corresponds to the average Shannon index than beta is equal to the average Kullback-Leibler divergence.

When there are two subunits, and presence-absence data are used, this can be calculated with the following equation:

where, S₁= the total number of species recorded in the first community, S₂= the total number of species recorded in the second community, and c= the number of species common to both communities.

Whittaker's species turnover

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