Despite their significance in soil ecosystems and their use for investigations of soil ecosystem functioning and in bioindication elsewhere, springtails (Collembola) have not been well investigated in
South Africa. Early recognition of their role in soil systems and sporadic systematic work has essentially characterised knowledge of the southern African fauna for some time. The situation is now
changing as a consequence of systematic and ecological work on springtails. To date this research has focused mostly on the Cape Floristic Region and has revealed a much more diverse springtail fauna
than previously known (136 identifiable species and an estimated 300 species for the Cape Floristic Region in total), including radiations in genera such as the isotomid Cryptopygus. Quantitative
ecological work has shown that alpha diversity can be estimated readily and that the group may be useful for demonstrating land use impacts on soil biodiversity. Moreover, this ecological work has
revealed that some disturbed sites, such as those dominated by Galenia africana, may be dominated by invasive springtail species. Investigation of the soil fauna involved in decomposition in
Renosterveld and Fynbos has also revealed that biological decomposition has likely been underestimated in these vegetation types, and that the role of fire as the presumed predominant source of nutrient
return to the soil may have to be re-examined. Ongoing research on the springtails will provide the information necessary for understanding and conserving soils: one of southern Africa’s major
The significance of soil organisms for ecosystem functioning and ecosystem service delivery is widely appreciated.1,2 Amongst the many arthropod taxa that contribute to soil ecosystem functioning,
the springtails (Collembola) have been identified as an important group. These small arthropods occur in most ecosystems and may reach densities of several hundred thousand individuals per square
metre.3 They form the prey of a wide variety of soil organisms and ground-living arthropods, actively contribute to soil formation and structure,4 and have major effects on
both plants and plant consumers; and thus link the above-ground and below-ground components of terrestrial systems.2,5 The ecological roles and abundance of springtails in terrestrial
systems have also led to their recognition as important organisms for bioindication,6,7 and as model organisms in ecotoxicology.8 Consequently, springtails have been the subject
of substantial and long-standing interest, especially in Europe and North America.3,9,10
The significance of springtails in soil systems in South Africa is unlikely to be different from that elsewhere, recognising that springtails reach their highest species richness and abundance in moist
habitats rather than in arid areas.11,12 However, in contrast with many other regions of the world,10 and with other components of the soil fauna in South Africa, such as the
ants,13,14,15,16 springtails have, until recently, been the subject of little attention. Early research recognised that springtails are likely to play an important role in various South
African biomes,17 and since the initial work of Womersley18, ongoing, but sporadic and typically restricted taxonomic investigations have been
Few investigations have sought to understand comprehensively the diversity of the group (at present the only available, but unpublished list by P. Greenslade, places the fauna at about 43
genera and 90 species for the Western Cape), their contribution to ecosystem functioning, and their utility for bioindication,29 although occasional attempts at doing so have been
made.30 The exception is research on the group undertaken in the sub-Antarctic Tundra and Polar Desert biomes of the Prince Edward Islands, which, at least geopolitically, form part
of South Africa.31 For these island biomes, the limited fauna of 16 species is relatively well understood.32,33,34,35,36 Perhaps more significantly, it is this island work
that has precipitated a recent and substantial change in current understanding of springtail ecology and systematics in continental South Africa.
On the basis of a comprehensive comparison of the likely effects of climate change on indigenous versus invasive springtails on the Prince Edward Islands, funded by a bilateral grant
under the South Africa–Norway agreement (Table 1),34 a second project was developed to compare life histories of species in polar systems (namely, the Prince Edward Islands
and Svalbard, Norway) and more temperate systems (southern Norway and the Western Cape Province, South Africa). This work sought also to investigate the implications thereof for ecosystem
functioning and drew in further expertise from Sweden and support from a South Africa–Sweden bilateral (Table 1). In both cases, it was recognised from the outset that the springtails
in South Africa are poorly known systematically, and that this taxonomic impediment37 would therefore constrain work substantially. Thus, additional collaborations were established
with systematic experts under a third bilateral agreement (South Africa–France) and with the support of the International Barcode of Life project (iBOL - http://ibol.org) (Table 1).
Here we report on research so far undertaken across these major projects, how systematic and ecological understanding of the group is progressing for South Africa, and the prospects for ongoing
research and future collaborations amongst researchers with an interest in soil diversity and ecosystem functioning.
TABLE 1: A summary of the springtail research undertaken over the past decade in South Africa, funded mostly through bilateral research agreements. In each case the
partner country, project title, areas of work and outputs (e.g. numbers of students advised, investigators involved and papers published) are indicated. The International
Barcode of Life (iBOL)-related work is not listed as a separate project because it was initiated within the South Africa–France bilateral agreement. This work is now being
pursued both through another project within the same bilateral agreement and through support from a larger, barcoding initiative established with funding from iBOL
and managed by the South African Institute for Aquatic Biodiversity.
