title: Corolla morphology influences diversification rates in bifid toadflaxes (Linaria sect. Versicolores).
abstract: OBJECTIVE
  The role of flower specialization in plant speciation and evolution remains controversial. In this study the evolution of flower traits restricting access to pollinators was analysed in the bifid toadflaxes (Linaria sect. Versicolores), a monophyletic group of ~30 species and subspecies with highly specialized corollas.
  
  
  METHODS
  A time-calibrated phylogeny based on both nuclear and plastid DNA sequences was obtained using a coalescent-based method, and flower morphology was characterized by means of morphometric analyses. Directional trends in flower shape evolution and trait-dependent diversification rates were jointly analysed using recently developed methods, and morphological shifts were reconstructed along the phylogeny. Pollinator surveys were conducted for a representative sample of species.
  
  
  RESULTS
  A restrictive character state (narrow corolla tube) was reconstructed in the most recent common ancestor of Linaria sect. Versicolores. After its early loss in the most species-rich clade, this character state has been convergently reacquired in multiple lineages of this clade in recent times, yet it seems to have exerted a negative influence on diversification rates. Comparative analyses and pollinator surveys suggest that the narrow- and broad-tubed flowers are evolutionary optima representing divergent strategies of pollen placement on nectar-feeding insects.
  
  
  CONCLUSIONS
  The results confirm that different forms of floral specialization can lead to dissimilar evolutionary success in terms of diversification. It is additionally suggested that opposing individual-level and species-level selection pressures may have driven the evolution of pollinator-restrictive traits in bifid toadflaxes.
authors.affiliation: Departamento de Biodiversidad y Conservación, Real Jardín Botánico (RJB-CSIC), Plaza de Murillo 2, 28014 Madrid, Spain.
authors.forename: Mario
authors.initials: M
authors.lastname: Fernández-Mazuecos
authors.forename: José Luis
authors.initials: JL
authors.lastname: Blanco-Pastor
authors.forename: José M
authors.initials: JM
authors.lastname: Gómez
authors.forename: Pablo
authors.initials: P
authors.lastname: Vargas
mesh_terms: D000818:Animals; D005075:Biological Evolution; D035264:Flowers / Q000033:anatomy & histology*; D032221:Linaria / Q000033:anatomy & histology / Q000235:genetics*; D057048:Plant Nectar / Q000502:physiology; D054817:Pollination
chemical_list: D057048:Plant Nectar
keywords: Convergence; Linaria sect. Versicolores; flower specialization; flower tube; nectar spur; pollination; reversal; speciation; species selection; toadflax; trait-dependent diversification