><td>On vegetation, litter</td><td></td><td></td></tr><tr><td>Rastriopes lineata Womersley, 1931</td><td>Womersley 1931, Paecl 1959, 1967</td><td>WC, NC, G</td><td>E</td><td>Under a fallen twig and on rainwater pool (Womersley 1931), on vegetation (Paecl 1959), moss and rotten leaves, grass on river banks (Paecl 1967)</td><td>Rastriopes schultzei in Paecl 1959</td><td>Paecl (1959) synonymized this species with R. schultzei, but in 1967 considered that the specimen he identified as schultzei in Paecl (1959) was in fact R. lineata, bona species.</td></tr><tr><td>Tritosminthurus schudi Snider, 1988*</td><td>Snider 1988</td><td>WC</td><td>E</td><td></td><td></td><td></td></tr><tr><td>Sminthuridae</td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Papirinus prodigiosum Yosii, 1954</td><td>Paecl 1959</td><td>KZN</td><td>D</td><td></td><td>Sphyrnatheca prodigiosa (Yosii, 1954)</td><td>The genus Papirinus, placed among Katianidae in Bretfeld (1999), is considered here as closer to Sminthuridae. This species is only known from Japan. Other species exist in Madagascar, Sumatra, Thailand and Congo. The South African species is probably new (Betsch 1980).</td></tr><tr><td>Sminthurus viridis (Linnaeus, 1758)</td><td>Lawrence 1953, Paecl 1959</td><td>WC</td><td>I</td><td>On vegetation</td><td></td><td>Mainly holarctic species, thought to have been introduced from Europe (via Australia) as eggs in soil through the importation of clover seed (Wallace 1968, Wallace and Walters 1974).</td></tr></tbody></table>
<figure><img src="image_3.png" /><figcaption>**Figure 1.** The cumulative number of Collembola species described from South Africa. The three major increases in described species are indicated by the author names (Womersley, Coates and Barra).</figcaption></figure>

**Discussion**

The number of Collembola species recorded for South Africa is low compared to well-studied regions such as Europe (Deharveng 2007), but is the highest of all African countries south of Sahara (Thibaud 2013). Low sampling intensity in Africa seems to be the main reason for this pattern. Based on new records and species discovered during recent systematic sampling in the Western Cape Province alone (Janion et al. 2011a, b, Potapov et al. 2011, Janion et al. 2012, Liu et al. 2012, Janion et al. 2013), it is clear that many species remain to be recorded and described for this province. Given low richness documented elsewhere in South Africa the same situation is likely to be the case both there and in other African countries. The spatial distribution of species richness records also suggests that incomplete sampling coverage lies at the heart of the current diversity patterns. Most records to date have come from those provinces where taxonomists were either based or hosted such as in Cape Town of the Western Cape Province (Womersley 1934, Paclt 1959, Yosii 1959), and in Pretoria of the Gauteng Province (Coates 1969), reflecting a recurrent bias in geographic patterns of diversity of poorly known groups (Deharveng et al. 2000). Although Collembola do generally prefer moist environments (Hopkin 1997), which may mean lower diversity in arid provinces such as the Northern Cape and North-West Provinces (see Mucina and Rutherford 2006),
<figure><img src="image_4.png" /><figcaption>**Figure 2.** Number of Collembola species recorded for each degree square grid in South Africa.</figcaption></figure>

low species richness in provinces such as Limpopo and Kwazulu-Natal is at odds with most other groups in the country (see e.g. Davis 1997 for dung beetles, Erasmus et al. 2000 for antlions, Foord et al. 2002 for spiders, Evans et al. 2006 for amphibians and birds, Schoeman and Foord 2012 for ants). The only exception to the poor knowledge of the fauna is for the sub-Antarctic Prince Edward Island group (consisting of Marion Island and the smaller Prince Edward Island), which is geopolitically a part of South Africa, and for which the fauna has been thoroughly investigated (Table 3, Gabriel et al. 2001, Hugo et al. 2006, Chown and Froneman 2008). Such a general situation of poor knowledge is typical for the Collembola in many parts of the world (e.g. Cicconardi et al. 2013), and will hamper efforts both to conserve this diversity (Cardoso et al. 2011) and to understand which components of it are non-indigenous and may be having impacts on the indigenous fauna (see discussion in Roques et al. 2009).

With the caveat in mind of undersampling, both in many parts of Africa and country-wide, it is worth considering what the current information on species in the country suggests. It appears that endemicity is likely to be high (currently 65%). This value is similar to that found for other invertebrate groups and plants in South Africa, with
<figure><img src="image_5.png" /><figcaption>**Figure 3.** A sample-based rarefaction curve for the Western Cape, for observed species richness, and the Chao1 and Jackknife2 Estimators.</figcaption></figure>

an extraordinary high number of endemic species found in the south-western Cape (see Colville et al. 2002, Goldblatt and Manning 2002, Herbert and Kilburn 2004, Rebelo et al. 2006, Pryke and Samways 2010). Endemicity is expected to increase with local sampling, but will likely decline if sampling is undertaken in neighbouring countries where information on the group is similarly low (e.g. Namibia, see Thibaud and Massoud 1988). Currently, sampling in the southern part of Africa mostly concerns sites within South Africa, generating a rapid increase in species richness and endemicity, as many additional endemic species have been obtained from samples as little as a few kilometres from already well sampled areas (Janion-Scheepers, Bedos and Deharveng unpublished results).

