The elevational gradient in Andean bird species richness at the local scale: a foothill peak and a high-elevation plateau

A monotonic decline in species richness with increasing elevation has often been considered a general pattern, but recent evidence suggests that the dominant pattern is hump-shaped with maximum richness occurring at some mid-elevation point. To analyse the relationship between species richness and elevation at a local scale we surveyed birds from lowlands to timberline in the Bolivian Andes. We divided the transect into 12 elevational belts of 250 m and standardized species richness in each belt with both individual- and sample-based rarefaction and richness estimation. The empirical data were then correlated to four explanatory variables: 1) area per elevational belt, 2) elevation (also representing ecosystem productivity), 3) a mid-domain effect (MDE) null model of geometrically constrained empirical range sizes, and 4) a hump-shaped model derived empirically for South American birds representing the regional species pool hypothesis. Local species richness peaked at ca 1000 m elevation, declined sharply to ca 1750 m, and then remained roughly constant. Elevation was the best single predictor, accounting for 78 - 85% of the variance in the empirical data. A multiple regression model with elevation, area, and MDE explained 85 - 90% of the variance. Monte Carlo simulations showed that the richness peak at 1000 m is the result of an overlap of two distinct avifaunas (lowland and highland) and that the correlation to MDE in the multiple regression was likely spurious. We recommend complementing correlation analyses involving MDE predictions with an examination of the distribution of range midpoints. The steep decline at mid-elevations was mainly due to a rapid loss of lowland species. The high-elevation plateau is striking and unexpected, but has also been found previously. It cannot be explained at present and exemplifies that despite several decades of research elevational gradients are still not well understood.