What controls the distribution of the Japanese endemic hemlock, Tsuga diversifolia? Footprint of climate in the glacial period on current habitat occupancy

Plant distributions are thought to be controlled by climate at large scales, and by non-climatic factors including soil conditions, topography and biotic interactions at smaller scales. However, not all plant distributions are explained by the current environment. Lags between current plant distributions and suitable environment for them are suggested to exist, which is often called empty habitat. To identify the existence and cause of lags between current climate and the distribution of Tsuga diversifolia, climatic conditions for the species distribution were clarified and potential habitats under current and the last glacial maximum (LGM; 21 ka) climates have been projected. The relationships between T. diversifolia distribution and climatic variables were explored using a classification tree model and a generalized additive model based on high-resolution (ca. 1 km) climatic data and a nationwide distribution database. The models were highly accurate. We revealed that T. diversifolia requires high summer precipitation even in humid Japanese environments. Areas with cool and wet summers were classified as potential habitat. Empty habitat for the focal species was identified in Hokkaido. Meanwhile, no potential habitat was projected in Hokkaido under the LGM. Additional experiments that varied temperature and summer precipitation during the LGM showed that the potential habitat was projected in Hokkaido irrespective of temperature decrease if summer precipitation increased nearly equal to the current climate. These results suggest that T. diversifolia vanished from Hokkaido, where fossil evidence indicated its occurrence until the late Neogene, during the glacial periods of the Pleistocene because of increased summer dryness.