Checklist
Print
Checklist
Updated list of Collembola species currently recorded from South Africa
expand article infoCharlene Janion–Scheepers, Louis Deharveng§, Anne Bedos§, Steven Chown|
‡ Stellenbosch University, Matieland, South Africa
§ Muséum National d’Histoire Naturelle, Paris, France
| Monash University, Melbourne, Australia
Open Access

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), Paclt (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 ScienceTM (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 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.

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).

Figure 2.

Number of Collembola species recorded for each degree square grid in South Africa.

A summary of the Collembola species recorded from South Africa based on the literature.

Number of species recorded from literature Number of species accepted from literature Introduced Endemic Widespread
PODUROMORPHA
Hypogastruridae 19 11 4 5 2
Brachystomellidae 6 6 1 5 0
Neanuridae 16 15 2 10 3
Odontellidae 3 2 0 1 1
Onychiuridae 5 1 0 1 0
Tullbergiidae 8 7 1 3 3
TOTAL 57 42 8 25 9
ENTOMOBRYOMORPHA
Isotomidae 23 19 5 8 6
Entomobryidae 49 36 8 25 3
Cyphoderidae 10 9 0 8 1
Paronellidae 1 1 0 0 1
Tomoceridae 1 1 0 1 0
TOTAL 84 66 13 42 11
NEELIPLEONA
Neelidae 1 1 0 0 1
TOTAL 1 1 0 0 1
SYMPHYPLEONA
Sminthurididae 2 1 0 0 1
Katiannidae 5 4 1 1 2
Dicyrtomidae 2 2 1 1 0
Bourletiellidae 7 7 1 6 0
Sminthuridae 2 1 1 0 0
TOTAL 18 15 4 8 3
TOTAL 160 124 25 75 24

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.

Figure 3.

A sample-based rarefaction curve for the Western Cape, for observed species richness, and the Chao1 and Jacknife2 Estimators.

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.