Currently, six genera are thought to be endemic to South Africa: *Najtafrica* Barra, 2002 (one species, Pseudachorutinae), *Probrachystomellides* Weiner & Najt, 1991 (one species, Brachystomellidae), *Capbrya* Barra, 1999 (two species, Entomobryidae), *Lepidokrugeria* Coates, 1969 (one species, Lepidocyrtinae), *Neophorella* Womersley, 1934 (one species, Tomoceridae) and *Tritosminthurus* Snider, 1988 (one species, Bourletielidae). *Neophorella dubia* was described from a single specimen by Womersley (1934) and is the only endemic species of the family Tomoceridae to occur in South Africa.
<table><caption>**Table 3.** Species recorded from the Prince Edward Islands, an island group geopolitically part of South Africa. Abbreviations used: E = endemic to Marion Island, S = sub-Antarctic distribution, I = introduced, D = dubious.</caption><thead><tr><th>Current species name</th><th>Source</th><th>Status</th><th>Name in source and comments</th></tr></thead><tbody><tr><td colspan="4"><strong>PODUROMORPHA</strong></td></tr><tr><td colspan="4"><strong>Hypogastruridae</strong></td></tr><tr><td>Ceratophysella denticulata (Bagnall, 1941)</td><td>Deharveng (1981)</td><td>I</td><td>Ceratophysella cf. denticulata (Bagnall, 1941)</td></tr><tr><td>Hypogastrura viatica (Tullberg, 1872)</td><td>Deharveng (1981)</td><td>D</td><td>Not found again since 1981, possible contamination (CJS pers. obs.)</td></tr><tr><td colspan="4"><strong>Neanuridae</strong></td></tr><tr><td>Friesea tilbrookii Wise, 1970</td><td>Deharveng (1981)</td><td>S</td><td>Friesea vienneyi Deharveng, 1981 (syn Greenslade 1986)</td></tr><tr><td colspan="4"><strong>Tullbergiidae</strong></td></tr><tr><td>Tullbergia bisetosa Börner, 1902</td><td>Deharveng (1981)</td><td>S</td><td></td></tr><tr><td colspan="4"><strong>ENTOMOBRYOMORPHA</strong></td></tr><tr><td colspan="4"><strong>Isotomidae</strong></td></tr><tr><td>Cryptopygus antarcticus travei Deharveng, 1981</td><td>Deharveng (1981)</td><td>E</td><td></td></tr><tr><td>Cryptopygus dubius Deharveng, 1981</td><td>Deharveng (1981)</td><td>S</td><td></td></tr><tr><td>Cryptopygus tricuspidis Enderlein, 1909</td><td>Deharveng (1981)</td><td>S</td><td></td></tr><tr><td>Folismotoma marionensis (Deharveng, 1981)</td><td>Deharveng (1981)</td><td>E</td><td>Isotoma (Sorensia) marionensis Deharveng, 1981</td></tr><tr><td>Isotomurus maculatus Müller, 1876</td><td>Deharveng (1981)</td><td>I</td><td>Isotomurus cf. palustris, confirmed as I. maculatus by Greenslade (2010)</td></tr><tr><td>Mucrosomia caeca (Wahlgren, 1906)</td><td>Deharveng (1981)</td><td>S</td><td>Cryptopygus caecus Wahlgren, 1906 (new comb. after Potapov 2001)</td></tr><tr><td>Parisotoma notabilis (Schäffer, 1896)</td><td>Deharveng (1981)</td><td>I</td><td>Isotoma (Parisotoma) notabilis</td></tr><tr><td colspan="4"><strong>Tomoceridae</strong></td></tr><tr><td>Pogonognathellus flavescens (Tullberg, 1871)</td><td>Gabriel et al. (2001)</td><td>I</td><td></td></tr><tr><td colspan="4"><strong>NEELIPLEONA</strong></td></tr><tr><td colspan="4"><strong>Neelidae</strong></td></tr><tr><td>Megalothorax minimus Willem, 1900</td><td>Deharveng (1981)</td><td>I</td><td>Megalothorax cf. minimus Willem, 1900, identification confirmed by C. Schneider (pers. comm.)</td></tr><tr><td colspan="4"><strong>SYMPHYPLEONA</strong></td></tr><tr><td colspan="4"><strong>Katiannidae</strong></td></tr><tr><td>Sminthurinus granulosus Enderlein, 1909</td><td>Deharveng (1981)</td><td>S</td><td>Sminthurinus cf. granulosus Enderlein, 1909 in Deharveng (1981)</td></tr><tr><td>Sminthurinus tuberculatus Delamare Deboutteville & Massoud, 1963</td><td>Gabriel et al. (2001)</td><td>S</td><td>Sminthurinus cf. kerguelensis Salmon, 1964 in Deharveng (1981)</td></tr><tr><td>Katianna sp.</td><td>Chown and Froneman (2008)</td><td>E</td><td></td></tr></tbody></table>

