Conservation implications: Invertebrates constitute a substantial and functionally significant component of terrestrial biodiversity and are valuable indicators of environmental condition. Although consideration of invertebrates has historically been neglected in conservation planning and management, substantial progress with surveys, systematics and bioindication means that it is now both feasible and advisable to incorporate them into protected area monitoring activities.

Monitoring has a long history in entomology with long-term data series being used, for example, to develop forecasting systems in pest management (Conrad, Fox & Woiwod 2007). The Rothamsted Insect Survey (UK) has, for example, been monitoring the relative abundance of insects since the 1960s (Woiwod & Harrington 1994). Indeed, butterfly monitoring in the UK is amongst the most comprehensive current monitoring programmes using insects. Here, four schemes provide complementary information on the distribution, abundance and diversity of butterfly assemblages. These schemes include mapping and multiscale atlases to monitor changes in species distributions, Red Data Books on species conservation status, transects that provide information on population trends and a system of occasional surveys of selected species (Thomas 2005). Together these data are extremely valuable for butterfly conservation and they provide substantial insight into annual and long-term trends in the diversity, abundance and distribution of the species concerned (Conrad et al. 2007).

Significant advances have also been made in South Africa, and elsewhere, over the last three decades (McGeoch 2002), both with regard to improvements in invertebrate systematics (Speight & Castella 2001; Telford & Littlewood 2008) and development of invertebrate bioindicators and their use in monitoring (McGeoch 2007; Samways, McGeoch & New 2010). In this article, we use a selection of taxa and case studies (which is by no means comprehensive) to demonstrate the value and feasibility of both monitoring invertebrates in PAs and including them in broader conservation assessments and biodiversity monitoring programmes. We discuss the objectives, opportunities and constraints specific to invertebrate monitoring in PAs, with a particular focus on South Africa. We identify suitable taxa for monitoring based on a range of monitoring objectives and experiences, and make recommendations for the implementation of an invertebrate monitoring programme for PAs in South Africa.

In November 1995, South Africa ratified the Convention on Biological Diversity (CBD) under which signatories are obligated to develop a strategic plan for the conservation and sustainable use of biodiversity. Meeting the goals of the Convention requires intensive national effort including (1) discovery, description and inventory of species, (2) analysis and synthesis of information into predictive classification systems, and (3) organisation of this information into an efficiently retrievable form that best meets the needs of science, conservation and society. Invertebrates necessarily form a significant component of this task as they not only perform critical roles in ecosystems (via pollination, decomposition, soil property maintenance, predation, parasitism, and herbivory; see for example, Lawrence (1953), Scholtz and Holm (1985) and Scholes and Walker (1993)) but also contribute disproportionately to biodiversity and outnumber plant and vertebrate richness several fold (Grimaldi & Engel 2005; Samways 2005).

Although lesser known than in the case of vertebrates and plants, several South African invertebrate taxa also display high levels of endemism. Many invertebrates have narrow ranges; for example, in KwaZulu-Natal alone seven species of butterfly, about 80 species of snail, and more than 200 species of millipede are found exclusively in the province (Pringle, Henning & Ball 1994; Hamer 1998; Herbert & Kilburn 2004). These species require conservation action to ensure their survival because of their small distribution ranges and increasing rates of land transformation (Thompson 2008). Another example is the very high concentration of taxa with a southern Gondwanan ancestry that are restricted to the Cape Fold mountains. This very rich South African Gondwanan fauna (800 species, 200 families) has its primary hotspot on the Cape Peninsula (a substantial proportion of which constitutes Table Mountain National Park [TMNP] in the Western Cape [Day 2005]). High levels of endemism (111 Cape Peninsula endemics, according to Picker and Samways [1996]) are concentrated in refugial habitats such as mountain streams, southern Afrotemperate forest and caves. The Cape Floristic Region, in general, has a distinctive invertebrate fauna, with high levels of endemism in gall-forming insects (Wright & Samways 1998), leafhoppers (Stiller 2002), dung beetles (Davis 2002), scorpions (Prendini 2005), millipedes (Hamer & Slotow 2002), velvet worms, (Onychophora) (Daniels et al. 2009; Hamer, Samways & Ruhberg 1997), heelwalkers (Mantophasmatodea) (Damgaard et al. 2003) and lycaenid butterflies (Cottrell 1985). Other biomes are probably less well studied in terms of their invertebrate fauna and levels of endemicity, but the more northern Afrotemperate forests, and even grassland and savanna have high levels of narrow endemism for some taxa that have been investigated in this respect (e.g. Hamer, Slotow & Lovell 2006; Hamer & Slotow 2009).

