Fossil fruits of Firmiana and Tilia from the middle Miocene of South Korea and the efficacy of the Bering land bridge for the migration of mesothermal plants

Determining whether the high-latitude Bering land bridge(BLB)was ecologically suitable for the migration of mesothermal plants is significant for Holarctic phytogeographic inferences.Paleobotanical studies provide a critical source of data on the latitudinal positions of different plant lineages at different times,permitting assessment of the efficacy of the BLB for migration.Here we report exceptionally preserved fossils of Firmiana and Tilia endochrysea from the middle Miocene of South Korea.This represents a new reliable record of Firmiana and the first discovery of the T.endochrysea lineage in the fossil record of Asia.The occurrence of these fossils in South Korea indicates that the two lineages had a distribution that extended much farther north during the middle Miocene,but they were still geographically remote from the BLB.In light of the broader fossil record of Asia,our study shows that,in the middle Miocene,some mesothermal plants apparently inhabited the territory adjacent to the BLB and thus they were possibly capable of utilizing the BLB as a migratory corridor.Some other mesothermal plants,such as Firmiana and the T.endochrysea lineages,however,are restricted to more southern regions relative to the BLB based on current fossil evidence.These lineages may have been ecologically unable to traverse the BLB,which raises questions about the efficacy of the BLB as a universal exchange route for mesothermal plants between Asia and North America during the middle Miocene.