Current species name Source Province recorded from in SA Status Habitat if given in source Name published in source if different from the current one Comments
PODUROMORPHA
Hypogastruridae
Acherontiella thibaudi Barra, 1994 Barra 1994 KZN W Beach sand South Africa and several tropical regions of East Africa and Southeast Asia (Thibaud 2010)
Austrogastrura lobata (Yosii, 1959) Yosii 1959 WC E Choreutinula lobata Yosii, 1959
Ceratophysella armata (Nicolet, 1842) Womersley 1934, Paclt 1959, 1967, Coates 1970 WC, KZN, EC, FS, G, NC D Damp soil, moss, litter Hypogastrura armata Nicolet, 1842 Western palaearctic distribution.
Ceratophysella armata trispina (Womersley, 1934) Womersley 1934 WC D Hypogastrura armata var. trispina Womersley, 1934 Described from a single specimen with three anal spines, could also have been Triacanthella sp.
Ceratophysella longispina (Tullberg, 1876) Womersley 1934 NC, KZN, D Hypogastrura longispina Tullberg, 1876 Northern hemisphere circumpolar distribution (Fjellberg 1998)
Hypogastrura manubrialis (Tullberg, 1876) Womersley 1934, Paclt 1959, 1967 NC, KZN, WC I Wet habitat Distributed worldwide, considered introduced in the Southern hemisphere
Hypogastrura manubrialis neglectus (Börner, 1901) Womersley 1934 WC D Hypogastrura manubrialis var. neglectus (Börner, 1901) Dubious: lacks two anal spines, no more information provided.
Hypogastrura purpurescens (Lubbock, 1868) Womersley 1934, Paclt 1959, 1967 WC I Wet leaves Hypogastrura pseudopurpurascens Womersley, 1928 in Womersley 1934
Hypogastrura (Hypogastrura) purpurescens (Lubbock, 1868) in Paclt 1959, 1967
The species can be considered as introduced from Northern hemisphere, as has recently been confirmed for Australia (Greenslade et al. 2014).
Hypogastrura sahlbergi (Reuter, 1895) Paclt 1959 WC D Near stream Dubious record: holarctic distribution (Bellinger et al. 1996–2014).
Hypogastrura sahlbergi rosea (Reuter, 1895) Womersley 1934 WC D Damp rocks Hypogastrura sahlbergi var. rosea (Reuter, 1895) Agrees with sahlbergi s. str. except for colour. Species inquirenda.
Hypogastrura viatica (Tullberg, 1872) Womersley 1934, Paclt 1959 WC I Littoral Nordic countries and Arctic, considered introduced in southern hemisphere (Greenslade 2002).
Mesogastura libyca (Caroli, 1914) Paclt 1959 WC D Forest litter Choreutinula libyca Caroli, 1914 Probably Austrogastrura lobata (Yosii, 1959), present in the same locality.
Triacanthella madiba Janion, D’Haese & Deharveng, 2012 Janion et al. 2012 WC E Cave guano
Willemia trilobata Barra, 1995 Barra 1995 KZN E Beach sand
Xenylla capensis Weiner & Najt, 1991 Weiner and Najt 1991 WC E Forest leaf litter
Xenylla maritima Tullberg, 1869 Paclt 1959, 1967 WC, EC, KZN, NWP I Wet habitat, forest Cosmopolitan distribution (Fjellberg 1998), probably introduced in the Southern hemisphere
Xenylla rhodesiensis Womersley, 1929 Coates 1970 MP E Wet habitat
Xenylla schillei Börner, 1903 Paclt 1959 Les D At stream Only recorded from Europe, while the collection locality in South Africa is very isolated and at a high altitude
Xenylla yucatana Mills, 1938 Barra 1995 KZN W Forest soil Pan-tropical distribution (Deharveng et al. 2011)
Brachystomellidae
Brachystomella africana Yosii, 1959 Yosii 1959 WC E Brachystomella parvula africana Yosii, 1959
Brachystomella coatesi Weiner & Najt, 1991 Weiner and Najt 1991 WC E Forest leaf litter
Brachystomella georgensis Weiner & Najt, 1991 Weiner and Najt 1991 WC E Forest leaf litter
Brachystomella parvula (Schäffer, 1896) Womersley 1934, Paclt 1959, 1967, Coates 1970 MP, WC, KZN, EC, FS I Wet litter Cosmopolitan distribution (Fjellberg 1998)
Probrachystomellides nicolaii Weiner & Najt, 1991* Weiner and Najt 1991 WC E Forest leaf litter
Setanodosa capitata (Womersley, 1934) Womersley 1934 WC E Brachystomella capitata Womersley, 1934
Neanuridae
Aethiopella capensis (Womersley, 1934) Womersley 1934, Paclt 1959 WC, KZN E Stony stream Ceratrimeria flavoantennatus var. capensis Womersley, 1934
Aethiopella handschini (Denis, 1924) Paclt 1959 Les, WC D Under stone, litter Described and previously only known from Ethiopia (Massoud 1967)
Anurida maritima (Guérin-Méneville, 1836) Womersley 1934, Paclt 1959, Yosii 1959, Lawrence 1953 WC, KZN W Littoral Cosmopolitan distribution (Fjellberg 1998)
Ectonura barrai Janion, Bedos & Deharveng, 2011 Janion et al. 2011b WC E Forest leaf litter
Ectonura coatesi Barra, 1994 Barra 1994 KZN E Litter on dunes
Ectonura monochaeta Janion, Bedos & Deharveng, 2011 Janion et al. 2011b WC E Forest leaf litter
Ectonura natalensis (Womersley, 934) Womersley 1934, Paclt 1959 KZN, WC, EC E Litter Achorutes natalensis Womersley, 1934
Neanura natalensis (Womersley, 1934)
Ectonura oribiensis (Coates, 1968) Coates 1968 KZN E Soil, litter Neanura oribiensis Coates, 1968
Friesea claviseta Axelson, 1900 Womersley 1934 KZN, WC I Litter Cosmopolitan, possibly introduced in the southern hemisphere
Friesea versabilis Barra, 1995 Barra 1995 KZN W Under vegetation Recorded from South Africa and Madagascar (Thibaud 2008)
Najtafrica riebi (Barra, 1994)* Barra 1994 KZN E Dune litter Stachorutes riebi Barra, 1994
Neanura muscorum (Templeton, 1835) Coates 1968a EC I Litter Sub-cosmopolitan, introduced in the southern hemisphere. All other species of the genus are in Europe.
Pseudachorutella africana Weiner & Najt, 1991 Weiner and Najt 1991 WC E Forest leaf litter
Pseudachorutes alluaudi (Delamare Deboutteville, 1946) Paclt 1959 KZN W Forest leaf litter Ceratrimeria alluaudi Delamare Deboutteville, 1946 Described and only known so far from Eastern Africa (Massoud 1967).
Pseudachorutes univesicatus Weiner & Najt, 1991 Weiner and Najt 1991 WC E Forest leaf litter
Vitronura joanna (Coates, 1968) Coates 1968a NWP E Soil Neanura joanna Coates, 1968
Odontellidae
Odontella sylvatica Weiner & Najt, 1991 Weiner and Najt 1991 WC E Forest leaf litter
Odontellina deharvengi Barra, 1995 Barra 1995 KZN W Soil Recorded from South Africa and Madagascar (Thibaud 2008)
Superodontella empodialis (Stach, 1934) Paclt 1959 KZN D Odontella empodialis Stach, 1934 Dubious identification, European distribution
Onychiuridae
Deuteraphorura inermis (Tullberg, 1869) Womersley 1934, Paclt 1959 WC D Under stones Onychiurus fimetarius (Linné, Lubbock) (sic) in Womersley 1934
Onychiurus pseudinermis Börner, Börner 1903 in Paclt 1959
Given the confusion around the species fimetarius, inermis and pseudinermis, and the age of the specimen slides, the identification given by authors (following Bellinger et al. 1996-2014) is uncertain.
Orthonychiurus camerunensis (Schött, 1926) Paclt 1967 G D Soil Onychiurus camerunensis Schött, 1926 The Schött description is insufficient to recognize the species.
Orthonychiurus saasveldensis (Weiner & Najt, 1991) Weiner and Najt 1991 WC E Forest, on bark Onychiurus saasveldensis Weiner & Najt, 1991
Protaphorura armata (Tullberg, 1869) Lawrence 1953 ? D Onychiurus armatus A holarctic distribution. Southern records of Protaphorura are usually Thalassaphorura species, or possible introductions.
Protaphorura matsumotoi (Kinoshita, 1923) Paclt 1959 FS D Soil Onychiurus matsumotoi Kinoshita, 1923 A species inquirenda after Yosii (1977), only recorded so far from Japan.
Tullbergiidae
Delamarephorura capensis Janion, Weiner & Deharveng, 2013 Janion et al. 2013 WC E Soil
Delamarephorura szeptyckii Barra & Weiner, 2009 Barra and Weiner 2009 EC E Dry grassland
Fissuraphorura miscellanea Barra, 1995 Barra 1995 KZN E Soil
Mesaphorura krausbaueri (Börner, 1901) Womersley 1934, Paclt 1959 WC, EC, FS D Soil, under stones Tullbergia krausbaueri Börner, 1901 Dubious identification, most Mesaphorura have been identified as M. krausbaueri before the split of this species by Rusek (1971). Older records are not reliable (Fjellberg 1998).
Mesaphorura yosii (Rusek, 1967) Barra 1995 KZN W Cosmopolitan distribution
Paratullbergia callipygos (Börner, 1902) Womersley 1934 WC I Tullbergia callipygos Börner, 1902 Holarctic distribution
Tullbergia meridionalis Cassagnau & Rapoport, 1962 Barra 1995 KZN W Dune sand Described from Argentina and later recorded from South Africa.
