Paliurus (Paliureae,Rhamnaceae) from the Miocene of East China and its Macrofossil-based Phylogenetic and Phytogeographical History

The extant genus Paliurus Miller is divided into P. ramosissimus and P. spina‐christi groups and is diagnostically characterized by distinctive orbicular‐winged fruits with a trilocular (or often bilocular) ovary and basally triveined leaves with a subentire to serrate margin. Here, structurally preserved winged fruits and associated compressed leaves of Paliurus (Paliureae, Rhamnaceae) are reported from the Middle‐Upper Miocene Shengxian Formation of Tiantai, Zhejiang, eastern China. The fruits can be identified as Paliurus microcarpa Xiang‐Chuan Li sp. nov., the diagnostic characters of external morphology and anatomy of which differ from those of both extant and fossil generic members; the leaves can be designated as Paliurus sp. The species based on winged fruits, P. microcarpa, might represent an ancient intermediate link exclusively known across the P. ramosissimus and P. spina‐christi groups, showing a closer affinity to the P. ramosissimus group based on the resemblant smaller fruits with a narrower wing, and to the P. spina‐christi group based on the similar wing structure and texture, respectively. Morphological analyses of the present and previous fossil fruit records around the world indicate that diversification of Paliurus had begun in northern transpacific regions by the Late Eocene. The macrofossil records, i.e., fruits and leaves of Paliurus globally when projected on paleoenvironmental maps in view of changing continental positions through time (from Eocene to Miocene) display that the past distribution of the genus was much wider than the present and belongs to a tropical and warm temperate realm of the Northern Hemisphere. Therefore, the modern restricted disjunction in tropical and warm temperate regions of East Asia and South Europe (even eastwards West Asia) might represent glacial refugia for some of the generic ancient lineages to survive severe cold at least since the terminal Pliocene.