19The diversity of species inhabiting freshwater relative to marine habitats is striking, given that 20 freshwater habitats make up less than 1% of Earth's water. The most commonly proposed 21 mechanism for this pattern is that freshwater habitats are more fragmented than marine habitats, 22 resulting in increased opportunities for allopatric speciation in freshwater and thus increased 23 speciation rates relative to marine habitats. However, some authors suggest that speciation is 24 faster in sympatry than allopatry, as illustrated by lacustrine radiations such as African cichlids. 25 Sympatric speciation may be common in lakes, while allopatric speciation may be dominant in 26 rivers. If allopatric speciation is inherently slower than sympatric speciation, we might expect 27 differences in rates of speciation between lineages primarily confined to riverine versus 28 lacustrine habitats. These differences within freshwater habitats may be important to consider 29 when comparing diversification rates between freshwater and marine lineages. Here I compared 30 rates of speciation associated with ray-finned fishes in marine, riverine and lacustrine habitats. I 31 also used geographic range data to compare rates of speciation associated with allopatric and 32 sympatric speciation modes in freshwater fishes. I found that lakes had faster rates of speciation 33 than other aquatic habitats. However, most freshwater fish diversity arose through allopatric 34 speciation in rivers, despite their slower speciation rate. Speciation rates were more strongly 35 correlated with habitat (riverine or lacustrine) than present-day range overlap (allopatry or 36 sympatry). Surprisingly, riverine and marine habitats had similar rates of speciation. Models of 37 evolution suggest that lacustrine habitats are evolutionary unstable, and biased transition rates 38 explain the dearth of lacustrine species. Collectively, these results suggest that while allopatric 39 speciation is important for generating freshwater diversity, this mechanism may not be the cause 40 of faster speciation rates between freshwater and marine habitats. 41 3 KEY WORDS 42 43 Fishes, allopatric speciation, sympatric speciation, speciation rates, lacustrine radiations, 44 diversification 45 46 47 48 49 50