Tullbergia kilimanjarica (Delamare Deboutteville, 1953) Paclt 1959, 1967, Coates 1970 WC, KZN, MP W Forest leaf litter, garden soil Mesaphorura kilimanjarica Delamare Deboutteville, 1953 Described from Tanzania and later recorded from South Africa.
ENTOMOBRYOMORPHA
Isotomidae
Archisotoma sabulosa Barra, 1997 Barra 1997 KZN E Littoral dune sand
Arlea tridens Barra, 1997 Barra 1997 KZN E Dune litter
Ballistura schoetti (Dalla Torre, 1895) Womersley 1934, Yosii 1959, Paclt 1959, 1967 WC, EC I Vegetation, rain pools Proisotoma schoetti (Dalla Torre, 1895) in Womersley 1934 and Paclt 1959, 1967 Cosmopolitan distribution
Clavisotoma africana (Womersley, 1934) Womersley 1934, Paclt 1959 WC E Wet leaves, rain pools Proisotoma africana (Womersley, 1934)
Folsomides americanus Denis, 1931 Paclt 1959, Barra 1997 KZN W From dry leaves Cosmopolitan distribution
Folsomina onychiurina Denis, 1931 Barra 1997 KZN W Pantropical distribution
Hemisotoma thermophila (Axelson, 1900) Womersley 1934, Paclt 1959, Coates 1970 KZN, WC W Under rotting leaves Isotoma bituberculata Wahlgren, 1906 in Womersley 1934 and in Paclt 1959
Isotomina thermophila (Axelson, 1900) in Coates 1970
Cosmopolitan distribution. Isotoma bituberculata 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.
Isotoma finitima Scherbakov, 1899 Paclt 1959 KZN D Sorensia finitima (Scherbakov, 1899) The species is described without PAO, but body pigment is present; as such, it does not fit any known genus (Potapov 2001). Species inquirenda.
Isotoma mauretanica Handschin, 1926 Womersley 1934 WC D Species inquirenda. Stach (1947) considered this Algerian species as a possible member of the genus Isotomurus, but the description is too brief to support such a statement. See also Isotomurus palustris.
Isotomiella sodwana Barra, 1997 Barra 1997 KZN E Litter and humus on sand dunes
Isotomodes productus (Axelson, 1906) Womersley 1934 WC I Under stones Subcosmopolitan, records from southern hemisphere scattered.
Isotomurus balteatus (Reuter, 1876) Womersley 1934 WC D Isotomurus palustris var. balteata (Reuter, 1876). I. balteatus 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 Isotomurus. The record of this species for South Africa is therefore dubious.
Isotomurus palustris (Müller, 1776) Womersley 1934, Paclt 1959, 1967 WC, EC, G, KZN I Specimens of Isotoma mauretanica Handschin, 1926 recorded in Womersley 1934 were re-identified as I. palustris by Paclt (1959). This change is probably wrong, as Paclt states that specimens lack bothriotrichia.
Isotomurus tricuspis Börner, 1906 Paclt 1959, 1967 WC D Damp moss Specimens of Isotomurus palustris var. balteata cited by Womersley (1934) are considered as I. tricuspis 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. tricuspis from Java.
Micranurophorus musci Bernard, 1977 Barra 1997 KZN W Humid sand 20 cm under pioneer vegetation Subcosmopolitan interstitial species.
Mucrosomia caeca (Wahlgren, 1906) Paclt 1959 KZN, WC W From wet debris Cryptopygus caecus Wahlgren, 1906 Current name after Potapov (2001).
Parisotoma mossopi (Womersley, 1934) Paclt 1959 FS E From soil containing organic material Isotoma notabilis ssp. mossopi Womersley, 1934
Parisotoma notabilis (Schäffer, 1896) Paclt 1959, 1967 WC I Wet leaves, leaf litter, Isotoma notabilis Schäffer, 1896 in Paclt 1959, 1967
Parisotoma obscurocellata Potapov, Janion & Deharveng, 2011 Potapov et al. 2011 WC E Litter under plants, coastal
Parisotoma sexsetosa Potapov, Janion & Deharveng, 2011 Potapov et al. 2011 WC E Forest leaf litter
Pauropygus caussaneli (Thibaud, 1996) Barra 1997 KZN W Littoral sand Cryptopygus riebi Barra, 1997 Synonymy after Potapov, Gao and Deharveng 2013. On the coasts of Indian and Atlantic Oceans
Proisotoma davidi Barra, 2001 Barra 2001 EC E Grassland soil
Proisotoma minuta (Tullberg, 1871) Paclt 1959, 1967 WC, KZN, FS, EC I Litter Cosmopolitan species.
Entomobryidae
Capbrya marshalli Barra, 1999* Barra 1999 EC E Grassland
Capbrya themeda Barra, 1999* Barra 1999 EC E Grassland
Coecobrya caeca (Schött, 1896) Goto 1953 WC D In cave Sinella coeca (Schött, 1896) C. coeca 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 C. tenebricosa (Folsom, 1902) (Zhang et al. 2009)
Coecobrya hoefti (Schäffer, 1896) Paclt 1959 WC D In cave 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 C. tenebricosa (Zhang et al. 2009)
Entomobrya atrocincta Schött, 1897 Paclt 1967 WC I? Litter 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).
Entomobrya decemfasciata (Packard, 1873) Womersley 1934 WC D 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.
Entomobrya lanuginosa (Nicolet, 1842) Womersley 1934 WC I? Entomobrya nivalis Linnaeus, 1758 f. immaculata Schäffer, 1896 The cited form is tentatively reported to E. lanuginosa. In that case it would be an introduced species.
Entomobrya minima Brown, 1926 Brown 1926 KZN E Under stone
Entomobrya multifasciata (Tullberg, 1871) Paclt 1967 WC, NC, G I Litter, next to stream Widespread in the holarctic region.
Entomobrya nicoleti (Lubbock, 1876) Womersley 1934 WC I? Entomobrya nivalis f. maculata Schäffer, 1896 The cited form is tentatively reported to E. nicoleti. In that case it would be an introduced species.
Entomobrya nivalis (Linnaeus, 1758) Paclt 1959, 1967, Coates 1970 WC, EC, FS, KZN I Litter, rainwater pool Cosmopolitan distribution, but most reliable records are in the holarctic region.
Lepidocyrtus cyaneus Tullberg, 1871 Paclt 1959 KZN, EC I? Dry leaves, damp soil Cosmopolitan distribution, but considered introduced in southern hemisphere where other related species are absent.
Lepidocyrtus ferrugineus (Schött, 1893) Paclt 1959 KZN D Dry leaves Described from Africa, the species needs a modern redescription to be recognizable.
Lepidocyrtus lanuginosus (Gmelin, 1788) Womersley 1934, Paclt 1967 WC D Litter Records of this species from the southern hemisphere need to be checked.
Lepidokrugeria meyerae Coates, 1969* Coates 1969 MP E Dead leaves
Orchesella hexfasciata (Harvey, 1896) Paclt 1959 FS, G D Litter Entomobrya hexfasciata Harvey, 1896 Assigned to the genus Entomobrya by Paclt (1959), today considered as an Orchesella (Christiansen and Bellinger 1998). All reliable records are from the USA.
Pseudosinella alba (Packard, 1873) Paclt 1959 WC, EC I Litter Cosmopolitan distribution, but most reliable records are in the holarctic region.
Pseudosinella biguttata Barra 1997 Barra 1997 KZN E Sand forest litter
Pseudosinella immaculata (Lie–Pettersen, 1897) Paclt 1959 KZN D 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’
Pseudosinella octopunctata Börner, 1901 Paclt 1959 WC, FS I? Wet litter Subcosmopolitan distribution, but most tropical and southern hemisphere records need confirmation.
Seira addoensis Coates, 1968 Coates 1968 EC E Soil and vegetation
Seira anncla Coates, 1968 Coates 1968, 1970 EC, WC E Shore vegetation
Seira annulicornis (Börner, 1903) Yosii 1959, Coates 1968, 1970 WC, MP, G, FS, KZN, W Seira (Lepidocyrtinus) annulicornis (Börner, 1903) in Yosii 1959 African distribution
Seira annulipes (Handschin, 1929) Womersley 1934 KZN, WC W On vegetation Lepidocyrtus annulipes, mispelling for Lepidocyrtinus annulipes Handschin, 1929 African distribution. Redescription needed on modern standards.
Seira annulosa (Wahlgren, 1906) Womersley 1934 WC D Shore vegetation Lepidocyrtinus flavovirens var. annulosa Wahlgren, 1906 Species previously known from Sudan; morphological features given by Wahlgren and Womersley do not allow reliable identification.
Seira barnardi (Womersley, 1934) Womersley 1934, Yosii 1959, Paclt 1959, 1967, Coates 1968, 1970 WC, NWP E Wet leaves Lepidocyrtinus cooperi var. barnardi Womersley, 1934
Seira (Lepidocyrtinus) barnardi (Womersley, 1933) (sic)
Seira capensis (Womersley, 1934) Womersley 1934, Yosii 1959, Coates 1968 WC, EC E On vegetation Lepidocyrtinus capensis Womersley, 1934
Seira (Seira) capensis (Womersley, 1934) in Yosii 1959
Seira damerella Coates, 1968 Coates 1968, 1970 L, MP E Litter
Seira dayi Yosii, 1959 Yosii 1959, Coates 1968 WC E Seira (Lepidocyrtinus) dayi Yosii, 1959