PAs play a vital role in the conservation of such narrow range endemics, and these species are generally considered ‘species of special concern’ in PA management objectives (Table 1). For example, in the TMNP several invertebrates require special attention, such as Colophon westwoodi (Coleoptera: Lucanidae) and Trimenia malagrida malagrida (Lepidoptera: Lycaenidae). Peripatopsis alba is an endangered cave-dwelling velvet worm confined to Table Mountain (Hamer et al. 1997). These taxa have been found only at very few localities and these sites thus require special conservation management.

As a charismatic group of taxa, butterflies have long received the attention of entomologists, collectors and conservationists across the world. Similar approaches and activities for other invertebrate groups are nonetheless possible. For example, South Africa has a rich arachnid fauna with about 5000 known species, representing 6% of global arachnid diversity. Although they constitute a diverse and ecologically important group of invertebrates in South Africa, they have traditionally been comparatively poorly sampled in some areas. However, in 1997 the South African National Survey of Arachnida (SANSA) was initiated by the Biosystematics Unit (ARC-Plant Protection Research Institute). SANSA aims to inventory the arachnofauna of the country and forms part of the Threatened Species Programme of the South African National Biodiversity Institute (SANBI) to determine the distribution and conservation status of spiders and scorpions.

One of the focus areas of SANSA is to survey PAs to obtain species-specific information, compile inventories and to determine which species in South Africa receive some protection. More than 86 surveys in PAs are currently underway, of which 25 have already resulted in published annotated checklists that provide information on abundance, behaviour and the distribution of arachnid species. Several forms of participation have involved PAs, including SANSA surveys, surveys by PA managers and rangers, student research projects (seven MSc projects completed), by-catch data from other research projects and records submitted by the public. This has dramatically increased the knowledge of the spider fauna of PAs and with material made available to taxonomists has resulted in the description of several new species (e.g. Wesolowska & Haddad 2009). During this survey a record 430 species were recorded, which represent more than 20% of the South African spiders and include several new and undescribed species (Haddad, Dippenaar-Schoeman & Wesolowska 2006; Wesolowska & Haddad 2009). Long-term surveys in PAs also increase the number of species sampled. For example, during the survey of the Kruger National Park (KNP) 152 species were reported in 2003, increasing the number currently listed for the park to 305 (Dippenaar-Schoeman & Leroy 2003, see also Dippenaar-Schoeman 1988, 2006). PAs have proven to be particularly valuable sites to SANSA, both from the perspective of encountering pristine habitat and high diversity, as well as for the safety of survey teams (Dippenaar-Schoeman et al. 1999, 2005).

Because of the importance of inventories in species conservation assessment and PA management, Engelbrecht (2010) investigated to what extent and how invertebrate species inventories are currently applied in PA management in South Africa. PA managers and ecologists were asked, using a questionnaire-based survey, to comment on whether (1) a documented management plan existed for the PA in question, (2) any invertebrate species inventories existed for the PA, and if so, (3) whether those inventories had been used to guide the development of any of the management goals or actions for the PA. Although the overwhelming majority considered inventories desirable, very few of the responses received indicated that invertebrates were specifically considered in management goals or actions. Those that did included the Brenton Blue Butterfly Reserve (Western Cape) and the Suikerbosrand Nature Reserve (Gauteng), which is home to the Heidelberg Copper (Engelbrecht 2010). In other cases, the principal value of inventories was to tourists and scientists visiting the PA. The outcome of this study illustrates that although insect conservation and monitoring have not yet become mainstream in PA planning and management processes, the potential value of inventories is (1) widely acknowledged and (2) would be more suited to this purpose where accompanied by information on richness patterns, distributions, and rare and indicator species (Engelbrecht 2010).

