Late Oligocene fossil acorns and nuts of Quercus section Cyclobalanopsis from the Nanning Basin,Guangxi,South China

Quercus is the largest genus within the Fagaceae and has a rich fossil record.Most of the fossil material is attributed to the subgenus Quercus based on leaves,pollen or rarely acorns and nuts.Fossil records of Q.section Cyclobalanopsis characterized by ring-cupped acorns are relatively few and especially those described based on nuts are scant.In this study,we described four new species of Quercus section Cyclobalanopsis based on mummified acorns and nuts:Q.paleodisciformis X.Y.Liu et J.H.Jin sp.nov.,Q.paleohui X.Y.Liu et J.H.Jin sp.nov.,Q.nanningensis X.Y.Liu et J.H.Jin sp.nov.and Q.yongningensis X.Y.Liu et J.H.Jin sp.nov.These species closely resemble the extant species Q.disciformis,Q.hui,Q.kerrii,and Q.dinghuensis.The occurrence of Q.section Cyclobalanopsis in the Oligocene stratum of Guangxi,South China,suggests that the section has diversified within its extant distribution center since the Oligocene.By combining records from other areas,we propose that the section first appeared in the middle Eocene of East Asia(Sino-Japan),has diversified in situ with a few elements scattering into West Asia and southern Europe since the Oligocene and Pliocene,respectively,and finally became restricted in East Asia since the Pleistocene.This indicates that the section originated and diversified in East Asia,before spreading into West Asia no later than the Oligocene and into southern Europe by the Pliocene.Subsequently it disappeared from South Europe and West Asia due to the appearance of the(summer dry)Mediterranean climate and widespread cooling during the Pleistocene.

Climatic envelope of evergreen sclerophyllous oaks and their present distribution in the eastern Himalaya and Hengduan Mountains

Evergreen sclerophyllous oaks （the E.S. oaks, Quercus section Heterobalanus） are the dominant species of the local ecosystem in the eastern Himalaya and the Hengduan Mountains, southwest China. In this study, we document the climatic envelope of the seven E.S. oak species and examine the relationships between climate and their distribution. This was done using a principal components analysis （PCA） and multiple regression analysis （MRA） of nine climatic indices. The main climatic envelope of the E.S. oaks were： mean temperature of the warmest month （MTW）= 12.0-19.5℃, warmth index （WI） = 33.2-88.9℃ month, annual biotemperature （BT）=-6.9- -0.3 ℃, coldness index （CI）=-30.4- -10.1 ℃ month, mean temperature of the coldest month （MTC）=-3.7-3.0℃ and annual precipitation （AP）=701-897 mm at the lower limits; and MTW=8.3-16.1℃, WI=15.7-59.1℃ month, BT=3.6-8.9℃, CI=-55.4-19.3℃ month, MTC=8.3-16.1 ℃ and AP=610-811 mm at the upper limits. The climatic range of the E.S. oaks is wide and includes two climatic zones, the cool-temperature zone and the subpolar zone. The PCA and MRA results suggest that the thermal climate plays a major role and precipitation plays a secondary role in controlling the large-scale distribution of the E.S. oaks, except Quercus monimotricha. In thermal regimes, BT and/or MTW are most important for both lower and upper limits of the E.S. oaks. Furthermore, our results indicate that the upper distribution limits of the E.S. oaks are less determined by low temperatures and their duration （CI） than by other factors.

Late Eocene sclerophyllous oak from Markam Basin, Tibet, and its biogeographic implications

Sclerophyllous evergreen broad-leaved forests,mainly made up of sclerophyllous oak,Quercus section Heterobalanus(Øerst.)Menitsky,Fagaceae,represent the most typical forest type in the Hengduan Mountains.Their distribution pattern is closely related to the growth and formation of the Qinghai-Tibetan Plateau(QTP).The oldest fossil record of Quercus sect.Heterobalanus so far discovered is from the middle Miocene of the Gazhacun Formation in Namling County,southern Tibet.However,our recent discovery of leaf fossils from the upper Eocene of Lawula Formation in Markam Basin,southeastern Tibet,illustrates that their origin is nearly 20 Myr older than previously assumed.By integrating the results from geometric morphometrics,geographical range expansion,and ecological niche shifts of this section in what is now the QTP and the Hengduan Mountains,we infer that the leaves of Quercus sect.Heterobalanus were already adapted to cool and dry conditions in some local regions no later than in the late Eocene.Then,with the growth of the QTP and late Cenozoic global cooling,the expansion of cooler and drier habitats benefited the spread and development of this section and their leaves exhibited morphological stasis through stabilizing selection.Based on published fossil records and recent discoveries,we argue that Quercus sect.Heterobalanus appeared in the subtropical evergreen and deciduous broad-leaved mixed forests of the southeastern margin of what is now the QTP no later than in the late Eocene.Some taxa spread westwards along the Gangdese Mountains and later the Himalaya,and others spread eastwards and southeastwards,gradually becoming a dominant group of species in the Hengduan Mountains.This dispersal route is contrary to the previous“northwards hypothesis”of this section,and further supports the hypothesis of an East Asian origin for Quercus section Ilex Loudon.