Seira eleana Coates, 1968 Coates 1968, 1970 MP W From dry vegetation Also recorded from Mozambique by Coates (1968) and from Yemen by Barra (2004)
Seira flavovirens (Börner, 1903) Womersley 1934, Yosii 1959, Coates 1968 WC D Lepidocyrtinus flavovirens Börner, 1903 in Womersley 1934; author should be (Börner, 1903)
Seira (Seira) flavovirens (Börner, 1903) in Yosii 1959
May correspond to several whitish species of Seira.
Seira grisea (Womersley, 1934) Womersley 1934, Coates 1968 WC E From vegetation Pseudosira grisea Womersley, 1934 Possibly a synonym of Seira flavovirens according to Yosii (1959)
Seira grisea annulata (Womersley, 1934) Womersley 1934 WC D Pseudosira grisea var. annulata Womersley, 1934 The taxonomic value of this form is uncertain. This variety might be synonym of S. flavovirens after Yosii (1959).
Seira incerta (Handschin, 1926) Womersley 1934 WC D Estuary Lepidocyrtinus incertus Handschin, 1926 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.
Seira laeta (Börner, 1908) Börner 1908 NC E Pseudosira (Mesira) laeta Börner, 1908
Seira lindei Coates, 1968 Coates 1968 EC, WC E Wet litter
Seira marephila Coates, 1968 Coates 1968 EC, WC E Litter
Seira mathewsi Coates, 1968 Coates 1968, 1970 EC, WC E From vegetation
Seira metala Coates, 1968 Coates 1968 WC E Litter
Seira metarsiosa Coates, 1968 Coates 1968 FS, NC E From grass
Seira munroi (Paclt, 1959) Paclt 1959 NC E In ants’ nest Diamantinum munroi Paclt, 1959 Transferred to Seira by Salmon (1964)
Seira nagatai Yosii, 1959 Yosii 1959 WC E Seira (Seira) nagatai Yosii, 1959
Seira pallens (Börner, 1908) Börner 1908 NC E Pseudosira nyassica var. pallens Börner, 1908
Seira pseudocoerulea (Denis, 1924) Womersley 1934, Yosii 1959 WC D Estuary Lepidocyrtinus pseudocoeruleus (Denis, 1924) in Womersley 1934 African species. A study of the chaetotaxy of Ethiopian specimens would be however necessary to confirm identification (Yosii 1959).
Seira rowani Yosii, 1959 Yosii 1959, Coates 1968, 1970 WC E On vegetation Seira (Afroseira) rowani Yosii, 1959
Seira rykei Coates, 1968 Coates 1968 WC E On vegetation
Seira squamoornata (Scherbakov, 1898) Paclt 1959, 1967 KZN, WC, FS, G, NC D Soil and vegetation The numerous records of this species by Paclt are all dubious, and concern various endemic species of the genus. S. squamoornata is today considered to be limited to the Palaearctic region.
Seira tsikama Coates, 1968 Coates 1968, 1970 WC E Forest leaf litter
Seira vaneedeni Coates, 1968 Coates 1968 KZN E From shrub and grass
Cyphoderidae
Calobatinus rhadinopus (Börner, 1913) Börner 1913, Paclt 1967 KZN, G E Termite nest Calobatella rhadinopus Börner, 1913
Cyphoda colura (Börner, 1908) Börner 1908 NC E Termite nest Cyphoderus colurus Börner, 1908
Cyphoda limboxiphia (Börner, 1913) Börner 1913, Paclt 1967 KZN, G E? Termite nest Cyphoderus limboxiphius Börner, 1913
Cyphoda natalensis (Börner, 1913) Börner 1913, Womersley 1934 KZN, WC E Termite nest Cyphoderus natalensis Börner, 1913
Cyphoderus assimilis (Börner, 1906) Paclt 1959 KZN W Ant nest Cosmopolitan distribution
Cyphoderus bidenticulatus Parona, 1888 Börner 1913 KZN E Termite nest
Cyphoderus omoensis Delamare Deboutteville, 1945 Paclt 1959, Womersley 1934 WC D In cave Cyphoderus arcuatus var. aethiopicus Hanschin, 1929 in Womersley 1934, Wrong identification of Womersley after Paclt (1959)
Cyphoderus squamidives Silvestri, 1918 Silvestri 1918, Paclt 1959, 1967 KZN, WC, G E? Termite nest Cyphoderus arcuatus var. squamidives in Silvestri 1918
Cyphoderus trinervoidis Paclt, 1965 Paclt 1965 G E Termite nest
Pseudocyphoderus wasmanni Börner, 1913 Börner 1913, Paclt 1967 KZN, G E Termite nest
Paronellidae
Dicranocentruga nigromaculata (Schött, 1903) Paclt 1959 KZN W Paronella nigromaculata Schött, 1903 in Paclt 1959 African species. The generic name Dicranocentruga Wray, 1953 was reactivated by Mitra (2002)
Tomoceridae
Neophorella dubia Womersley, 1934* Womersley 1934 WC E
NEELIPLEONA
Neelidae
Megalothorax minimus (Willem, 1900) Paclt 1967 WC W Damp soil, moss Cosmopolitan species, currently in course of splitting. South Africa specimens will have to be re-examined.
SYMPHYPLEONA
Sminthurididae
Denisiella serroseta (Börner, 1908) Börner 1908, Paclt 1959 NC W Sminthurides (Stenacidia) serroseta in Börner 1908; Sminthurides (Denisiella) serroseta in Paclt 1959 African species
Sphaeridia minima (Schött, 1893) Paclt 1959, 1967 FS, WC D From soil Sminthurides (Sphaeridia) minimus (Schött, 1893) S. minima is distributed in western Africa. It is very similar, if not identical, to the cosmopolitan species S. pumilis Krausbauer, 1898. Bretfeld (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.
Katiannidae
Katianna kerguelenensis Denis, 1947 Paclt 1959 KZN D The South African records of this sub-Antarctic species need confirmation.
Sminthurinus mime (Börner, 1907) Womersley 1931, Paclt 1959, Paclt 1967 WC W Beneath vegetation Sminthurinus terrestris Womersley, 1931 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.
Sminthurinus niger (Lubbock, 1873) Womersley 1931, Paclt 1959 WC I Under loose bark Mostly holarctic. Tropical and Australian records may be the result of introductions.
Sminthurinus pallidus Womersley, 1931 Womersley 1931, Paclt 1959 WC E Beneath vegetation Sminthurinus terrestris Womersley, 1931 in Paclt 1959 The synonymy of S. pallidus Womersley 1931 with S. terrestris proposed by Paclt (1959) is based on unsufficient ground and not accepted here.
Stenognathellus stenognathus (Börner, 1907) Paclt 1959 WC, KZN W Litter Sminthurinus stenognathus (Börner, 1907) Africa and Argentina.
Dicyrtomidae
Dicyrtomina africana Womersley, 1931 Womersley 1931 WC E On vegetation Dicyrtomina minuta form africana Womersley, 1931 The validity of this form needs confirmation.
Dicyrtomina minuta (O. Fabricius, 1783) Paclt 1959, 1967 WC I At stream, on vegetation Northern hemisphere, probably introduced in southern regions. Paclt considered Dicyrtoma minuta f. africana as identical with D. minuta.
Bourletiellidae
Bourletiella arvalis (Fitch, 1863) Paclt 1959 WC I Lucerne pasture Bourletiella (Bourletiella) arvalis (Fitch, 1863) Northern hemisphere, with local occurrence in southern hemisphere where it has been probably introduced.
Prorastriopes barnardi (Womersley, 1931) Womersley 1931, Paclt 1959 WC E Amongst grass Deuterosminthurus marmoratus var. barnardi Womersley, 1931 A colour form of P. marmoratus. Paclt (1959) synonymized this form with R. schultzei on insufficient evidence.
Prorastriopes marmoratus (Womersley, 1931) Womersley 1931, Paclt 1959 WC E Rainwater pools Deuterosminthurus marmoratus Womersley, 1931 Paclt (1959) synonymized this species with R. schultzei on insufficient evidence.
Prorastriopes schultzei (Börner, 1908) Börner 1908 WC, G, NC E Among vegetation, wet habitat Bourletiella schultzei in Börner, 1908 Generic assignation after Betsch (1980). Paclt (1959) proposes to synonymize P. marmoratus, P. banardi and P. schultzei, with R. lineata on weak morphological evidence as all these species are too briefly described. The same author considers in 1967 that his previous citation of schultzei (in Paclt 1959) as Rastriopes lineatus (here R. lineata).
Prorastriopes webbi Paclt, 1964 Paclt 1964, Coates 1970 KZN, MP, EC E On vegetation, litter
Rastriopes lineata Womersley, 1931 Womersley 1931, Paclt 1959, 1967 WC, NC, G E Under a fallen twig and on rainwater pool (Womersley 1931), on vegetation (Paclt 1959), moss and rotten leaves, grass on river banks (Paclt 1967) Rastriopes schultzei in Paclt 1959 Paclt (1959) synonymized this species with R. schultzei, but in 1967 considered that the specimen he identified as schultzei in Paclt (1959) was in fact R. lineata, bona species.
Tritosminthurus schuhi Snider, 1988* Snider 1988 WC E
Sminthuridae
Papirinus prodigiosum Yosii, 1954 Paclt 1959 KZN D Sphyrotheca prodigiosa (Yosii, 1954) The genus Papirinus, placed among Katiannidae 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).
Sminthurus viridis (Linnaeus, 1758) Lawrence 1953, Paclt 1959 WC I On vegetation 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).

