The rise and fall of the Refugial Hypothesis of Amazonian speciation: a paleoecological perspective

The refugial hypothesis is treated as the definitive history of Amazonian forests in many texts. Surprisingly, this important theoretical framework has not been based on paleoecological data. Consequently, a model of Amazonian aridity during the northern hemispheric glaciation has been accepted uncritically. Ironically, the Refuge Hypothesis has not been tested by paleobotanical data. We present a revision of the concept of Neotropical Pleistocene Forest Refuges and test it in the light of paleocological studies derived from pollen analysis of Amazonian lake sediments deposited during the last 20,000 years. Our analysis is based primarily on paleoenvironmental data obtained from sites in Brazil and Ecuador. These data are contrasted with those that favor the hypothesis of fragmented tropical forests in a landscape dominated mainly by tropical savannas under an arid climate. The Ecuadorian data set strongly suggests a 5ºC cooling and presence of humid forests at the foot of the Andes, during the last Ice Age. The same climatic and vegetational scenario was found in the western Brazilian Amazon. On the other hand, somewhat drier conditions were observed in the central Amazon, but the landscape remained a forested landscape during the supposedly arid phases of the Late Quaternary. Data obtained from the Amazon Fan sediments containing pollen derived from extensive sections of the Amazon Basin, were fundamental to the conclusion that the predominance of savannas in this region is not supported by botanical data. Our revision of the assumptions derived from the Refuge Hypothesis indicates that it has succumbed to the test now permitted by a larger paleocological data set, which were not available during the golden age of this paradigm, when indirect evidence was considered satisfactory to support it.