Owing to estimates of the likely large size of the South African springtail fauna, the collaborative research was established to investigate the diversity of the Cape Floristic Region, focusing mostly on
fynbos, renosterveld and forest vegetation. From the outset, the research approach emphasised collection across a range of sitesin a systematic fashion (Figure 1) for estimates of richness and abundance,
supplemented by ad hoc collections from as wide a range of habitats as possible for the systematic and barcoding studies. Importantly, our investigations combined traditional taxonomy and barcoding
to ensure an integrated, modern approach to the systematics of the group.
Initial assessments of the published taxonomic research indicated a faunal assortment of 90 species in 16 families, but, excluding incorrect identifications, only 57 valid and recognisable species
(Table 2a). Now, based on a geographically extensive collection of more than 450 samples from the Western Cape (Figure 1), it is clear that the faunal component is much larger. The collection methods
included high gradient extraction of litter bags; Berlese–Tullgren extraction of leaf litter, moss and rotten wood; vegetation beating and collection by hand (especially in caves and streams);
pitfall trapping; litter sifting; and soil washing (of beach and deep soil sand). The current faunal assemblage stands at 136 morphospecies in 19 families (Table 2b), but only half of the samples
have been sorted to species sofar and not all microhabitats have been surveyed to completion. Moreover, assessments of previous lists (in particular Paclt19) for the country, the
examination of museum specimens and barcoding work have indicated that many species have been misidentified and that several groups have either cryptic species or have shown substantial radiations.
For example, more than 11 Cryptopygus and 7 Parisotoma species were found during our survey, in contrast to only 2 species that have been previously recorded – Cryptopygus
caecus and Parisotoma notabilis.19 In addition, 23 described species of Seira are found in South Africa,22 whilst only 1 species is described from
Australia.38 These groups that have radiated in the Cape Floristic Region are now the subject of closer investigation from the classical taxonomic39,40 and barcoding
perspectives. Thus, although research to date has more than doubled the number of known species, based on our experience from elsewhere it seems likely that at least 300 species will be
characteristic of the regional (i.e. Cape Floristic Region) fauna. For South Africa as a whole the richness is certainly much higher. Even so, the Collembola likely has fewer species than
are found in the highly diverse insect orders, such as the Coleoptera or Lepidoptera.41 Many sequences obtained from barcoding have been connected to traditional taxonomic species,
although certain species with poor morphological descriptions have been more difficult(as in the case of several Seira species). The barcoding results have also led to careful re-examination
of the individuals traditionally assigned to the widespread species Parisotoma notabilis, which actually represent a group of several similar species of endemic Parisotoma. In addition,
the barcoding has proven not only to be of particular importance to detect introduced species of complex groups (e.g. Isotomurus maculatus), but also to show that some forms expected to be
European actually do not match any European cluster (such as one cluster of Isotomurus and several of Ceratophysella).
FIGURE 1: Systematic (circle with dot in centre) and ad hoc (black circle) sampling sites for the current springtail diversity assessments in the Western Cape Province, South Africa. Protected areas are overlaid in shading.
TABLE 2a: Number of springtail species in the Western Cape Province, South Africa in each of the springtail families as listed in the systematic literature as of November
TABLE 2b: Number of springtail species in the Western Cape Province, South Africa added from our own collections (as of November 2010).
Amongst the 136 species-level taxa identified for the Western Cape, at least 34 represent species either with wide distributions or known or suspected to have been introduced from elsewhere. Based on
our preliminary molecular analysis, two of these species – Neanura muscorum and Isotomurus cf. maculatus – have already been shown to be invasive, occurring locally in fynbos
or forest habitat. Similar comparisons between South African and European populations of other suspected invasive species are in progress. For example, in an investigation of diversity and decomposition
rates in renosterveld fragments in the Piketberg area,42 the springtail assemblages were dominated (60% by abundance) by a single species, Hypogastrura manubrialis, which is widely
distributed in Europe,43 and clearly is an invasive species in the Western Cape. Similarly, two Isotomurus species, thought to be introduced to the region, were also found in high
abundance in a preliminary assessment of the diversity of springtails in Cape Flats Sand Fynbos and an adjacent pine plantation in the Tokai Forest Reserve, which forms part of Table Mountain National
Park; these species were restricted to the pine plantation. This study also showed that springtail abundance and species richness (but not species identity) differ significantly amongst these two major
habitat types, and, that for complete estimation of the fauna using a litter sampling technique (for each sample: 1 L oflitter collected over a standardised 1 m2 and then extracted by a
Berlese–Tullgren funnel in the laboratory), the extent of sampling is not so onerous that it precludes reasonably straightforward estimation of local (alpha) diversity (Figure 2). In consequence,
the springtails could be used for assessments of soil health, as they are elsewhere.3,9
FIGURE 2: Sample-based species rarefaction curves for the (a) Cape Flats Sand
Fynbos and (b) Pinus radiata sites.
