The evolutionary species pool hypothesis and patterns of freshwater diatom diversity along a pH gradient

Aim  To interpret the unimodal relationship between diatom species richness and lake pH within the context of the evolutionary species pool hypothesis (SPH). We test the following primary prediction arising from the SPH: the size of the potential species pool (PSP) will increase along a gradient representing the historical commonness of different pH environments (pH commonness). To do this we assume that the present‐day spatial dominance of near‐neutral pH conditions compared with acidic and alkaline conditions reliably mimics the relative spatial availabilities of historical pH conditions among freshwater lakes. We also determine whether local richness represents a constant proportion of PSP size along the pH commonness gradient. Location  Two hundred and thirty‐four lakes distributed over a 405,000 km 2 region of the north‐eastern United States of America. Methods  Sediment diatom morphospecies lists and pH data were acquired from the US Environmental Protection Agency's Environmental Monitoring and Assessment Program (EMAP) website. Using 248 morphospecies that occurred in at least 10 of the 234 lakes, four different measures of PSPs were calculated along the pH gradient. Local species richness was equated with the number of species occurring within the lake. Alpha diversity was equated with the average species richness of lakes with similar pH values. A combination of statistical methods were employed, including correlations, quadratic regression and piecewise regression. Results  PSP size increased significantly with pH commonness for all four measures of PSP size, thus supporting the primary prediction of the evolutionary SPH. Local richness comprised a larger proportion of the PSP within acidic lakes than within circumneutral lakes. Alpha diversity and lake species richness both increased significantly with pH commonness, but the former did so in a two‐step fashion. We test and reject several alternative contemporary time‐scale explanations for our findings. Main Conclusions  Our findings are consistent with the hypothesis that diatom taxonomic richness is presently lower within acidic and highly alkaline lakes than in circumneutral lakes owing to the limited opportunity in space and/or time for the evolution of suitably adapted species. Whereas ecological processes can explain why certain species are excluded from particular habitats, e.g. acidic lakes, they cannot account for why so few species are adapted to those habitats in the first place.