EVOLUTIONARY theory has not explained how competition among lower level units is suppressed in the formation of higher-level evolutionary units1,2. For example, the key problem of early evolution is how small, individual replicators formed cooperative groups of sufficient complexity to allow accurate copying of the genetic material3. The puzzle is why parasites did not subvert the formation of cells by obtaining benetics benefits from the group without contributing to shared traits that enhance reproduction4. These parasites would outcompete other replicators within the cell, disrupting reproductive fairness among subunits and destroying the functional coherence of the group. A similar problem arose at a later evolutionary stage with the orderly mendelian segregation of subunits (chromosomes) within cells, and reproductive fairness continued to be a problem in the evolution of insect5 and human societies6. Here I present a simple model to show how reproductive fairness evolves among subunits to create functional coherence and higher-level units. Self-restraint, which evolves according to the kin-selection coefficient of relatedness, is not sufficient: mutual policing and enforcement of reproductive fairness are also required for the evolution of increasing social complexity.
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