Taxonomic, ecological and biogeographic re-evaluation of temperate stream fishes in Southern Africa : a case study of the enteromius anoplus complex
Authors:   , Kambikambi Manda Juliet 
Published: November 2021

Worldwide, cryptic species, which comprise two or more distinct species that are classified as one due to their morphological similarity, are more common than previously thought. In South Africa, the chubbyhead barb, Enteromius anoplus (Weber, 1897), is the most widespread freshwater fish species. The widespread occurrence of this species across different river basins raised questions on its taxonomic status due to the likely occurrence of hidden diversity. This further raised the need to evaluate the ecological patterns associated with likely occurrence of such hidden diversity. Phylogenetic analyses of the cytochrome b gene of E. anoplus populations across South Africa revealed the occurrence of four genetically distinct species. These findings supported the restoration Enteromius cernuus (Barnard 1943) stat. rev. from the Olifants River system; the restoration of Enteromius anoplus sensu stricto (Weber 1897) stat. rev. from the Gouritz River system; the redescription of Enteromius oraniensis (Barnard 1943) stat. rev. from the Orange River system; and the description of a new species, Enteromius mandelai sp. nov. from the Eastern Cape rivers. Analyses of morphometric and meristic data revealed that these species were differentiated based on barbel length and number of lateral line scales. Ecological studies on Enteromius mandelai were conducted within headwater streams of major river systems in the Eastern Cape region. These headwater streams had different catchment characteristics ranging from near-natural to anthropogenically-perturbed mainly due to agriculture-related disturbances and invasion of riparian zones by non-native plants. Stable isotope analysis provided evidence of trophic plasticity for E. mandelai across the different headwater streams. Specifically, E. mandelai exhibited large isotopic niches in agriculture-disturbed headwater streams, whereas small isotopic niches were observed in headwater streams invaded by non-native plants. Because of the evidence of trophic plasticity for E. mandelai, this study further hypothesized that this species would exhibit variable environmental niche patterns related to its distribution, abundance and the presence of contra-specifics due to different abiotic factors across its range. Enteromius mandelai was generally most abundant at sites with high total dissolved solids and sand substratum, which was characteristic of large pool habitats, and least abundant at sites with high turbidity. In the Great Fish River, E. mandelai occurred as the only species in most headwater sites, whereas in the Keiskamma and Buffalo River systems, it co-occurred with Amatolacypris trevelyani and Sandelia bainsii. Enteromius mandelai was, nevertheless, found to be ubiquitous together with having wide environmental niches, and exhibited no discernible patterns in its habitat associations. The taxonomic re-evaluation of the E. anoplus complex, which provided evidence of the existence of new species, has implications for the biogeographic zonation of the ichthyofauna of the Cape Fold Ecoregion (CFE), Amathole-Winterberg Highlands Ecoregion (AWHE) and the Southern Temperate Highveld Ecoregion (STHE). For example, the currently recognised southern limit of the STHE, which encompasses the AWHE, is based on the occurrence of E. anoplus, Enteromius pallidus and Labeo umbratus. However, evidence from this and other recent studies indicate the occurrence of either new species or distinct lineages in the southern-most section of the STHE, which is consistent with the patterns observed in the CFE. Comparison of taxonomic composition showed that species in the eastern part of the CFE were similar to species in the AWHE and the southern-most section of the STHE. Because of the close affinities among most of the new taxa in these regions, this study proposes that the current delimitation of the CFE should be reviewed by extending it to include the AWHE and sections of the STHE.