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), 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).

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.

Current species name Source Status Name in source and comments
PODUROMORPHA
Hypogastruridae
Ceratophysella denticulata (Bagnall, 1941) Deharveng (1981) I Ceratophysella cf. denticulata (Bagnall, 1941)
Hypogastrura viatica (Tullberg, 1872) Deharveng (1981) D Not found again since 1981, possible contamination (CJS pers. obs.)
Neanuridae
Friesea tilbrooki Wise, 1970 Deharveng (1981) S Friesea viennei Deharveng, 1981 (syn Greenslade 1986)
Tullbergiidae
Tullbergia bisetosa Börner, 1902 Deharveng (1981) S
ENTOMOBRYOMORPHA
Isotomidae
Cryptopygus antarcticus travei Deharveng, 1981 Deharveng (1981) E
Cryptopygus dubius Deharveng, 1981 Deharveng (1981) S
Cryptopygus tricuspis Enderlein, 1909 Deharveng (1981) S
Folsomotoma marionensis
(Deharveng, 1981)
Deharveng (1981) E Isotoma (Sorensia) marionensis Deharveng, 1981
Isotomurus maculatus Müller, 1876 Deharveng (1981) I Isotomurus cf. palustris, confirmed as I. maculatus by Greenslade (2010)
Mucrosomia caeca (Wahlgren, 1906) Deharveng (1981) S Cryptopygus caecus Wahlgren, 1906 (new comb. after Potapov 2001)
Parisotoma notabilis (Schäffer, 1896) Deharveng (1981) I Isotoma (Parisotoma) notabilis
Tomoceridae
Pogonognathellus flavescens (Tullberg, 1871) Gabriel et al. (2001) I
NEELIPLEONA
Neelidae
Megalothorax minimus Willem, 1900 Deharveng (1981) I Megalothorax cf. minimus Willem, 1900, identification confirmed by C. Schneider (pers. comm.)
SYMPHYPLEONA
Katiannidae
Sminthurinus granulosus Enderlein, 1909 Deharveng (1981) S Sminthurinus cf. granulosus Enderlein, 1909 in Deharveng (1981)
Sminthurinus tuberculatus Delamare Deboutteville & Massoud, 1963 Gabriel et al. (2001) S Sminthurinus cf. kerguelensis Salmon, 1964 in Deharveng (1981)
Katianna sp. Chown and Froneman (2008) E

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 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, Bourletiellidae). 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. 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.

