Five palaeobiological laws needed to understand the evolution of the living biota.

The foundations of several disciplines can be expressed as simple quantitative laws, for example, Newton's laws or the laws of thermodynamics. Here I present five laws derived from fossil data that describe the relationships among species extinction and longevity, species richness, origination rates, extinction rates and diversification. These statements of our palaeobiological knowledge constitute a dimension largely hidden from view when studying the living biota, which are nonetheless crucial to the study of evolution and ecology even for groups with poor or non-existent fossil records. These laws encapsulate: the critical fact of extinction; that species are typically geologically short-lived, and thus that the number of extinct species typically dwarfs the number of living species; that extinction and origination rates typically have similar magnitudes; and, that significant extinction makes it difficult to infer much about a clade's early history or its current diversity dynamics from the living biota alone. Although important strides are being made to integrate these core palaeontological findings into our analysis of the living biota, this knowledge needs to be incorporated more widely if we are to understand their evolutionary dynamics.