Involucre fossils of Carpinus, a northern temperate element, from the Miocene of China and the evolution of its species diversity in East Asia

East Asia has long been recognized as a major center for temperate woody plants diversity. Although several theories have been proposed to explain how the diversity of these temperate elements accumulated in the region, the specific process remains unclear. Here we describe six species of Carpinus, a typical northern hemisphere temperate woody plant, from the early Miocene of the Maguan Basin, southwestern China, southern East Asia. This constitutes the southernmost, and the earliest occurrence that shows a high species diversity of the genus. Together with other Carpinus fossil records from East Asia, we show that the genus had achieved a high diversity in East Asia at least by the middle Miocene. Of the six species here described, three have become extinct, indicating that the genus has experienced apparent species loss during its evolutionary history in East Asia. In contrast, the remaining three species closely resemble extant species, raising the possibility that these species may have persisted in East Asia at least since the early Miocene. These findings indicate that the accumulation of species diversity of Carpinus in East Asia is a complex process involving extinction, persistence, and possible subsequent speciation.