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Knowing the number of species ( S ) in an ecological system (i.e., species richness) is of primary importance to understanding biodiversity. Willis (1) states that the only “statistically admissible” way to estimate S is by modeling the number of species with n = 1, 2, 3, ... individuals, which is known as the frequency ratio method (2). Once this is accomplished, Willis (1) states that S can then be estimated via extrapolation through probabilistic models. However, extrapolations of richness estimates across orders of magnitude can be unsound (1, 3) and the data required by statistical estimators, such as frequency ratios, do not … [↵][1]1To whom correspondence may be addressed. Email: ken{at}weecology.org or lennonj{at}indiana.edu. [1]: #xref-corresp-1-1

Reply to Willis: Powerful predictions of biodiversity from ecological models and scaling laws