References

  • Abrantes EA, Bellini BC, Bernardo AN, Fernandes LH, Mendonça MC, Oliveira EP, Queiroz GC, Sautter KD, Silveira TC, Zeppelini D (2010) Synthesis of Brazilian Collembola: an update to the species list. Zootaxa 2388: 1–22. http://www.mapress.com/zootaxa/2010/f/z02388p022f.pdf
  • Abrantes EA, Bellini BC, Bernardo AN, Fernandes LH, Mendonça MC, Oliveira EP, Queiroz GC, Sautter KD, Silveira TC, Zeppelini D (2012) Errata Corrigenda and update for the ‘Synthesis of Brazilian Collembola: an update to the species list.’ ABRANTES et al. (2010), Zootaxa, 2388: 1–22. Zootaxa 3168: 1–21. http://www.mapress.com/zootaxa/2012/f/z03168p021f.pdf
  • Annecke DP, Moran VC (1982) Insects and mites of cultivated plants in South Africa. Butterworths, Durban.
  • Barra JA (1994) Nouveaux Collemboles Poduromorphes de la Province du Natal (Rép. Sud Africaine) (Insecta: Collembola). Journal of African Zoology 108: 181–189.
  • Barra JA (1995) Nouveaux Collemboles Poduromorphes des sables littoraux (partie terrestre) de la Province du Natal (Rép. Sud Africaine) (Insecta: Collembola). Journal of African Zoology 109: 125–139.
  • Barra JA (1997) Nouveaux Collemboles Entomobryomorphes des sables littoraux (partie terrestre) de la Province du Natal (Rép. Sud Africaine) (Insecta : Collembola). Journal of African Zoology 111: 465–480.
  • Barra JA (1999) Un nouveau genre Capbrya avec deux nouvelles espèces de la Province du Cap (Rep. Sud Africaine) (Collembola: Entomobryidae). Bulletin de l’Institut Royal des Sciences Naturelles de Belgique, Entomologie 69: 19–24.
  • Barra JA (2001) Proisotoma davidi sp. n. from Cape Province (South Africa) (Collembola). Deutsche Entomologische Zeitschrift 48: 23–26. doi: 10.1002/mmnd.4800480103
  • Barra JA (2002) Un nouveau genre de Pseudachorutinae (Collembola) des sables littoraux de la Province du Natal (République Sud africaine). Zoosystema 24: 177–180.
  • Barra JA, Weiner WM (2009) A new species of Delamarephorura Weiner & Najt, 1999 (Collembola,Tullbergiidae) from Cape Province (South Africa). Acta Zoologica Cracoviensia 52B: 57–60. doi: 10.3409/azc.52b_1-2.57-60
  • Bellinger PF, Christiansen KA, Janssens F (1996–2014) Checklist of the Collembola of the World. http://www.collembola.org [accessed 10 November 2014]
  • Bedos A (1994) Les Collemboles édaphiques du massif du Doi Inthanon (Thailande): biodiversité et écologie en forêt tropicale. PhD thesis, Toulouse III University, Toulouse, France.
  • Bengtsson J, Janion C, Chown SL, Leinaas HP (2011) Variation in decomposition rates in the fynbos biome, South Africa: the role of plant species and plant stoichiometry. Oecologia 165: 225–235. doi: 10.1007/s00442-010-1753-7
  • Bengtsson J, Janion C, Chown SL, Leinaas HP (2012) Litter decomposition in fynbos vegetation, South Africa. Soil Biology and Biochemistry 47: 100–105. doi: 10.1016/j.soilbio.2011.11.023
  • Berlese JE (1905) Apparecchio per racogliere press ed in gran numiro piccoli artropodi. Redia 2: 85–89.
  • Betsch JM (1980) Eléments pour une monographie des Collemboles Symphypléones (Hexapodes, Apterygotes). Mémoires du Muséum national d'Histoire naturelle 116: 1–227.
  • Biggs R, Simons H, Bakkenes M, Scholes RJ, Eickhout B, van Vuuren D, Alkemade R (2008) Scenarios of biodiversity loss in southern Africa in the 21st century. Global Environmental Change 18: 296–309. doi: 10.1016/j.gloenvcha.2008.02.001
  • Börner C (1908) Collembolen aus Südafrika, nebst einer Studie über die I. Maxille der Collembolen. In: Schultze L (Ed.) Forschungsreise im westlichen und zentralen Südafrika. Denkschriften der Medicinisch-naturwissenschaftlichen Gesellschaft zu Jena 13: 53–68.
  • Börner C (1913) Neue Cyphoderinen. Zoologisher Anzeiger 41: 274–284.
  • Bretfeld G (1999) Symphypleona. In: Dunger W (Ed.) Synopses on Palaearctic Collembola, Vol. 2. Abhandlungen und Berichte des Naturkundemuseums Görlitz 71: 1–318.
  • Brown JM (1926) Some African Apterygota. Annals and Magazine of Natural History 9: 34–44.
  • Butchart SHM, Walpole M, Collen B, van Strien A, Scharlemann JPW, Almond REA, Baillie JEM, Bomhard B, Brown C, Bruno J, Carpenter KE, Carr GM, Chanson J, Chenery AM, Csirke J, Davidson NC, Dentener F, Foster M, Galli A, Galloway JN, Genovesi P, Gregory RD, Hockings M, Kapos V, Lamarque J-F, Leverington F, Loh J, McGeoch MA, McRae L, Minasyan A, Morcillo MH, Oldfield TEE, Pauly D, Quader S, Revenga C, Sauer JR, Skolnik B, Spear D, Stanwell-Smith D, Stuart SN, Symes A, Tierney M, Tyrrell TD, Vié J-C, Watson R (2010) Global biodiversity: indicators of recent declines. Science 328: 1164–1168. doi: 10.1126/science.1187512
  • Cardoso P, Erwin TL, Borges PAV, New TR (2011) The seven impediments in invertebrate conservation and how to overcome them. Biological Conservation 14: 2647–2655. doi: 10.1016/j.biocon.2011.07.024
  • Chen JX, Christiansen K (1997) Subgenus Coecobrya of the genus Sinella (Collembola: Entomobryidae) with special reference to the species of China. Annals of the Entomological Society of America 90: 1–19. doi: 10.1093/aesa/90.1.1
  • Christiansen K, Bellinger P (1998) The Collembola of North America north of the Rio Grande, second edition. Grinnell College, Grinnell, IA, 1520 pp.
  • Chown SL (2010) Temporal biodiversity change in transformed landscapes: a southern African perspective. Philosophical Transactions Royal Society B 365: 3729–3742. doi: 10.1098/rstb.2010.0274
  • Chown SL, Froneman PW (2008) The Prince Edward Islands. Land-sea Interactions in a changing climate. African Sun Media, Stellenbosch.
  • Chytrý M, Pyšek P, Tichý L, Knollová I, Danihelka J (2005) Invasions by alien plants in the Czech Republic: a quantitative assessment across habitats. Preslia 77: 339–354. http://www.preslia.cz/P054CChy.pdf
  • Cicconardi F, Fanciulli PP, Emerson BC (2013) Collembola, the biological species concept and the underestimation of global species richness. Molecular Ecology 22: 5382–5396. doi: 10.1111/mec.12472
  • Coates TJ (1970) Check-list of the Collembola of South African Parks. Koedoe 13: 181–184. doi: 10.4102/koedoe.v13i1.740
  • Colville J, Picker MD, Cowling RM (2002) Species turnover of monkey beetles (Scarabaeidae: Hopliini) along environmental and disturbance gradients in the Namaqualand region of the succulent Karoo, South Africa. Biodiversity and Conservation 11: 243–264. doi: 10.1023/A:1014520226989
  • Costello MJ, May RM, Stork NE (2013) Can we name Earth’s species before they go extinct? Science 339: 413–416. doi: 10.1126/science.1230318
  • Culik MP, Zeppelini D (2003) Diversity and distribution of Collembola (Arthropoda: Hexapoda) of Brazil. Biodiversity Conservation 12: 1119–1143. doi: 10.1023/A:1023069912619
  • Davis ALV (1997) Climatic and biogeographical associations of southern African dung beetles (Coleoptera: Scarabaeidae s. str.). African Journal of Ecology 35: 10–38. doi: 10.1111/j.1365-2028.1997.051-89051.x
  • Deharveng L (1981) Collemboles des îles subantarctiques de l’océan Indien. Comité National Française des Recherches Antarctiques 48: 33–108.