As shown in Table 2, despite this and other available and related tools in KwaZulu-Natal, invertebrates have been explicitly included in only a small proportion of land use change applications under the National Environmental Management Act (NEMA) (Act 107 of 1998), where the proposed land use change may have significant impacts on species of special concern. Various mitigation measures were proposed by Ezemvelo KZN Wildlife as befitting the range of invertebrate taxa concerned. The recommendations ranged from no mitigation specific to invertebrates, through restriction of the proposed land use to certain portions of the land (e.g. to already transformed portions of the land), to set-asides for invertebrate conservation. In one case, refusal of the permit was recommended on invertebrate conservation grounds. Some of these recommendations were incorporated into the Record of Decision issued by the provincial Department of Agriculture and Environmental Affairs (now the Department of Agriculture, Environmental Affairs and Rural Development), whilst others were not.

One of the insights gained from the aforementioned process is the need to strengthen the profile of invertebrates in conservation planning, both within and outside of PAs, so that charismatic umbrella vertebrate taxa are not relied upon to effect the conservation of invertebrates and their habitats. The Ezemvelo KZN Wildlife IEM team now communicates to all relevant parties when umbrella species are used for invertebrate conservation purposes. The need to conserve invertebrates is not readily accepted or appreciated by some permitting authorities, applicants and environmental consultants, and some lack expertise on invertebrates and therefore ignore them. Profiling of invertebrates and training of consultants are required to successfully integrate invertebrate conservation into processes legislated by NEMA and its subsidiary Acts. This will allow invertebrates to be considered explicitly and conserved in areas subject to change in land use.

Monitoring invertebrate responses to fire (e.g. frequency and intensity) is also important in PAs within biomes where fire is a key habitat management tool, such as savanna and fynbos (Parr et al. 2002). Fire is a particular conservation risk in Mediterranean-type biomes and, although it is an essential element in the maintenance of biodiversity, fire regimens (frequency, intensity and timing) have been significantly altered by increasing human population densities, resulting in too frequent, too infrequent and other ecologically undesirable fire properties (Forsyth & Van Wilgen 2008). A study by Pryke (2008) showed that the invertebrates in the TMNP demonstrated differential short-term resilience to fire, especially with regard to species richness and abundance. Some components of the invertebrate assemblage were remarkably resilient to fire (particularly ants), whilst others were far more conservative (pollinators and detritivores). Invertebrates were also found to recolonise high-altitude grasslands in the Drakensberg within a short space of time after burning (Uys, Hamer & Slotow 2006). However, although not well studied, fire in Afrotemperate forest may have a negative impact on some less mobile taxa such as molluscs (Uys, Hamer & Slotow 2010). In terms of fire-related monitoring objectives in PAs, invertebrates are thus an important element of monitoring the status of and trends in habitat persistence, degradation and rehabilitation (Table 1).

Monitoring or research programmes that include accurate taxonomic identifications are the most likely way of detecting new alien species and structured monitoring programmes for target species allow for early detection and potentially rapid control. Those involved in invertebrate monitoring should not only be familiar with known invasives, but be on the lookout (early detection) for species that are problematic in other countries, but have not yet been recorded in South Africa. Examples include the yellow crazy ant (Anoplolepis gracilipes) and red imported fire ant (Solenopsis invicta) (Lowe et al. 2000).

