Dated molecular phylogenies are the basis for understanding species diversity and for linking changes in rates of diversification with historical events such as restructuring in developmental pathways, genome doubling, or dispersal onto a new continent. Valid fossil calibration points are essential to the accurate estimation of divergence dates, but for many groups of flowering plants fossil evidence is unavailable or limited. Arabidopsis thaliana, the primary genetic model in plant biology and the first plant to have its entire genome sequenced, belongs to one such group, the plant family Brassicaceae. Thus, the timing of A. thaliana evolution and the history of its genome have been controversial. We bring previously overlooked fossil evidence to bear on these questions and find the split between A. thaliana and Arabidopsis lyrata occurred about 13 Mya, and that the split between Arabidopsis and the Brassica complex (broccoli, cabbage, canola) occurred about 43 Mya. These estimates, which are two-to threefold older than previous estimates, indicate that gene, genomic, and developmental evolution occurred much more slowly than previously hypothesized and that Arabidopsis evolved during a period of warming rather than of cooling. We detected a 2-to 10-fold shift in species diversification rates on the branch uniting Brassicaceae with its sister families. The timing of this shift suggests a possible impact of the Cretaceous-Paleogene mass extinction on their radiation and that Brassicales codiversified with pierid butterflies that specialize on mustard-oil-producing plants.he most important genetic model in plant biology is Arabidopsis thaliana. It is the first plant to have its entire genome sequenced, and it serves as a key comparison point with other eukaryotic genomes. A. thaliana is diploid and has a small genome distributed on just five chromosomes, considerations in its choice as a model (1). The age of the Arabidopsis crown group (CG), previously estimated at 5.8-3 Mya (2, 3), and of splits within Brassicaceae have been used to understand the pace of evolution in genes affecting self-incompatibility (4, 5), the rate of change in signal transduction and gene expression (6, 7), the persistence of shared chromosomal rearrangements in A. thaliana and Brassica oleracea (8), the tempo of evolution of miRNA sequences (9), the evolution of pierid butterflies specializing in plants that produce mustard oils (10), and the ages of wholegenome duplication (WGD) events giving rise to gene pairs in Arabidopsis (11). As genomes of additional Brassicaceae (e.g., Capsella rubella) and other Brassicales (e.g., Carica papaya) (12) are sequenced, the importance of robust estimates of divergence dates relating these genomes to one another and to the geological record increases substantially.The accuracy of divergence times inferred from sequence data depends on valid, verifiable fossils to calibrate phylogenetic trees. Previous dates for the origin of Arabidopsis relied on the report of fossil pollen assigned to the genus Rori...

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Flowering plants represent the most significant branch in the tree of land plants, with respect to the number of extant species, their impact on the shaping of modern ecosystems and their economic importance. However, unlike so many persistent phylogenetic problems that have yielded to insights from DNA sequence data, the mystery surrounding the origin of angiosperms has deepened with the advent and advance of molecular systematics. Strong statistical support for competing hypotheses and recent novel trees from molecular data suggest that the accuracy of current molecular trees requires further testing. Analyses of phytochrome amino acids using a duplicate gene-rooting approach yield trees that unite cycads and angiosperms in a clade that is sister to a clade in which Gingko and Cupressophyta are successive sister taxa to gnetophytes plus Pinaceae. Application of a cycads þ angiosperms backbone constraint in analyses of a morphological dataset yields better resolved trees than do analyses in which extant gymnosperms are forced to be monophyletic. The results have implications both for our assessment of uncertainty in trees from sequence data and for our use of molecular constraints as a way to integrate insights from morphological and molecular evidence.