The rate of evolution of an RNA plant virus has never been estimated using temporally spaced sequence data, by contrast to the information available on an increasing range of animal viruses. Accordingly, the evolution rate of Rice yellow mottle virus (RYMV) was calculated from sequences of the coat protein gene of isolates collected from rice over a 40-year period in different parts of Africa. The evolution rate of RYMV was estimated by pairwise distance linear regression on five phylogeographically defined groups comprising a total of 135 isolates. It was further assessed from 253 isolates collected all over Africa by Bayesian coalescent methods under strict and relaxed molecular clock models and under constant size and skyline population genetic models. Consistent estimates of the evolution rate between 4 ؋ 10 ؊4 and 8 ؋ 10 ؊4 nucleotides (nt)/site/year were obtained whatever method and model were applied. The synonymous evolution rate was between 8 ؋ 10 ؊4 and 11 ؋ 10 ؊4 nt/site/year. The overall and synonymous evolution rates of RYMV were within the range of the rates of 50 RNA animal viruses, below the average but above the distribution median. Experimentally, in host change studies, substitutions accumulated at an even higher rate. The results show that an RNA plant virus such as RYMV evolves as rapidly as most RNA animal viruses. Knowledge of the molecular clock of plant viruses provides methods for testing a wide range of biological hypotheses.The mutation rates of RNA viruses (i.e., the number of nucleotide misincorporations per site and per round of replication) are 10 4 to 10 5 times higher than those of their DNA hosts (8, 9). Such a high mutation rate is attributed to the lack of repair function of the RNA polymerase of these viruses, the short replication times, and the large populations in infected hosts (7). A high mutation rate often results in rapid evolution of RNA animal viruses. This allowed the measure of the evolution rates of an extensive range of animal viruses through the analysis of heterochronous sequences, i.e., sequences of viral genes isolated at different times (11,36). A large variation in the evolution rates of RNA animal viruses was subsequently found and was attributed mostly to differences in replication rates (24,26). Estimates of evolution rates are used increasingly to date the emergence and to reconstruct the population dynamics of major viral epidemics (6).Interestingly, some RNA viruses change little or not at all over time. The best-documented example is an RNA plant virus, Tobacco mild green mosaic virus, which showed no increase in genetic diversity over the 90 years considered, in the longest series of isolates with known isolation times for any virus (20). Indeed, many studies have shown the remarkable genetic stability of RNA plant virus populations from different geographical regions, hosts, and collection times (21). It was claimed that most tobamovirus populations are very stable and do not evolve at a measurable rate (22). It was even observed that populations...