Decisions on what to monitor must also necessarily consider the focal taxa and habitats along with logistic feasibility of the proposed exercise. Different invertebrate taxa are obviously more, or less, appropriate for monitoring in particular environments and are differentially sensitive to particular stressors. Table 3 provides an assessment of the value of selected higher-level taxa to monitoring based on a range of key generic criteria. The outcome of this assessment provides guidance for selecting which higher taxa to focus on a priori, before proceeding with more specific indicator testing as would be required to achieve more specific monitoring objectives (see McGeoch [1998] and McGeoch, Van Rensburg & Botes [2002] for general approach). We selected 16 objectives and criteria that are most relevant in a PA context and assessed a range of higher taxa against these. Rather than assessing all higher taxa, a subgroup of seven, which were considered to be those broadly most feasible in a South African context (based on the collected specialist opinion and experience of the authors), was selected a priori. Other taxa may well be relevant and suitable for more narrowly defined purposes. The taxa were scored for each criterion (see explanation in Table 3) and ranked according to a descending score (left to right, Table 3). Odonata, Formicidae, Lepidoptera (butterflies) and Scarabaeidae were considered best suited for monitoring in PAs. Interestingly, Coleoptera, Hymenoptera and Lepidoptera (along with molluscs and arachnids) currently receive most research attention in South African National Parks (Figure 1). For example, dung beetles, whilst potentially useful indicators as a result of their specialised distribution patterns, require fine-scale spatial data to realise this potential (Davis 2002). Given the rapid increase in knowledge and expertise on South Africa’s arachnid fauna, particularly in PAs, the score for this taxon is likely to increase in the foreseeable future. We consequently use the Odonata and Formicidae (Hymenoptera) as two of the highest-scoring taxa in our assessment to illustrate aspects of the suitability and application of invertebrates for monitoring in PAs.

FIGURE 1: Percentage of research projects (n = 63) on invertebrates (a) per taxon and (b) per park (including nontaxon specific projects) currently registered with South African National Parks (as at November 2009).

In addition to the well-known and widely used SASS system (Dallas & Day 1993; Revenga et al. 2005), the Dragonfly Biotic Index (DBI) provides a measure of ecological integrity for freshwater systems. The DBI is a weighted index (see Table 4) based on the quantitative assessment of three subindices of species distribution, threat status and sensitivity to disturbance (Simaika & Samways 2008, 2009a). The total DBI of a water body (stream, river or pool) reflects the total odonate assemblage, thus allowing for water bodies to be compared and restoration success to be monitored (Simaika & Samways 2008). Every South African odonate species has been assigned a score (Samways 2008). The DBI has been tested and applied in biomonitoring (Simaika & Samways 2009a), 2011 to measure habitat recovery (Simaika & Samways 2008) and select sites for conservation (Simaika & Samways 2009b). Previous work has shown a strong correlation between adult dragonfly scores and macroinvertebrate scores (Simaika & Samways 2011; Smith, Samways & Taylor 2007). An advantage of the DBI over conventional macroinvertebrate indices is that it operates at the species level and is therefore highly sensitive to habitat condition. It, therefore, has good potential for environmental assessment and monitoring freshwater biodiversity and quality, alongside SASS (Simaika & Samways 2011). The low field effort required to obtain a DBI score for a site also makes this a low-cost and readily applied method. Odonata are therefore particularly useful for monitoring freshwater quality and landscape physiognomy around riverine and other aquatic habitats in South Africa’s PAs (Samways 1993b).

