TitleFive palaeobiological laws needed to understand the evolution of the living biota O ne of the cornerstones of evolutionary theory is the idea of differential survival. The fact of death, the termination of the lives of individuals, is central to evolutionary change. For biologists working on the timescales of generations, the importance of differential survival has been well understood and well studied since Darwin. On longer timescales, however, although it is well recognized that there has been extensive termination of unique morphologies and lineages, study of the living biota alone offers relatively little when trying to determine the importance of these extinctions, given that among the living we only have direct data on the survivors. This presents significant challenges when trying to elucidate the evolutionary process that produced the living biota.Fortunately, the fossil record provides unique and invaluable data on the nature of extinction, including its selectivity, recurrence times, magnitudes and causes. While there has been a dramatic increase in the number of integrated palaeontological and neontological studies [1][2][3][4] , from a palaeobiological perspective, the field of biology has not yet fully assimilated the fundamental findings from the fossil record. Here I summarize these findings in the context of their relevance to our understanding of the living biota. I express this knowledge in the form of simple laws (Box 1), in the hope of facilitating the incorporation of the longstanding and fundamental discoveries of palaeontology into the foundations of thinking in evolutionary biology and ecology, something that has yet to be fully realized.
The first law of palaeobiologyLineages become extinct. While the notion of immortality at the level of individuals has been rejected for millennia, it was not until around 1800 that the idea that species could become extinct was established through the discovery of fossils that could not be ascribed to any living species 5 . More formally, the first law can be expressed in terms of the probability that clade will ultimately go extinct, P e :Five palaeobiological laws needed to understand the evolution of the living biota
Charles R. MarshallThe 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 extinc tion and longevity, species richness, origination rates, extinction rates and diversification. These statements of our palaeo biological 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 orig...