Dynamical analysis of the global elevational diversity gradient in passerine birds reveals a prominent role for highlands as species pumps

Low elevation regions harbor the majority of the world’s species diversity compared to high elevation areas. This global elevational diversity gradient, suggests that lowland species have had more time to diversify, or that net diversification rates have been higher in the lowlands (either due to higher ecological limits or intrinsically higher diversification rates). However, highlands seem to be cradles of diversity as they contain many young endemics, suggesting that their rates of speciation are exceptionally fast. Here, we use a phylogenetic diversification model that accounts for the dispersal of species between different elevations to examine the evolutionary dynamics of the elevational diversity gradient in passerine birds, a group that has radiated globally to occupy almost all elevations and latitudes. We find strong support for a model where passerines diversify at the same rate in the highlands and the lowlands but where the rate of dispersal from high to low elevations is more than twice as fast as in the reverse direction. This suggests that while there is no consistent trend in diversification across elevations, highland regions act as species pumps because the diversity they generate migrates into the lowlands, thus setting up the observed gradient in passerine diversity. This species pump is particularly strong in the tropics, where the inferred rate of speciation is 1.4 times faster than in the temperate zone. We conclude that despite their lower diversity, highland regions are disproportionally important for maintaining diversity in the adjacent lowlands. The extinction of species in the tropical highlands due to rapid climate change this century could thus have major and long-lasting impacts on global passerine diversity.