reflecting widespread extinction during climate cooling at the Oligocene/Miocene boundary
[∼ 23 million years ago (Ma)]. Recent biogeographic studies have revealed many instances
of long-distance dispersal in gymnosperms as well as in angiosperms. Acting together,
extinction and long-distance dispersal are likely to erase historical biogeographic signals.
Notwithstanding this problem, we show that phylogenetic relationships in the gymnosperm …