In the context of soil health, the influence of springtails on litter decomposition rates and nutrient cycling, including the ways in which litter type, spatial position and home-field advantage
influence decomposition, has long been of interest to soil ecologists.45,46,47,48,49 As a consequence of the research priorities identified within our South Africa–Sweden and South
Africa–Norway bilateral projects (Table 1), we have also been concerned with these questions. The dominant paradigm for the Fynbos biome has been that loss of organic matter and nutrientcycling
take place largely as a consequence of fires, which, with an average fire frequency of 11 years,50 return nutrients accumulated in litter to the soil. Biological decomposition was largely
relegated to a less significant role.51,52,53
During the investigation of springtail diversity in renosterveld fragments, decomposition of the litter of three representative plant species was also investigated. Decomposition of these species
varied between 0.00674/day for Galenia africana, a shrub favoured by disturbances such as overgrazing, to 0.00222/day for renosterbos Elytropappus rhinocerotis and 0.00029/day for the
sturdy geophyte Watsonia borbonica, corresponding to litter half-lives of 0.3, 0.8 and 6 years, respectively.42 The rates for Galenia and renosterbos are much faster than
those previously found for litter of the fynbos species Leucadendron parile and Protea repens.51,52 However, they are not unusual compared with other fynbos species we have
studied (Figure 3; personal observation).
If the mean fire return time for fynbos systems is about 11 years,50 and if decomposition of the less hardy species proceeds such that many litters have half-lives that are less than half
this time, then biological decomposition as a nutrient recycling process is much more significant than previously estimated.53 Ultimately, the significance of this decomposition will depend
on the relative contributions of species with more readily or less readily decomposed litter to the litter pool of any given system. To date, such estimates have not been made. Species with high
decomposition rates of the kinds we investigated clearly make a significant contribution to the fynbos flora, and it is obvious that litter of many species do not accumulate on the ground in the
way that some Protea species do (personal observations). Thus, the current research on ecosystem functioning has demonstrated that important as fire is in the Fynbos biome, it may be
complemented and sometimes surpassed by other processes.
FIGURE 3: Decomposition rates per day (mean ± s.d.) of various renosterveld
and fynbos species: Galenia africana, renosterbos (Elytropappus rhinocerotis)
and Watsonia borbonica42 (black triangle); Protea exima53 (black circle); and Leucospermum parile
and Protea repens51,52 (outlined circle).
Our research on springtails has thus far revealed not only a hitherto undocumented diversity in South Africa, but also that the group may be much more significant for ecosystem functioning than
previously thought. Given a growing human population and its impacts on the environment,54 national requirements for sustainable development and conservation, and the need to provide
measures of conservation and sustainable development success internationally, ongoing work on the Collembola will prove both useful and valuable. The current collaborations described here will
continue throughout the duration of the International Barcode of Life initiative (iBOL - http://ibol.org), with support from several institutions, both in South Africa
and abroad, and growing interest from various sectors in demonstrating soil health and conservation success. Moreover, as this knowledge develops so more information will be made available
through the projecthome page (www.sun.ac.za/cib/collembola). Our long-term aims are to encourage additional work on the group in the southern
African region, and to foster collaborations that can enhance understanding of this significant group of organisms. Underlying these aims is the realisation that sustainable development and
conservation must continue to focus on one of southern Africa’s major natural assets: the soil.
We are grateful for support, through the National Research Foundation, from the South Africa–France, South Africa–Norway and South Africa–Sweden bilateralgrants, and the International
Barcode of Life project. We thank Melodie McGeoch, Brigitte Brashler and two anonymous reviewers for comments on a previous version of the manuscript. The map was kindly produced by Dian Spear. Cape
Nature and SANParks provided collection permits.
We declare that we have no financial or personal relationships which may have inappropriately influenced us in writing this article.
S.L.C. was the project leader; A.B., L.D., H.P.L. and C.J. provided taxonomic data; H.P.L., J.B., A.M. and C.J. provided decomposition data; A.L. provided rarefaction curves; and D.P. and B.J.V.V.
provided data on barcoding.
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