title: Five major shifts of diversification through the long evolutionary history of Magnoliidae (angiosperms).
abstract: BACKGROUND
  With 10,000 species, Magnoliidae are the largest clade of flowering plants outside monocots and eudicots. Despite an ancient and rich fossil history, the tempo and mode of diversification of Magnoliidae remain poorly known. Using a molecular data set of 12 markers and 220 species (representing >75% of genera in Magnoliidae) and six robust, internal fossil age constraints, we estimate divergence times and significant shifts of diversification across the clade. In addition, we test the sensitivity of magnoliid divergence times to the choice of relaxed clock model and various maximum age constraints for the angiosperms.
  
  
  RESULTS
  Compared with previous work, our study tends to push back in time the age of the crown node of Magnoliidae (178.78-126.82 million years, Myr), and of the four orders, Canellales (143.18-125.90 Myr), Piperales (158.11-88.15 Myr), Laurales (165.62-112.05 Myr), and Magnoliales (164.09-114.75 Myr). Although families vary in crown ages, Magnoliidae appear to have diversified into most extant families by the end of the Cretaceous. The strongly imbalanced distribution of extant diversity within Magnoliidae appears to be best explained by models of diversification with 6 to 13 shifts in net diversification rates. Significant increases are inferred within Piperaceae and Annonaceae, while the low species richness of Calycanthaceae, Degeneriaceae, and Himantandraceae appears to be the result of decreases in both speciation and extinction rates.
  
  
  CONCLUSIONS
  This study provides a new time scale for the evolutionary history of an important, but underexplored, part of the tree of angiosperms. The ages of the main clades of Magnoliidae (above the family level) are older than previously thought, and in several lineages, there were significant increases and decreases in net diversification rates. This study is a new robust framework for future investigations of trait evolution and of factors influencing diversification in this group as well as angiosperms as a whole.
authors.affiliation: Laboratoire Ecologie, Systématique, Evolution, Université Paris-Sud, CNRS UMR 8079, 91405, Orsay, France. massoni.julien@gmail.com.
authors.forename: Julien
authors.initials: J
authors.lastname: Massoni
authors.affiliation: Institut de Recherche pour le Développement (IRD), UMR-DIADE, 911, avenue Agropolis, BP 64501, Cedex 5, F-34394, Montpellier, France. thomas.couvreur@ird.fr.|Département des Sciences Biologiques, Université de Yaoundé I, Ecole Normale Supérieure, Laboratoire de Botanique systématique et d'Ecologie, B.P. 047, Yaoundé, Cameroon. thomas.couvreur@ird.fr.
authors.forename: Thomas L P
authors.initials: TL
authors.lastname: Couvreur
authors.affiliation: Laboratoire Ecologie, Systématique, Evolution, Université Paris-Sud, CNRS UMR 8079, 91405, Orsay, France. herve.sauquet@u-psud.fr.
authors.forename: Hervé
authors.initials: H
authors.lastname: Sauquet
mesh_terms: D001499:Bayes Theorem; D005075:Biological Evolution; D005580:Fossils; D016013:Likelihood Functions; D019684:Magnoliopsida / Q000145:classification* / Q000235:genetics*; D010802:Phylogeny