Ant monitoring in South Africa has gained significant momentum over the last decade. For example, the Iimbovane project (http://academic.sun.ac.za/iimbovane) is a spatially extensive and temporarily replicated ant monitoring programme (currently focussed in the Western Cape), where equivalent information on invertebrates is rare (Driver et al. 2005; Koch et al. 2000). It thus delivers baseline data of species distributions in different vegetation types in two biodiversity hotspots (the Fynbos biome and the Succulent Karoo biome), against which the impact of future environmental change can be assessed (Braschler et al. 2010). The project uses sites that are disturbed in a variety of ways and control sites that are typically located within PAs. Spatially and temporally replicated surveys are used to detect trends and assess natural variation, including variation due to seasonality, climatic events or fire events. However, such surveys face many challenges, including securing expertise and funding for long periods. Continuity was built into the Iimbovane project by combining monitoring with a long-term outreach programme (teaching biodiversity to high school learners participating in the monitoring; see Braschler [2009]) and by involving technical staff on long-term contracts. Including both pristine and disturbed sites means that the data may be used for a variety of purposes. It is inevitable that much conservation will have to be done outside PAs, including transformed landscapes, which makes monitoring disturbed sites necessary. Control sites in PAs are particularly valuable and serve to separate various fine-scale and large-scale disturbance effects. Furthermore, where restoration is the goal for conservation, baseline data are needed on what is natural to an area and this information is often best obtained from control sites with natural vegetation such as those found inside PAs.

Faced with adopting and implementing an invertebrate monitoring programme, the KNP has also assessed the feasibility, resource requirements and cost of ant monitoring in the park. Ants were selected as one of three possible groups (the others were dung beetles and butterflies) because of the presence of taxonomic expertise in the park, a history of active collaboration with external researchers on this taxon, good baseline data on ant assemblages in the KNP being available and the ubiquitousness and relevance of the taxon in the area (Parr & Chown 2001; Parr et al. 2004). To assess the costs associated with ant monitoring, ants were sampled from 34 plots (2000 m² each) using 20 pitfall traps per plot. Traps were left open in the field for 5 days. Pitfall trapping was chosen because it is widely used for ant sampling and is considered efficient in savanna ecosystems (Parr & Chown 2001; Underwood & Fisher 2006). Sampling was done from 2003 to 2005 and again in 2009. The optimum workforce needed for the field work of a single 2000 m² area was found to be four people, which is similarly labour intensive to other monitoring projects in SANParks (Figure 2). The total time spent sampling and identifying ants on a single plot was 460 min, including 80 min spent on setting and removing traps, 80 min for screening an average of 300 individual ants, and 300 min for identifying them to species level under the microscope. Although comparatively inexpensive (see below), ant sampling was found to be more time consuming than other monitoring projects in the KNP (Figure 2). This may be mitigated by constantly using the same trained assistants and applying simplified assessments such as those developed for Australia (Andersen et al. 2002). The resources required for postcollection sorting, identification, curation and analyses are often not carefully considered and these are all critical for the collection and long-term use and storage of monitoring information. Considered per appropriate sample area, ant monitoring was also found to be one of the least expensive monitoring projects in the KNP (Figure 2).

Ants have been demonstrated to provide particularly valuable management-based information across a range of regions of the world, in particular for invasive species monitoring, detecting trends in functionally important species and those of specific conservation concern and assessing long-term changes in ecosystems and the consequences of management action (Underwood & Fisher 2006). This series of factors makes the Formicidae particularly suitable for inclusion in PA monitoring systems in South Africa.

FIGURE 2:Estimated total resource costs per unit sampling area of the ant monitoring project in the Kruger National Park compared with six other monitoring projects in SANParks (different area for each project, but the minimum relevant unit area is used in each case).

