SMILAX (SMILACACEAE) FROM THE MIOCENE OF WESTERN EURASIA WITH CARIBBEAN BIOGEOGRAPHIC AFFINITIES

Premise of the study: Recent molecular studies provide a phylogenetic framework and some dated nodes for the monocot genus Smilax. The Caribbean Havanensis group of Smilax is part of a well-supported New World clade with a few disjunct taxa in the Old World. Although the fossil record of the genus is rich, it has been difficult to assign fossil taxa to extant groups based on their preserved morphological characters. Methods: Leaf fossils from Europe and Asia Minor were studied comparatively and put into a phylogenetic and biogeographic context using a molecular phylogeny of the genus. Key results: Fossils from the early Miocene of Anatolia represent a new species of Smilax with systematic affinities with the Havanensis group. The leaf type encountered in the fossil species is exclusively found in species of the Havanensis group among all modern Smilax. Scattered fossils of this type from the Miocene of Greece and Austria, previously referred to Quercus (Fagaceae), Ilex (Aquifoliaceae), and Mahonia (Berberidaceae) also belong to the new species. Conclusions: The new Smilax provides first fossil evidence of the Havanensis group and proves that this group had a western Eurasian distribution during the Miocene. The age of the fossils is in good agreement with the (molecular-based) purported split between the Havanensis and Hispida groups within Smilax. The Miocene Smilax provides evidence that all four subclades within the New World clade had a disjunct intercontinental distribution during parts of the Neogene involving trans-Atlantic crossings (via floating islands or the North Atlantic land bridge) and the Beringia land bridge.