Parasite introduction to the endangered western leopard toad : spill over or spill back?
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South Africa has a diverse anuran fauna consisting of 161 described species. Amphibians are suffering large-scale regressions due to various threats: (1) pollution, (2) habitat engineering and (3) invasive species. The endangered Western Leopard toad (Sclerophrys pantherina) endemic to the Western Cape is currently experiencing major external pressures from these above-mentioned threats. The latter being a local invader, namely the Guttural toad (Sclerophrys gutturalis), which (through human-assisted translocation) was introduced from KwaZulu-Natal into the native range of the Western Leopard Toad. The direct effects such as predation and competition have received extensive attention, seeing that invasive species can have devastating effects on native fauna. However, a more neglected field of research is the indirect threats invasive species pose to the native fauna such as interactions with infectious agents, which include spill back and spill over of parasites. The present study aimed to understand this interaction by focusing on the relationship between parasites, native species, and invasive species. The research examined mechanisms such as spill back and spill over for a better understanding of the indirect drives and consequences invasive species may hold. Morphological markers (light microscopy and scanning electron microscopy) as well as molecular markers (COI and 28S) were applied to survey parasites found in collected toads from five populations: (1) native Guttural Toad in KwaZulu-Natal; (2) invasive Guttural Toad isolated in Western Cape; (3) invasive Guttural Toad and native Western Leopard Toad in Western Cape; (4) native Western Leopard Toad isolated in Western Cape; and (5) native Guttural Toad from Potchefstroom. Parasites that were observed was a nasal mite Lawrencarus eweri (Lawrence, 1952); a lung nematode Rhabdias cf. africana Kuzmin, 2001; intestinal nematode Cosmocerca sp. Diesing, 1861; intestinal trematode Mesocoelium cf. monodi Dolfus, 1929; two blood parasites Hepatozoon ixoxo Netherlands, Cook, & Smit 2014, as well as Trypanosoma sp. Gruby, 1843. It was found that the introduced species may have vacated their parasites throughout the invasion, possibly due to the ‘enemy release hypothesis’ (Marr et al. 2008). This can enhance the invaders competitive ability as defence costs against parasites can be decreased and reproductive rates can increase (Hatcher & Dunn, 2011). The native Western Leopard Toad population co-existing with invasive Guttural Toad was found to contain less parasites than the isolated native Western Leopard Toad population. In this case, it is possible that the invader decreased the parasite loads of the native population by acting as a ‘sink’ for native parasites (Kelly et al., 2009). If so, native parasites are taken up by the invader but fail to complete their life cycle due to disorientation and lack of co-evolutionary history (Hempel et al., 2003). Encysted nematodes (presumably third-stage larvae), collected from native Western Leopard Toads and invasive Guttural Toads from the Western Cape, appear to have a host-size and niche specificity rather than a specificity to the host itself. Few life cycles have been described for nematodes in South African fauna to which a toad acts as an intermediate host for the third-stage larvae. Thus, without further identification and molecular studies it is uncertain whether these cysts in native Western Leopard Toads and invasive Guttural Toads are a result of spill back or spill over. However, each case of parasite-host relationship is unique since the relationship can be dynamic. This makes it difficult to predict the consequences. Furthermore, as is the case with restoration ecology, irreversible changes may have to be accepted in certain ecosystems that are subject to invading parasites and their introduced hosts (Dunn & Hatcher, 2014).