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...
The rosid clade (70,000 species) contains more than one-fourth of all angiosperm species and includes most lineages of extant temperate and tropical forest trees. Despite progress in elucidating relationships within the angiosperms, rosids remain the largest poorly resolved major clade; deep relationships within the rosids are particularly enigmatic. Based on parsimony and maximum likelihood (ML) analyses of separate and combined 12-gene (10 plastid genes, 2 nuclear; >18,000 bp) and plastid inverted repeat (IR; 24 genes and intervening spacers; >25,000 bp) datasets for >100 rosid species, we provide a greatly improved understanding of rosid phylogeny. Vitaceae are sister to all other rosids, which in turn form 2 large clades, each with a ML bootstrap value of 100%: (i) eurosids I (Fabidae) include the nitrogen-fixing clade, Celastrales, Huaceae, Zygophyllales, Malpighiales, and Oxalidales; and (ii) eurosids II (Malvidae) include Tapisciaceae, Brassicales, Malvales, Sapindales, Geraniales, Myrtales, Crossosomatales, and Picramniaceae. The rosid clade diversified rapidly into these major lineages, possibly over a period of <15 million years, and perhaps in as little as 4 to 5 million years. The timing of the inferred rapid radiation of rosids [108 to 91 million years ago (Mya) and 107-83 Mya for Fabidae and Malvidae, respectively] corresponds with the rapid rise of angiosperm-dominated forests and the concomitant diversification of other clades that inhabit these forests, including amphibians, ants, placental mammals, and ferns. community assembly ͉ divergence time estimates ͉ phylogeny ͉ rapid radiation G reat progress has been made in elucidating deep-level angiosperm relationships during the past decade. The eudicot clade, with Ϸ75% of all angiosperm species, comprises several major subclades: rosids, asterids, Saxifragales, Santalales, and Caryophyllales (1-3). Investigations have converged on the branching pattern of the basalmost angiosperms, revealing that Amborellaceae, Nymphaeales [in the sense of APG II (3) and including Hydatellaceae (4)], and Austrobaileyales are successive sisters to all other extant angiosperms (reviewed in ref.2). Analyses of complete plastid genome sequences have resolved other problematic deep-level relationships, suggesting that Chloranthaceae and magnoliids are sister to a clade of monocots and eudicots plus Ceratophyllaceae (5, 6). Likewise, progress has been made in clarifying relationships within the large monocot (7) and asterid (8) clades.Despite these successes, the rosids stand out as the largest and least-resolved major clade of angiosperms; basal nodes within the clade have consistently received low internal support (1, 2, 9, 10). The rosid clade comprises Ϸ70,000 species and 140 families (2, 11). Containing more than a quarter of total angiosperm and Ϸ39% of eudicot species diversity, the rosid clade is broader in circumscription than the traditional Rosidae or Rosanae (e.g., 12; reviewed in ref.2). The oldest fossil flowers conforming to the rosids are from the late S...
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