Indices of reproductive skew depend on average reproductive success

Several indices of reproductive skew, which quantify the degree of unequal partitioning of reproductive output among members in an animal society, have been proposed. Here we point out the drawbacks of these indices. The most serious problem is the dependence of the indices on mean reproductive success: skew values tend to be larger, as average numbers of offspring decrease, due to random sampling error in numbers of offspring. Thus it is difficult to compare societies with different average lifetime reproductive success using these indices, even though we have presented methods to calculate the expected reproductive skew caused by random sampling error, especially when average numbers of offspring are small, as is often the case with cooperatively breeding vertebrates. As an alternative, we propose using the spatial dispersion indices of population ecology (Morisita's index or its standardized version) for the measurement of reproductive skew. These indices are almost independent of average fecundity and have their own method of testing for random variation in offspring numbers.