The evolution of methods for establishing evolutionary timescales

The fossil record is well-known to be incomplete. If read literally, it will also give a distorted view of the history of species divergence and extinction, because different species have different propensities to fossilise, the amount of rock fluctuates over geological time scales, and so on. Even so patterns in the fossil evidence allow us to assess the incompleteness of the fossil record. While the molecular clock can be used to extend the time estimates from fossil species to lineages not represented in the fossil record, fossils are the only source of information concerning absolute (geological) times in molecular dating analysis. We review different ways of incorporating fossil evidence in modern clock dating analyses, including node calibrations where lineage divergence times are constrained using probability densities and tip calibrations where fossil species at the tips of the tree are assigned dates from dated rock strata. While node calibrations are often constructed by a crude assessment of the fossil evidence and thus involves arbitrariness, tip calibrations may be too sensitive to the prior on divergence times or the branching process and too much affected by well-known problems of morphological character evolution, such as environmental influence on morphological phenotypes, correlation among traits and convergent evolution in disparate species. We discuss the utility of time information from fossils in phylogeny estimation and the search for ancestors in the fossil record.