Paclt (1959) mentioned that besides the single holotype specimen, this species was not found again and he synonymised it with the Paronellidae *Dicranocentruga nigromaculata* (Schött, 1903). Ireson and Greenslade (1990) re-examined the type specimen and re-assigned the species to Tomoceridae, stressing however its similarity with Isotomidae
(Skaife 1954). In spite of intensive sampling in its type locality of Table Mountain (Janion-Scheepers, Bedos and Deharveng unpublished results), the species was not retrieved in any of our samples, and is considered here as a *species inquirenda*.

The current information also suggests that approximately 20% of the Collembola species found in South Africa may have been introduced by humans to the region and should therefore be considered alien (see Pyšek et al. 2004 for terminology). Understanding what the proportion of introduced species in the fauna actually is will depend on additional comprehensive sampling, and on further consideration of species currently though to be alien. Thus, several species resembling well-known European taxa had previously been mistakenly assigned to these taxa. For example, *Seira squamoornata*, which was originally described from the Ukraine, was thought to be a common polymorphic species in South Africa after Paclt (1959). However, Yosii (1959) did not even include this species in his list, while Coates (1968b) found that specimens labelled as one species (*S. squamoornata*) by Paclt (1959), could actually be identified as several endemic species described by Yosii (1959) or Coates (1968b), and concluded that this European species does not occur in South Africa. Indeed, to date 25 indigenous species of *Seira* have been described from South Africa (Yosii 1959, Coates 1968b), and the richness of the genus is likely much larger.

Nonetheless, that several alien species are present, especially of European origin, is not surprising given the close historical links between South Africa and Europe (Giliomee and Mbenga 2007). Most of the invasive species were collected in disturbed environments, in gardens or close to human settlements (Supplementary Material Suppl. material 1) bearing out findings for a range of other groups that disturbance may favour alien species establishment (Chytrý et al. 2005, MacDougall and Turkington 2005, Richardson and Pyšek 2006). Perhaps the best known of the alien species is *Sminthurus viridis*, also known as the Lucerne flea (Wallace 1964, Wallace and Walters 1974), which received considerable attention in South Africa during the late 1960s due to its pest status. It is thought to have arrived from Australia as eggs in soil through the importation of clover seed (Walters 1968, Wallace and Walters 1974). It was first collected in 1951 near Somerset West and by 1959 over 50 000 hectares of Lucerne were infested (Wallace and Walters 1974). The problem now appears largely to have been resolved, although the species is still listed as a pest of Lucerne (Annecke and Moran 1982).

In conclusion, based on published knowledge only, the Collembola species richness of South Africa is high compared with other African countries (Thibaud 2013), but low compared with non-African countries (Deharveng 2007) and with the richness of other invertebrate groups in the South African region (Scholtz and Chown 1995). This is likely due to undersampling, as recent discoveries (e.g. Janion et al. 2011b, Potapov et al. 2011, Janion et al. 2012, 2013) have indicated. Owing to a recent, large and comprehensive ecological and systematic study, accompanied by DNA Barcoding (Porco et al. 2012) largely focused on the country's Western Cape Province (Bengtsson et al. 2010, Janion et al. 2011a, Liu et al. 2012), a substantial increase in the number of species is expected. With 67 species recognised for the Western Cape from the recorded literature, the richness estimates indicating at least 6–7 times that
number being present, and based on experience in other undersampled countries such as Thailand (Bedos 1994), we expect that species richness for the country will exceed 1000. Improvement of systematic knowledge through studies such as these, and improvements in ecological understanding of the impacts of both landscape change and invasive species on the springtail fauna (e.g. Gabriel et al. 2001, Liu et al. 2012), will help South Africa meet its commitments to biodiversity conservation especially as set out in the 2020 Aichi Biodiversity Targets.

## Acknowledgements

Grant Duffy assisted with the production of the map. Clement Schneider identified the *Megalothorax minimus* from Marion Island. We are grateful for support from the South Africa-France bilateral grants (PROTEA I and II), and the DST-NRF Centre of Excellence for Invasion Biology. SLC is supported by ARC DP140102815.

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**Supplementary material I**

**Table S1**

Authors: Charlene Janion-Scheepers, Louis Deharveng, Anne Bedos, Steven L. Chown

Data type: occurrence

Explanation note: Collection details of Collembola recorded from continental South Africa.

Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.