Historical contingency and the purported uniqueness of evolutionary innovations.

Many events in the history of life are thought to be singular, that is, without parallels, analogs, or homologs in time and space. These claims imply that history is profoundly contingent in that independent origins of life in the universe will spawn radically different histories. If, however, most innovations arose more than once on Earth, histories would be predictable and replicable at the scale of functional roles and directions of adaptive change. Times of origin of 23 purportedly unique evolutionary innovations are significantly more ancient than the times of first instantiation of 55 innovations that evolved more than once, implying that the early phases of life's history were less replicable than later phases or that the appearance of singularity results from information loss through time. Indirect support for information loss comes from the distribution of sizes of clades in which the same minor, geologically recent innovation has arisen multiple times. For three repeated molluscan innovations, 28-71% of instantiations are represented by clades of five or fewer species. Such small clades would be undetectable in the early history of life. Purportedly unique innovations either arose from the union and integration of previously independent components or belong to classes of functionally similar innovations. Claims of singularity are therefore not well supported by the available evidence. Details of initial conditions, evolutionary pathways, phenotypes, and timing are contingent, but important ecological, functional, and directional aspects of the history of life are replicable and predictable.