The Great Karoo and Namaqualand of South Africa are home to a species complex of morphologically conserved lizards that occur in allopatry (Karoo: Cordylus aridus, Cordylus cloetei, Cordylus minor; Namaqualand: Cordylus imkeae). However, there are negligible morphological differences and a lack of obvious physical or climatic barriers, particularly among the three Karoo species. We hypothesized that poor geographic coverage in previous studies and lack of an explicit species concept has caused taxonomic inflation. We therefore tested species boundaries by examining multiple criteria: multi-gene phylogenetics, niche distribution modelling and re-examination of diagnostic morphological features with a larger sample size. We found that C. aridus, C. cloetei and C. minor lack diagnosable differences for both genetics and morphology. Distribution modelling, ranging from present day to the last interglacial period, show connectivity has been maintained especially during cooler periods. Conversely, C. imkeae is morphologically diagnosable, genetically distinct and lacks connectivity with the other taxa. By evaluating multiple operational criteria, we conclude that the C. minor species complex comprises only two species, C. minor (with C. aridus and C. cloetei as junior synonyms) and C. imkeae, demonstrating that species defined from inadequate data and lack of an explicit species concept can lead to taxonomic inflation.

Climate shifts during the Quaternary Period have driven changes in regional range dynamics for many species, influencing population structure of species and in some cases promoting speciation. Within southern Africa, the psammophine snakes Psammophis trinasalis and P. namibensis were historically considered subspecies of P. leightoni but were elevated to species rank based on ecological differences. Preliminary phylogenetic analyses suggested intraspecific, not interspecific genetic variation between these taxa, but this finding was based on very limited data and could not be confirmed. To assess the level of genetic differentiation within the P. leightoni species complex, we explored the evolutionary history of these snakes by combining phylogenetic analyses, species distribution modelling and an examination of morphology. We generated a comprehensive, multi-gene phylogeny for Psammophis that included wider geographic sampling of the three species in the complex. Using this phylogeny, Bayesian and distance-based species delimitation analyses showed intraspecific, not interspecific divergences between taxa in the complex, suggesting that they collectively represent a single taxon. Furthermore, non-metric multidimensional scaling analysis of scalation characters showed no differences between the species. Moreover, palaeo-modelling at three time periods since the last interglacial period suggest that there have been varying levels of connectivity between these taxa, which has likely facilitated gene flow between them. Given the evidence, we propose that the P. leightoni complex represents a single species and therefore formally synonymise the three species.

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