The phylogeography of Amazonia revisited: new evidence from riodinid butterflies.

A fully resolved cladogram for 19 species in the Charis cleonus group of riodinid butterflies, which have closely parapatric ranges throughout the Amazon basin, is used to derive an area cladogram for the region. This represents the first comprehensive species-level analysis using insects and results in a hypothesis of Amazonian area relationships that is the most detailed to date. The Charis area cladogram is interpreted as supporting an historical vicariant split between the Guianas and the remainder of the Amazon and then between the upper and lower Amazon. The latter two clades can be further divided into the six most widely recognized areas of endemism and even smaller endemic centers within these, some of which, especially along the Madeira and lower Amazon Rivers, have never been previously hypothesized for butterflies. The overall pattern of historical interrelationships indicated is Guiana + ((Rondĵnia + (Pará + Belém)) + (Imeri + (Napo + Inambari))). The area relationships for riodinid butterflies show substantial congruence with those presented from the literature for amphibians, reptiles, birds, primates, rodents, and marsupials, suggesting a common vicariant history for these organisms. A summary area cladogram generated by combining area cladograms for all the aforementioned groups of organisms indicated the pattern of historical interrelationships to be (Guiana + (Rondĵnia + (Pará + Belém))) + (Imeri + (Napo + Inambari)). Charis cleonus group species distributions are noticeably larger around the upland periphery of Amazonia and smaller in the central and lower regions. A significant positive correlation between the proportion of range area above 100 m and total range size for each species is used to suggest that past sea-level rises may explain smaller range sizes in low-lying regions and that riverine barriers have been important in shaping the current distribution of C. cleonus group species.