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Background: Cichlids are a prime model system in evolutionary research and several of the most prominent examples of adaptive radiations are found in the East African Lakes Tanganyika, Malawi and Victoria, all part of the East African cichlid radiation (EAR). In the past, great effort has been invested in reconstructing the evolutionary and biogeographic history of cichlids (Teleostei: Cichlidae). In this study, we present new divergence age estimates for the major cichlid lineages with the main focus on the EAR based on a dataset encompassing representative taxa of almost all recognized cichlid tribes and ten mitochondrial protein genes. We have thoroughly re-evaluated both fossil and geological calibration points, and we included the recently described fossil †Tugenchromis pickfordi in the cichlid divergence age estimates., Results: Our results estimate the origin of the EAR to Late Eocene/Early Oligocene (28.71 Ma; 95% HPD: 24.43-33.15 Ma). More importantly divergence ages of the most recent common ancestor (MRCA) of several Tanganyika cichlid tribes were estimated to be substantially older than the oldest estimated maximum age of the Lake Tanganyika: Trematocarini (16.13 Ma, 95% HPD: 11.89-20.46 Ma), Bathybatini (20.62 Ma, 95% HPD: 16.88-25.34 Ma), Lamprologini (15.27 Ma; 95% HPD: 12.23-18.49 Ma). The divergence age of the crown haplochromine H-lineage is estimated to 22.8 Ma (95% HPD: 14.40-26.32 Ma) and of the Lake Malawi radiation to 4.07 Ma (95% HDP: 2.93-5.26 Ma). In addition, we recovered a novel lineage within the Lamprologini tribe encompassing only Lamprologus of the lower and central Congo drainage with its divergence estimated to the Late Miocene or early Pliocene. Furthermore we recovered two novel mitochondrial haplotype lineages within the Haplochromini tribe: 'Orthochromis' indermauri and 'Haplochormis' vanheusdeni., Conclusions: Divergence time estimates of the MRCA of several Tanganyika cichlid tribes predate the age of the extant Lake Tanganyika basin, and hence are in line with the recently formulated "Melting-Pot Tanganyika" hypothesis. The radiation of the 'Lower Congo Lamprologus clade' might be linked with the Pliocene origin of the modern lower Congo rapids as has been shown for other Lower Congo cichlid assemblages. Finally, the age of origin of the Lake Malawi cichlid flock agrees well with the oldest age estimate for lacustrine conditions in Lake Malawi.

Five new rheophilic haplochromine cichlid species are described from the Upper Congo drainage of Zambia and the Democratic Republic of the Congo: Orthochromis mporokoso sp. nov. and O. katumbii sp. nov. from the Bangwelu-Mweru ecoregion, O. kimpala sp. nov. and O. gecki sp. nov. from the Upper Lualaba ecoregion, and O. indermauri sp. nov. from the Lufubu River of the Lake Tanganyika ecoregion. Orthochromis kimpala sp. nov, O. gecki sp. nov., and O. indermauri sp. nov. are distinguished from all currently valid species of the genus Orthochromis Greenwood 1954, except for O. torrenticola (Thys van den Audenaerde 1963), by the presence of eggspots or eggspot-like maculae on the anal fin (vs. no eggspots). The three species can be easily distinguished from O. torrenticola by having three anal spines (vs. four anal spines). Moreover, all five new species can be individually distinguished from all currently known rheophilic taxa placed in the genera Orthochromis, Schwetzochromis Poll 1948 and the rheophilic species of the genus Haplochromis Hilgendorf 1888 (e.g. H. bakongo Thys van den Audenaerde 1964, H. snoeksi Wamuini Lunkayilako Vreven 2010, H. vanheusdeni Schedel et al. 2014) either based on meristic values, morphometric distances and colouration patterns, or on a combination of them.