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The timing of angiosperm origin and diversifi cation is crucial to understand the evolution of terrestrial ecosystems as we know them today. The fossil record and relaxed molecular clocks provide substantially different estimates of the time in which angiosperms started to diversify into lineages that originated living species. The roles of different methodological and systematic factors on ages estimated with relaxed clocks across land plants, with a focus on angiosperms, are here reviewed. Different re- laxed clocks and molecular data provided similar age estimates for nodes across land plants. Breaking the long branch subtending angiosperms typically did not result in substantially younger age estimates for angiosperms. Including or excluding fossil-derived calibrations on internal nodes had a pronounced effect on ages across the tree, and among examined factors, calibrations were the most infl uential in resulting ages. However, interactions between calibrations and genes were detected as different genes converged on similar ages when the same set of calibrations was applied. Estimated angiosperm ages were much older than angiosperm ear- liest fossils. Although a missing fossil history cannot be ruled out, this explanation alone is insuffi cient, considering the gradually increasing abundance, diversity and geographical distribution of angiosperms in the fossil record; the ordered progression of their morphological and functional diversifi cation; and the agreement in the sequence of appearance of lineages in the stratigraphic record and in molecular phylogenies. It is hypothesized that model misspecifi cation may play a role in the observed molecular clock/fossil discrepancy. Resumen: La edad del origen y diversifi cacion de las angiospermas son cruciales para entender la evolucion de los ecosistemas terrestres modernos. El registro fosil y los relojes moleculares relajados calculan edades substancialmente diferentes para el inicio de la diversifi cacion de las angiospermas. Se revisa el efecto de diferentes factores metodologicos y sistematicos en los estima- dos de relojes relajados para las plantas terrestres, con enfasis en las angiospermas. Diferentes metodos de reloj relajado y datos moleculares estimaron edades similares en todo el arbol fi logenetico. Romper la rama larga que subyace a las angiospermas no resulto en edades substancialmente mas jovenes. La inclusion o exclusion de calibraciones derivadas del registro fosil tuvo un efecto pronunciado en las edades estimadas, ademas de ser el factor mas relevante entre los que se evaluaron. Sin embargo, existen interacciones entre las calibraciones y los genes, pues diferentes genes convergieron en edades similares cuando las mismas cali- braciones fueron aplicadas. Las edades estimadas para las angiospermas son mucho mas viejas que sus primeros fosiles. Aunque la posibilidad de un registro fosil incompleto no puede excluirse, esta sola explicacion es insufi ciente, dado el aumento gradual en la abundancia y distribucion geografi ca de las angiospermas en el registro fosil; la progresion ordenada de su diversifi cacion morfologica y funcional; y la concordancia en la secuencia de aparicion de linajes en la secuencia estratigrafi ca y en las fi logenias moleculares. Se sugiere que la especifi cacion erronea del modelo evolutivo posiblemente juega un papel en la discrepancia entre los relojes relajados y el registro fosil. Palabras clave: calibracion, efecto del linaje, modelos mal especifi cados, plantas terrestres, registro fosil.

A Review of the Effect of Relaxed Clock Method, Long Branches, Genes, and Calibrations in the Estimation of Angiosperm Age