We have estimated the time for the last commoancor of extant seed plants by using molecular clocks constructed from the ofthe chloroplastic gene coding for the large subunit of ribulose-1,5-blphosphate carboxylase/oxygenase (rbcL) and the nuclear gene coding for the snail subunit of rRNA (Rrnl8). Phylogenetic analyses of nucleodde sequences indicated that the earliest divergence of extant seed plants is likely represented by a split between conufer-cycad and ansperm linea . Relative-rate tests were used to assess homogeneity of substution rates among lineages, and annual anglosperms were found to evolve at a faster rate than other taxa for rbcL and, thus, these sequences were excluded from construction of molecular clocks. Five distinct molecular cocks were calibrated using substitution rates for the two genes and four divergence times based on fossil and published molecular cock estimates. The five mated times for the last common ancestor of extant seed plants were in agreement with one another, with an average of 285 million years and a range of 275-290 million years. This implies a subsantily more recent ancestor of all extant seed plants than suggsed by some theories of plant evolution.Although progymnosperms are accepted as the ancestral group to all seed plants, the time when the five groups of extant seed plants (cycads, conifers, Ginkgo, Gnetales, and angiosperms) shared their last common ancestor is unclear because of uncertainty about ancestor-descendant relationships between progymnosperms and seed plants (1, 2). Beck (2-5), although not discussing angiosperms, proposed that the two main lineages of gymnosperms, coniferopsids (conifers, Ginkgo, and Gnetales) and cycadopsids (cycads), diverged independently from two lineages of the progymnosperms, with coniferopsids from Archaeopteridales and cycads from Aneurophytales. Because Archaeopteridales and Aneurophytales appeared in the Middle Devonian, Beck's hypothesis implies that the two gymnosperm lineages diverged about 375 million years (Myr) ago. Alternatively, Rothwell (6, 7) hypothesized that the two gymnosperm lineages both evolved from Aneurophytales via the seed fern complex, suggesting a more recent diversification of extant seed plants.Phylogenetic analysis of conserved gene sequences from extant plant taxa has contributed tangibly to our understanding of the early diversification of seed plants (8-11). Chloroplast and nuclear gene sequences have also been used to construct molecular clocks and to estimate the times for key events in the evolutionary history of flowering plants (12).However, before applying molecular clocks to date evolutionary events, approximate constancy of evolutionary rate over time needs to be established (13) by using procedures for assessing rate homogeneity over taxa (14,15) or over lineages (16).Here we estimate the time when extant seed plants shared their last common ancestorby using molecular clocks derived from nucleotide sequences of the chloroplast gene coding for the large subunit of ribulose-1,5-bispho...