Age estimation from molecular sequences has emerged as a powerful tool for inferring when a plant lineage arrived in a particular area. Knowing the tenure of lineages within a region is key to understanding the evolution of traits, the evolution of biotic interactions, and the evolution of floras. New analytical methods model change in substitution rates along individual branches of a phylogenetic tree by combining molecular data with time constraints, usually from fossils. These 'relaxed clock' approaches can be applied to several gene regions that need not all have the same substitution rates, and they can also incorporate multiple simultaneous fossil calibrations. Since 1995, at least 100 plant biogeographic studies have used molecular-clock dating, and about a fifth has used relaxed clocks. Many of these report evidence of long-distance dispersal. Meta-analyses of studies from the same geographic region can identify directional biases because of prevailing wind or water currents and the relative position and size of landmasses.Inferring past plant dispersal: from which data and why? The geographic ranges of the lineages seen today are often interrupted by large expanses of inhospitable habitat, for example, an ocean. This raises questions about how and when they acquired their disjunct habitats. When lineages arrive in new habitats they will usually diverge and sometimes speciate, therefore, dating dispersal and lineage splits has broad implications for understanding rates of diversification [1], the evolution of biota [2,3] and associated faunas [4][5][6], and the time taken for species to adapt to novel environments [7][8][9][10]. Recent biogeographic studies that have included time estimates have revealed strikingly fast diversification rates in some floras, such as the Cape Region of South Africa, the recency of species accumulation even in ancient lineages, such as ferns [1], and the persistence of precarious mutualisms over many millions of years, such as the interactions between figs and fig wasps and between yucca and yucca moths [4,6].The ideal materials from which to estimate the ages of lineages and their presence in particular places are abundant fossils. However, isolated fossils often provide only screenshots from which we extrapolate in time and space. Extinction followed by recolonization is particularly difficult to infer from the fossil record. It is often more parsimonious to interpret the history of taxa with extended fossil records in a framework of continuity [11,12]. Yet, there is ample direct evidence of rapid species expansions following dispersal from the study of invasive species, phylogeographic molecular data, and natural experiments such as Krakatau, an island sterilized by a massive eruption in 1883 [13][14][15][16].The idea of dating the divergence of lineages from the time-dependent change of their genomes, the molecular clock hypothesis, is 40 years old [17], and plant geographers have applied sequence-or protein-based dating since the 1990s [18][19][20][21][22][23][24][25][...