Coexistence of savanna and rainforest on the ice-age Sunda Shelf revealed by pollen records from southern South China Sea

Exposure of the presently submerged continental shelf of Southeast Asia during low sea-level stands of the Quaternary is thought to be deeply involved in the biogeographic evolution of this biodiversity hotspot. Inland proxy records are mounting to argue grassland expansions over Southeast Asia during glacial periods, lending supports to the existence of a transequatorial “savanna corridor” as hypothesized. However, pollen records retrieved from marine sediments have indicated widespread rainforests on the exposed northern Sunda Shelf. To reconcile these seemingly conflicting observations, here we present new, multisite marine pollen records from the southern South China Sea focusing on the Last Glacial Maximum (LGM) time interval. It is found that the sites in front of the Sunda Shelf palaeo-rivers contain substantially more Poaceae pollen, in particular a smaller size fraction, than the site offshore North Borneo. Among the sites, abundant lowland tree pollen with similar major taxa is also observed. The co-occurrence of grass and rainforest pollen suggests a longitudinal climate gradient with increasing aridity toward the interior of glacial Sundaland. Combining our results with various palynological evidence from marine sediments seems to indicate that the vegetation on the exposed Sunda Shelf might not be fully recognized by previous studies. This is likely due to an admixture of pollen from other source regions, particularly Borneo where the rainforest was retained during glacial periods. Therefore, the extant vegetation records, from both terrestrial and marine archives, support the hypothesis of a “savanna corridor” over the LGM sundaland.