Arrival order and release from competition does not explain why haplochromine cichlids radiated in Lake Victoria

The frequent occurrence of adaptive radiations on oceanic islands and in lakes is often attributed to ecological opportunity resulting from release from competition where arrival order among lineages predicts which lineage radiates. This priority effect occurs when the lineage that arrives first expands its niche breadth and diversifies into a set of ecological specialists with associated monopolization of the resources. Later-arriving species do not experience ecological opportunity and do not radiate. While theoretical support and evidence from microbial experiments for priority effects are strong, empirical evidence in nature is difficult to obtain. Lake Victoria (LV) is home to an exceptional adaptive radiation of haplochromine cichlid fishes, where 20 trophic guilds and several hundred species emerged in just 15 000 years, the age of the modern lake that was preceded by a complete desiccation lasting several thousand years. However, while about 50 other lineages of teleost fish also have established populations in the lake, none of them has produced more than two species and most of them did not speciate at all. Here, we test if the ancestors of the haplochromine radiation indeed arrived prior to the most competent potential competitors, 'tilapias' and cyprinids, both of which have made rapid radiations in other African lakes. We assess LV sediment core intervals from just before the desiccation and just after refilling for the presence of fossil fish teeth. We show that all three lineages were present when modern LV began to fill with water. We conclude that the haplochromines' extraordinary radiation unfolded in the presence of potentially competing lineages and cannot be attributed to a simple priority effect.