LAUSR

Aim: The evolutionary causes of the latitudinal diversity gradient are debated. Hypotheses have ultimately invoked either faster rates of diversification in the tropics, or more time for diversification due to the tropical origins of higher taxa. Here we perform the first test of the diversification rate and time hypotheses in freshwater ray-finned fishes, a group comprising nearly a quarter of all living vertebrates. Location: Global. Time period: 368-0 mya. Major taxa studied: Extant freshwater ray-finned fishes. Methods: Using a mega-phylogeny of actinopterygian fishes and a global database of occurrence records, we estimated net diversification rates, the number of colonizations and regional colonization times of co-occurring species in freshwater drainage basins. We used Generalized Additive Models to test whether these factors were related to latitude. We then compared the influence of diversification rates, colonization numbers, colonization times and surface area on species richness, and how these factors are related to each other. Results: While both diversification rates and time were related to richness, time had greater explanatory power and was more strongly related to latitude than diversification rates. Other factors (basin surface area, number of colonizations) also helped explain richness but were unrelated to latitude. The world9s most diverse freshwater basins (Amazon, Congo rivers) were dominated by lineages with Mesozoic origins. The temperate groups dominant today arrived near the K-Pg boundary, leaving comparatively less time to build richness. Diversification rates and colonization times were inversely related: recently colonized basins had the fastest rates, while ancient species-rich faunas had slower rates. Main conclusions: We concluded that time is the lead driver of latitudinal richness disparities in freshwater fish faunas. We suggest that the most likely path to building very high species richness is through diversification over long periods of time, rather than diversifying quickly.