Quantifying the level of eusociality

The evolution of animal societies in which some individuals forego direct reproduction to help others to reproduce poses an evolutionary paradox. Societies where all individuals reproduce equally and societies where a single individual completely monopolizes reproduction represent the end points of a continuum of variance in the reproductive output among group members. This led Sherman et al. (1995) to propose that cooperative breeding and eusociality (a term originally applied only to insects) are not discrete phenomena. Rather they form a continuum whose main difference is the extent to which individuals forego their own reproductive opportunity to help other members of the group. Here we present a new index: the eusociality index. It quantifies the decrease in direct reproduction of group members as a resut of altruistic acts directed to other members of the group (i.e. a measure of the level of eusociality). The rationale for this index lies in the fundamental duality of the reproductive process, in which organisms supply two distinct elements: (i) genetic material (genes); and (ii) power (energy). In non-eusocial animals, all individuals transmit genes and power in the same ratio (notwithstanding individual variance in offspring size and parental investment). By contrast, amongst eusocial animals some individuals contribute proportionally more to gene transfer, and others more to energy, resulting in high interindividual variation in the ratio of gene to power transfer.