Earliest orchid macrofossils: Early Miocene Dendrobium and Earina (Orchidaceae: Epidendroideae) from New Zealand.

Fossil leaves of two Early Miocene orchids (Dendrobium and Earina) are reported from New Zealand. The distinctive, raised tetra- to cyclocytic stomatal subsidiary cells of Earina and characteristic papilla-like absorbing glands and "ringed" guard cells of Dendrobium support the placement of the fossils into these genera. These therefore represent the first Orchidaceae macrofossils with cuticular preservation, the oldest records for subfamily Epidendroideae, as well as the first New Zealand and southern hemisphere fossil records for Orchidaceae. These taxa belong in basal clades to the Vandeae/Cymbideae or Epidendreae (Earina) and the Australasian clade of Dendrobium sensu lato. This phylogenetic placement demonstrates expansion of epiphytic orchids into Zealandia by the mid-Cenozoic and an important role for southern continents in the diversification of Orchidaceae.