Niche-dependent forest and savanna fragmentation in Tropical South America during the Last Glacial Maximum.

The refugia hypothesis, often used to explain Amazonia's high biodiversity, initially received ample support but has garnered increasing criticism over time. Palynological, phylogenetic, and vegetation model reconstruction studies have been invoked to support the opposing arguments of extensive fragmentation versus a stable Amazonian Forest during Pleistocene glacial maxima. Here, we test the past existence of forest fragments and savanna connectivity by bias-correcting vegetation distributions from a Dynamic Vegetation Model (DVM) driven by paleoclimate simulations for South America during the Last Glacial Maximum (LGM). We find evidence for fragmented forests akin to refugia with extensive tropical humid forests to the west and forest islands in central/southern Amazonia. Drier ecosystems of Northern Llanos, Caatinga and Cerrado may have merged into continuous savanna/grasslands that dominated the continent. However, our reconstructions suggest taller, dense woodland/tropical savanna vegetation and areas of similar bioclimate connected disparate forest fragments across Amazonia. This ecotonal biome may have acted as a corridor for generalist forest and savanna species, creating connectivity that allows for range expansion during glacial periods. Simultaneously, it could have served as a barrier for specialists, inducing diversification through the formation of 'semi-refugia'.