  • Deharveng L, Lips J, Rahmadi C (2011) Focus on guano. In: Bouchet P, Le Guyader H, Pascal O (Eds) The Natural History of Santo: Caves and soils. MNHN, Paris; IRD, Marseille; PNI, Paris. Patrimoines naturels 70: 300–305.
  • Dippenaar-Schoeman A (2014) The field guide to spiders of South African. Lapa Publishers, Pretoria.
  • Dippenaar-Schoeman AS, González Reyes AX (2006) South African National Survey (SANSA): Solifugae (sun-spiders) of the national parks and reserves of South Africa (Arachnida, Solifugae). Koedoe 49: 29–38. doi: 10.4102/koedoe.v49i2.114
  • Erasmus BFN, van Jaarsveld AS, Chown SL, Kshatriya M, Wessels KJ (2002) Vulnerability of South African animals taxa to climate change. Global Change Biology 8: 679–69. doi: 10.1046/j.1365-2486.2002.00502.x
  • ESRI (Environmental Systems Resource Institute) (2011) ArcGIS Desktop Release 10.1. ESRI, Redlands, California.
  • Evans KL, Rodrigues ASL, Chown SL, Gaston KJ (2006) Protected areas and regional avian species richness in South Africa. Biology Letters 2: 184–188. doi: 10.1098/rsbl.2005.0435
  • Foord SH, Dippenaar-Schoeman AS, Haddad CH (2011) South African spider diversity: African perspectives on the conservation of a mega-diverse group. In: Grillo O (Ed.) Changing Diversity in Changing Environment. InTech, Croatia. doi: 10.5772/24775
  • Foord SH, Dippenaar-Schoeman AS, van der Merwe M (2002) A check list of the spider fauna of the Western Soutpansberg, South Africa (Arachnida: Araneae). Koedoe 45: 35–43. doi: 10.4102/koedoe.v45i2.25
  • Gabriel AGA, Chown SL, Barendse J, Marshall DJ, Mercer RD, Pugh PJA, Smith VR (2001) Biological invasions on Southern Ocean islands: the Collembola of Marion Island as a test of generalities. Ecography 24: 421–430. doi: 10.1111/j.1600-0587.2001.tb00477.x
  • Godfray HCJ (2002) Challenges for taxonomy. Nature 417: 17–19. doi: 10.1038/417017a
  • Goto H (1953) A species of Collembola, Sinella coeca (Schött) (Entomobryidae), new to South Africa. Entomologist’s Monthly Magazine 89: 165–166.
  • Greenslade P (1986) Additions to collembolan fauna of Heard Island. Records of the South Australian Museum 197: 91–96.
  • Greenslade P (1994) Collembola. In: Houston WWK (Ed.) Zoological catalogue of Australia. Volume 22. Protura, Collembola, Diplura. CSIRO, Melbourne, 19–138.
  • Greenslade P (2002) Assessing the risk of exotic Collembola invading subantarctic islands: prioritising quarantine management. Pedobiologia 46: 338–344. doi: 10.1078/0031-4056-00141
  • Greenslade P (2010) Collembola fauna of the South Shetland Islands revisited. Antarctic Science 22: 233–242.
  • Greenslade P, Ireson J, Skarzynski D (2014) Biology and key to the Australian species of Hypogastrura and Ceratophysella (Collembola: Hypogastruridae). Austral Entomology 53: 53–74. doi: 10.1111/aen.12048
  • Herbert D, Kilburn D (2004) Field guide to the land snails and slugs of eastern South Africa. Natal Museum, Pietermaritzburg.
  • Hlavac P (2007) Revision of the subtribe Clavigerodina and an annotated catalogue of South African Clavigeritae (Coleoptera: Staphylinidae: Pselaphinae). African Invertebrates 48: 65–92.
  • Hogg ID, Hebert PDN (2004) Biological identification of springtails (Hexapoda: Collembola) from the Canadian Arctic, using mitochondrial DNA barcodes. Canadian Journal of Zoology 82: 749–754. doi: 10.1139/z04-041
  • Hopkin S (1997) Biology of the Springtails. Insecta: Collembola. Oxford University Press, Oxford.
  • Hugo EA, Chown SL, McGeoch MA (2006) The microarthropods of sub-Antarctic Prince Edward Island: a quantitative assessment. Polar Biology 30: 109–119. doi: 10.1007/s00300-006-0166-x
  • Huntley B, Barnard P (2012) Potential impacts of climatic change on southern African birds of fynbos and grassland biodiversity hotspots. Diversity and Distributions 18: 769–781. doi: 10.1111/j.1472-4642.2012.00890.x
  • Ireson JE, Greenslade P (1990) Lasofinius gen. n. (Collembola: Tomoceridae) from Tasmania and a re-examination of Neophorella dubia Womersley (Tomoceridae). Journal of the Australian Entomological Society 29: 205–214. doi: 10.1111/j.1440-6055.1990.tb00350.x
  • Janion C, Bedos A, Bengtsson J, Deharveng L, Jansen van Vuuren B, Leinaas HP, Liu A, Malmström A, Porco D, Chown SL (2011a) Springtail diversity in South Africa. South African Journal of Science 107: Art. #582. doi: 10.4102/sajs.v107i11/12.582
  • Janion C, Bedos A, Deharveng L (2011b) The genus Ectonura Cassagnau, 1980 in South Africa (Collembola: Neanuridae: Neanurinae), with a key to South African Neanurinae. ZooKeys 136: 31–45. doi: 10.3897/zookeys.136.1744
  • Janion C, D’Haese C, Deharveng L (2012) A new species and first record of the genus Triacanthella Schaffer, 1897 (Collembola, Hypogastruridae) for Africa. ZooKeys 163: 57–68. doi: 10.3897/zookeys.163.2298
  • Joppa LN, Roberts DL, Pimm SL (2011) The population ecology and social behaviour of taxonomists. Trends in Ecology and Evolution 26: 551–553. doi: 10.1016/j.tree.2011.07.010
  • Jordana R (2012) Synopses on Palaearctic Collembola. Capbryinae and Entomobryini. Soil Organisms 84: 1–390.
  • Lawrence RF (1953) The biology of the cryptic fauna of forest. AA Balkema, Cape Town.
  • Liu A, Janion C, Chown SL (2012) Collembola diversity in the critically endangered Cape Flats Sand Fynbos and adjacent pine plantations. Pedobiologia 55: 203–209. doi: 10.1016/j.pedobi.2012.03.002
  • MacDougall AS, Turkington R (2005) Are invasive species the drivers or passengers of change in degraded ecosystems? Ecology 86: 42–55. doi: 10.1890/04-0669
  • Magurran AE (2004) Measuring Biological Diversity. Blackwell Publishing, Oxford.
  • Massoud Z (1967) Monographie des Neanuridae, Collemboles Poduromorphes à pièces buccales modifiées. In: Biologie de l’Amérique Australe, vol.III. C.N.R.S. (Ed.), Paris, 7–399.
  • Mitra SK (2002) Status of Dicranocentruga Wray, 1953 with the description of a new species (Collembola: Entomobryidae). Records of the Zoological Survey of India 100: 105–116.
  • Paclt J (1959) Collembola. South African Animal Life. Results of the Lund University Expedition in 1950–1951, Vol. VI. Almqvist and Wiksells Boktryckeri AB, Uppsala, 24–78.
  • Paclt J (1964) Ein neuer Prorastriopes aus Zululand und zur Nomenklatur due südafrikianischen Rastropes-Art. Senckenbergiana Biologica 45: 661–663.
  • Paclt J (1965) Cyphoderus trinervoidis n.sp., ein neuer Termitophile aus Transvaal (Ins., Collembola). Senckenbergiana Biologica 46: 59–60.
  • Paclt J (1967) On South and Central African Collembola. Journal of the Entomological Society of South Africa 29: 135–47.
  • Parr CL, Robertson HG, Chown SL (2003) Apomyrminae and Aenictogitoninae: two new subfamilies of ant (Hymenoptera: Formicidae) for southern Africa. African Entomology 11: 128–129.
  • Petersen H, Luxton M (1982) A comparative analysis of soil fauna populations and their role in decomposition processes. Oikos 39: 287–388. doi: 10.2307/3544689
  • Pimm SL, Jenkins CN, Joppa LN, Roberts DL, Russell GJ (2010) How many endangered species remain to be discovered in Brazil? Natureza and Conservação 8: 71–77. doi: 10.4322/natcon.00801011
