The Concept of Evenness/unevenness: Less Evenness or More Unevenness?

Acta Biotheoretica 70 (1):1-28 (2021)
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Abstract

While evenness is understood to be maximal if all types (species, genotypes, alleles, etc.) are represented equally (via abundance, biomass, area, etc.), its opposite, maximal unevenness, either remains conceptually in the dark or is conceived as the type distribution that minimizes the applied evenness index. The latter approach, however, frequently leads to conceptual inconsistency due to the fact that the minimizing distribution is not specifiable or is monomorphic. The state of monomorphism, however, is indeterminate in terms of its evenness/unevenness characteristics. Indeed, the semantic indeterminacy also shows up in the observation that monomorphism represents a state of pronounced discontinuity for the established evenness indices. This serious conceptual inconsistency is latent in the widely held idea that evenness is an independent component of diversity. As a consequence, the established evenness indices largely appear as indicators of relative polymorphism rather than as indicators of evenness. In order to arrive at consistent measures of evenness/unevenness, it seems indispensable to determine which states are of maximal unevenness and then to assess the position of a given type distribution between states of maximal evenness and maximal unevenness. Since semantically, unevenness implies inequality among type representations, its maximum is reached if all type representations are equally different. For given number of types, this situation is realized if type representations, when ranked in descending order, show equal differences between adjacent types. We term such distributions “stepladders” as opposed to “plateaus” for uniform distributions. Two approaches to new evenness measures are proposed that reflect different perspectives on the positioning of type distributions between the closest stepladders and the closest plateaus. Their two extremes indicate states of complete evenness and complete unevenness, and the midpoint is postulated to represent the turning point between prevailing evenness and prevailing unevenness. The measures are graphically illustrated by evenness surfaces plotted above frequency simplices for three types, and by transects through evenness surfaces for more types. The approach can be generalized to include variable differences between types (as required in analyses of functional evenness) by simply replacing types with pairs of different types. Pairs, as the new types, can be represented by their abundances, for example, and these can be modified in various ways by the differences between the two types that form the pair. Pair representations thus consist of both the difference between the paired types and their frequency. Omission of pair frequencies leads to conceptual ambiguity. Given this specification of pair representations, their evenness/unevenness can be evaluated using the same indices developed for simple types. Pair evenness then turns out to quantify dispersion evenness.

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