Can the evolutionary-rates hypothesis explain species-energy relationships?

1. There is growing consensus that much of the marked broad-scale spatial variation in species richness is associated with variation in environmental energy availability, but at least nine principal mechanisms have been proposed that may explain these patterns. 2. The evolutionary-rates hypothesis suggests that high environmental energy availability elevates rates of molecular evolution, promoting faster speciation, so that more species occur in high-energy areas because more evolve there. Direct tests of this hypothesis are rare and their conclusions inconsistent. Here we focus on assessing the support for its underlying assumptions. 3. First, the evolutionary-rates hypothesis assumes that high energy levels promote mutation. There is certainly evidence that high levels of ultraviolet radiation increase mutation rates. High temperatures may also reduce generation times and elevate metabolic rates, which may promote mutation. On balance, data support a link between rates of metabolism and mutation, but a link between the latter and generation time is more equivocal and is particularly unlikely in plants. 4. Second, the evolutionary-rates hypothesis assumes that mutation rates limit speciation rates. This may be true if all else was equal, but correlations between mutation and speciation are probably very noisy as many other factors may influence rates both of sympatric and allopatric speciation, including the occurrence of physical isolation barriers, the magnitude of selection and population size. 5. Third, the evolutionary-rates hypothesis assumes that there is a strong correlation between current and historical energy levels. Factors such as tectonic drift may weaken such relationships, but are likely to have had negligible effects over the time period during which the majority of extant species evolved. 6. Fourth, the evolutionary-rates hypothesis assumes that changes in species ranges following speciation do not sufficiently weaken the correlation between the rate of speciation in an area and species richness. The ranges of many species appear to alter dramatically following speciation, and this may markedly reduce the strength of the relationship, but to what extent is unclear. 7. In sum, the degree to which the evolutionary-rates hypothesis can explain spatial variation in species richness remains surprisingly uncertain. We suggest directions for further research.