A Review on the Taxonomic, Evolutionary and Phytogeographic Studies of the Lotus Plant (Nelumbonaceae: Nelumbo)

Nelumbo Adans. （Nelumbonaceae） is an important member of the early-diverging eudicots. It contains two extant species： N. nucifera Gaertn. （the Sacred lotus）, distributed in Asia and Australia and N. lutea Willd. （the American lotus）, occurring in North America. This paper reviews the taxonomic, evolutionary and phytogeographic studies of the genus Nelumbo, and also raises scientific questions about it in further paleobotanic research. There are about 30 fossil species of Nelumbo established since the Early Cretaceous. Based on fossil studies, the ancestors of the extant N. nucifera and N. lutea are respectively considered to be N. protospeciosa from the Eocene to Miocene of Eurasia, and N. protolutea from the Eocene of North American. However, molecular systematic studies indicate that N. nucifera and N. lutea are probably split from a common ancestor during the Late Miocene to Early Pliocene, or even the Pleistocene, rather than separate relicts from extinct species on different continents. The characters of lotus stomatal development, seedling morphology as well as its flowering, pollination and fertilization in air reveal that it evolves from the land plants. Fossil data of Nelumbo indicates that the genus first occurs in mid-latitude area of Laurasia in the Early Cretaceous, then becomes widespread in North America and Eurasia and expands into Africa and South America during the Late Cretaceous; the genus probably colonizes the Indian Subcontinent from Asia during the Early Eocene after the collision of India and the Asian plates; the genus becomes extinct in Europe, but survives in Asia and North America during the Quaternary Ice Age, and later forms the present East Asia and North Australia-North America disjunctive distribution.