An example of habitat-focused monitoring is the streams in the TMNP. Because of the evolutionary significance and high levels of endemism, both streams and their associated Gondwanan biota could be prioritised for monitoring. Further reasons for doing so include the fact that Gondwanan taxa tend to be stenothermal (cold-adapted) (McKie, Cranston & Pearson 2004) and thus are most at risk from elevated temperatures and the synergistic influence of reduced precipitation predicted by models of global warming for the Western Cape (De Wit & Stankiewicz 2006). Baseline data for future monitoring in this habitat would include a thermal and biotic profile of a selected stream, encompassing both spatial (longitudinal stream profile) and seasonal variability. Against this baseline, future monitoring would then be conducted to detect the influences of climate change on both water temperature and invertebrate biota. Because water temperature is also influenced by flow, monitoring would simultaneously provide a baseline against which effects of the possible abstraction of the Table Mountain aquifer on the stream and its biota can be assessed. A year’s worth of baseline data have already been collected for this project at Window Gorge Stream (Ketley 2009) and the project has been transferred to SANParks for future monitoring. Regular, ongoing monitoring is essential in this system if trends in change drivers (especially changing flow rates, water temperatures and biotic responses) are to be captured.

Both temporal and small-scale turnover, as well as habitat association, are generally poorly understood in invertebrate communities (Lovell et al. 2010) and would need to be carefully considered in establishing and maintaining a habitat-based or community-based monitoring programme. For example, turnover of epigaeic invertebrates in Afrotemperate forest in the Drakensberg ranged from complete to 50%, even in forests within the same valley (Hamer & Slotow 2009; Uys, Hamer & Slotow 2009). Similarly, temporal turnover between seasons and years may be high for epigaeic invertebrates and flying insects, particularly (Lovell et al. 2010).

Importantly, sufficient information and support are now available for particular taxa to be included practically in monitoring in PAs, and in many instances significant levels of endemism render invertebrates important for monitoring in their own right. Indeed, several current initiatives are set to improve this situation substantially. Good data on species richness, species distributions and population trends remain essential if invertebrates are to be considered in conservation management decisions and if effectiveness of these decisions for their protection is to be assessed. Currently such data are available for only a few key taxa.

The South African Biosystematics Initiative of the National Research Foundation currently provides some funding for invertebrate survey and monitoring research, if projects are well justified and placed in an appropriate conceptual framework. Significant opportunities thus exist to develop collaborations between PA agencies needing data and researchers skilled to provide these data. Collaboration between PA and external researchers on PA invertebrate research can produce valuable baseline data, taxonomic knowledge and inventories, which then enable PAs to (1) integrate invertebrates into PA monitoring programmes in a feasible manner and (2) continue key invertebrate monitoring in the longer term.

• Although invertebrates are relevant and important across a range of PA management objectives, they should be explicitly and clearly linked to these objectives.

• Monitoring activities that involve invertebrates should focus on taxonomically well-known groups that are supported by, for example, good keys, available expertise and published evidence of their value in the specific context. The success of initiatives such as SANSA, Iimbovane and SABCA illustrates the feasibility and value of invertebrate surveys and monitoring in PAs.

• An important point to communicate to conservation planners is that vegetation or other surrogates do not adequately capture invertebrate diversity and that setting targets without considering at least some invertebrates is unlikely to capture invertebrate diversity (e.g. Hamer & Slotow 2009).

• In a South African PA context, dragonflies, ants, butterflies and dung beetles (and increasingly also spiders) are appropriate for monitoring a broad range of management objectives.

• Invertebrate species of special concern represent a particular knowledge gap for PAs and here research is required to identify localised endemics and threatened species along with surveys targeting potentially important areas and taxa. For threatened species, some idea of habitat requirements and basic biology is required for adequate management, as well as approaches to monitoring threatened invertebrate species.

• Similarly, surveys of invasive alien invertebrate species are required along with research on their impacts and approaches to prevent further introductions.

• Profiling of invertebrates and training specialist invertebrate consultants are required to successfully integrate invertebrate conservation into processes legislated by NEMA and its subsidiary Acts. This will allow invertebrates to be explicitly considered and conserved in areas subject to change in land use.

• Collaborative relationships between entomologists, particularly systematists, and PA agencies should be fostered to maximise both research opportunities and opportunities to improve our knowledge of the invertebrate fauna of PAs. Citizen scientists continue to make an extremely valuable contribution to invertebrate surveys and monitoring in South Africa and their potential role in PA monitoring is significant.

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