title: Zygomorphy evolved from disymmetry in Fumarioideae (Papaveraceae, Ranunculales): new evidence from an expanded molecular phylogenetic framework.
abstract: OBJECTIVE
  Fumarioideae (20 genera, 593 species) is a clade of Papaveraceae (Ranunculales) characterized by flowers that are either disymmetric (i.e. two perpendicular planes of bilateral symmetry) or zygomorphic (i.e. one plane of bilateral symmetry). In contrast, the other subfamily of Papaveraceae, Papaveroideae (23 genera, 230 species), has actinomorphic flowers (i.e. more than two planes of symmetry). Understanding of the evolution of floral symmetry in this clade has so far been limited by the lack of a reliable phylogenetic framework. Pteridophyllum (one species) shares similarities with Fumarioideae but has actinomorphic flowers, and the relationships among Pteridophyllum, Papaveroideae and Fumarioideae have remained unclear. This study reassesses the evolution of floral symmetry in Papaveraceae based on new molecular phylogenetic analyses of the family.
  
  
  METHODS
  Maximum likelihood, Bayesian and maximum parsimony phylogenetic analyses of Papaveraceae were conducted using six plastid markers and one nuclear marker, sampling Pteridophyllum, 18 (90 %) genera and 73 species of Fumarioideae, 11 (48 %) genera and 11 species of Papaveroideae, and a wide selection of outgroup taxa. Floral characters recorded from the literature were then optimized onto phylogenetic trees to reconstruct ancestral states using parsimony, maximum likelihood and reversible-jump Bayesian approaches.
  
  
  RESULTS
  Pteridophyllum is not nested in Fumarioideae. Fumarioideae are monophyletic and Hypecoum (18 species) is the sister group of the remaining genera. Relationships within the core Fumarioideae are well resolved and supported. Dactylicapnos and all zygomorphic genera form a well-supported clade nested among disymmetric taxa.
  
  
  CONCLUSIONS
  Disymmetry of the corolla is a synapomorphy of Fumarioideae and is strongly correlated with changes in the androecium and differentiation of middle and inner tepal shape (basal spurs on middle tepals). Zygomorphy subsequently evolved from disymmetry either once (with a reversal in Dactylicapnos) or twice (Capnoides, other zygomorphic Fumarioideae) and appears to be correlated with the loss of one nectar spur.
authors: Université Paris-Sud, Laboratoire Écologie, Systématique, Évolution, CNRS UMR 8079, 91405 Orsay, France and CNRS, UMR 0320/UMR 8120 Génétique Quantitative et Evolution - Le Moulon, INRA/Université Paris-Sud/CNRS/AgroParisTech, Ferme du Moulon, 91190 Gif-sur-Yvette, France herve.sauquet@u-psud.fr.
authors: Hervé
authors: H
authors: Sauquet
authors: Université Paris-Sud, Laboratoire Écologie, Systématique, Évolution, CNRS UMR 8079, 91405 Orsay, France and CNRS, UMR 0320/UMR 8120 Génétique Quantitative et Evolution - Le Moulon, INRA/Université Paris-Sud/CNRS/AgroParisTech, Ferme du Moulon, 91190 Gif-sur-Yvette, France.
authors: Laetitia
authors: L
authors: Carrive
authors: Université Paris-Sud, Laboratoire Écologie, Systématique, Évolution, CNRS UMR 8079, 91405 Orsay, France and CNRS, UMR 0320/UMR 8120 Génétique Quantitative et Evolution - Le Moulon, INRA/Université Paris-Sud/CNRS/AgroParisTech, Ferme du Moulon, 91190 Gif-sur-Yvette, France.
authors: Noëlie
authors: N
authors: Poullain
authors: Université Paris-Sud, Laboratoire Écologie, Systématique, Évolution, CNRS UMR 8079, 91405 Orsay, France and CNRS, UMR 0320/UMR 8120 Génétique Quantitative et Evolution - Le Moulon, INRA/Université Paris-Sud/CNRS/AgroParisTech, Ferme du Moulon, 91190 Gif-sur-Yvette, France.
authors: Julie
authors: J
authors: Sannier
authors: Université Paris-Sud, Laboratoire Écologie, Systématique, Évolution, CNRS UMR 8079, 91405 Orsay, France and CNRS, UMR 0320/UMR 8120 Génétique Quantitative et Evolution - Le Moulon, INRA/Université Paris-Sud/CNRS/AgroParisTech, Ferme du Moulon, 91190 Gif-sur-Yvette, France.
authors: Catherine
authors: C
authors: Damerval
authors: Université Paris-Sud, Laboratoire Écologie, Systématique, Évolution, CNRS UMR 8079, 91405 Orsay, France and CNRS, UMR 0320/UMR 8120 Génétique Quantitative et Evolution - Le Moulon, INRA/Université Paris-Sud/CNRS/AgroParisTech, Ferme du Moulon, 91190 Gif-sur-Yvette, France.
authors: Sophie
authors: S
authors: Nadot
mesh_terms: D005075:Biological Evolution; D035264:Flowers / Q000033:anatomy & histology*; D016013:Likelihood Functions; D028101:Papaveraceae / Q000033:anatomy & histology*; D010802:Phylogeny
keywords: Character evolution; Fumarioideae; Papaveraceae; Papaveroideae; Pteridophyllum; Ranunculales; disymmetry; floral evolution; floral symmetry; nectar spur; phylogeny; reversible-jump Markov chain Monte Carlo; zygomorphy