  • Porco D, Bedos A, Greenslade P, Janion C, Skarżyński D, Stevens M, Jansen van Vuuren B, Deharveng L (2012) Challenging species delimitation in Collembola: cryptic diversity among common springtails unveiled by DNA barcoding. Invertebrate Systematics 26: 470–477. doi: 10.1071/IS12026
  • Potapov M (2001) Synopses on Palaearctic Collembola, Volume 3. Isotomidae. Abhandlungen und Berichte des Naturkundemuseums Görlitz, Band 73, Heft 2, 2001, 603.
  • Potapov M, Gao Y, Deharveng L (2013) Taxonomy of the Cryptopygus complex. I. Pauropygus – a new worldwide littoral genus (Collembola, Isotomidae). ZooKeys 304: 1–16. doi: 10.3897/zookeys.304.4083
  • Potapov M, Janion C, Deharveng L (2011) Two new species of Parisotoma (Collembola: Isotomidae) from the Western Cape, South Africa. Zootaxa 2771: 17–24. PMid:21547000, PMCid:3086760
  • Pryke JS, Samways MJ (2010) Significant variables for the conservation of mountain invertebrates. Journal of Insect Conservation 13: 627–641. doi: 10.1007/s10841-009-9213-6
  • Pyšek P, Richardson DM, Rejmánek M, Webster GL, Williamson M, Kirschner J (2004) Alien plants in checklists and floras: towards better communication between taxonomists and ecologists. Taxon 53: 131–143. doi: 10.2307/4135498
  • Rebelo AG, Boucher C, Helme N, Mucina L, Rutherford MC (2006) Fynbos Biome. In: Mucina L, Rutherford MC (Eds) The vegetation of South Africa, Lesotho and Swaziland. Strelitzia 19, South African National Biodiversity Institute, Pretoria, 52–219.
  • Richardson DM, Pyšek P (2006) Plant invasions: merging the concepts of species invasiveness and community invasibility. Progress in Physical Geography 30: 409–431. doi: 10.1191/0309133306pp490pr
  • Robertson HG (2000) Afrotropical ants (Hymenoptera: Formicidae): taxonomic progress and estimation of species richness. Journal of Hymenoptera Research 9: 71–84.
  • Robertson HG (2002) Revision of the ant genus Streblognathus (Hymenoptera: Formicidae: Ponerinae). Zootaxa 97: 1–16.
  • Roques A, Rabitsch W, Rasplus J-Y, Lopez-Vaamonde C, Nentwig W, Kenis M (2009) Alien terrestrial invertebrates in Europe. In: Handbook of alien species in Europe. Springer, Heidelberg, 63–79. doi: 10.1007/978-1-4020-8280-1_5
  • Rougerie R, Decaens T, Deharveng L, Porco D, James SW, Chang C-H, Richard B, Potapov M, Suhardjono Y, Hebert PDN (2009) DNA barcodes for soil animal taxonomy. Pesquisa Agropecuária Brasileira 44: 789–801. doi: 10.1590/S0100-204X2009000800002
  • Rouget M, Richardson DM, Cowling RM, Lloyd JW, Lombard AT (2003) Current patterns of habitat transformation and future threats to biodiversity in terrestrial ecosystems of the Cape Floristic Region, South Africa. Biological Conservation 112: 63–85. doi: 10.1016/S0006-3207(02)00395-6
  • Rousse P, van Noort S (2013) Revision of the Afrotropical Lycorininae (Ichneumonidae, Hymenoptera) with description of a new species from South Africa. Zootaxa 3666: 252–266. doi: 10.11646/zootaxa.3666.2.8
  • Rusek J (1971) Zur Taxonomie der Tullbergia (Mesaphorura) krausbaueri (Börner) und ihrer verwandten (Collembola). Acta Entomologica Bohemoslovaca 68: 188–206.
  • Salmon JT (1964) An Index to the Collembola, Volume 1. Royal Society of New Zealand. Bulletin No. 7, Wellington, 1–144.
  • Samper C (2004) Taxonomy and environmental policy. Philosophical Transactions of the Royal Society B 359: 721–728. doi: 10.1098/rstb.2004.1476
  • Schoeman CS, Foord SH (2012) A checklist of ants (Hymenoptera: Formicidae) from the Marakele National Park, Limpopo, South Africa. Koedoe 54: Art. # 1030. doi: 10.4102/koedoe.v54i1.1030
  • Scholtz CH, Chown SL (1995) Insects in southern Africa: How many species are there? South African Journal of Science 91: 124–126.
  • Silvestri F (1918) Contribuzione alla conoscenza dei Termitidi e Termitofili dell’ Africa occidentale. II. Termitofili. Parte prima. Bollettino del Laboratorio di Zoologia General e Agraria, Portici 12: 287–346.
  • Skaife SH (1954) African Insect Life. Longmans Green, Cape Town.
  • Stach J (1947) The Apterygotan fauna of Poland in relation to the world-fauna of this group of insects. Family Isotomidae, 488 pp.
  • Terauds A, Chown SL, Bergstrom DM (2011) Spatial scale and species identity influence the indigenous-alien diversity relationship in springtails. Ecology 92: 1436–1447. doi: 10.1890/10-2216.1
  • Thibaud JM (2008) Les collemboles des sables littoraux de Madagascar. Annales de la Société Entomologique de France 44: 503–519. doi: 10.1080/00379271.2008.10697586
  • Thibaud JM (2010) Les collemboles des sables littoraux de l’île de Mayotte. Essai de synthèse sur les collemboles des sables littoraux d’îles de l’Océan Indien (zone ouest). Revue Française d'Entomologie 32: 113–121.
  • Thibaud JM, Massoud Z (1988) Recherche sur la faune interstitielle aérienne des sables fins: les Collemboles. II - Désert de Namibie. Annales de la Société Entomologique de France 24: 211–214.
  • van Noort S (2004–2014) Wasps, bees and ants of Africa and Madagascar. Iziko Museums of South Africa. www.waspweb.org [accessed 02 April 2014]
  • van Straalen NM (1998) Evaluation of bioindicator systems derived from soil arthropod communities. Applied Soil Ecology 9: 429–437. doi: 10.1016/S0929-1393(98)00101-2
  • Wallace MMH (1964) Present and probable world distribution of Sminthurus viridis and prospects for its biological control. Pedobiologia 14: 238–243.
  • Wallace MMH, Walters MC (1974) The introduction of Bdellodes lapidaria (Acari: Bdellidae) from Australia into South Africa for the biological control of Sminthurus viridis (Collembola). Australian Journal of Zoology 22: 505–517. doi: 10.1071/ZO9740505
  • Walters MC (1968) A study of Sminthurus viridis (L.) (Collembola) in the Western Cape Province. Entomology Memoirs 16: Department of Agriculture Technical Services. University of Stellenbosch, Stellenbosch, 99 pp.
  • Wardle DA, Bardgett RD, Klironomos JN, Setälä H, van der Putten WH, Wall DH (2004) Ecological linkages between aboveground and belowground biota. Science 304: 1629–1633. http: 10.1126/science.1094875
  • Weiner WM, Najt J (1998) Collembola (Entognatha) from East Africa. European Journal of Entomology 95: 217–237.
  • Weiner WM, Najt J (1999) New genus of Tullbergiinae (Collembola). Annales de la Société Entomologique de France 35: 183–187.
  • Womersley H (1931) Some Collembola of the family Sminthuridae from South Africa. Annals of the South African Museum 30: 137–156.
  • Womersley H (1934) On some Collembola-Arthropleona from South Africa and Southern Rhodesia. Annals of the South African Museum 30: 441–475.
  • Yosii R (1959) Collembolan fauna of the Cape Province, with special reference to the genus Seira Lubbock. Special Publication from the Seto Marine Biological Laboratory. Biological results from the Japanese Antarctic Research Expeditions 6: 3–24.
  • Yosii R (1977) Critical checklist of the Japanese species of Collembola. Contributions of the Biological Laboratory of Kyoto University 25: 141–170.
  • Zhang F, Deharveng L, Chen JX (2009) New species and rediagnosis of Coecobrya (Collembola: Entomobryidae), with a key to the species of the genus. Journal of Natural History 43: 2597–2615.