Altitudinal range-size distribution of breeding birds and environmental factors for the determination of species richness: An empirical test of altitudinal Rapoport's rule and non-directional rescue effect on a local scale.

Range-size distributions are important for understanding species richness patterns and led to the development of the controversial Rapoport's rule and Rapoport-rescue effect. This study aimed to understand the relationship between species richness and range-size distribution in relation to environmental factors. The present study tested the following: (1) altitudinal Rapoport's rule, and a subsequent test on climatic and ambient energy hypotheses, (2) non-directional rescue effect, and a subsequent test on effect of environmental factors associated with the distribution of narrowest to widest-range species. Altitudinal species range-size distribution increased with increasing altitude and showed a negative relationship with climatic variables. These results support the altitudinal Rapoport's rule and climatic hypothesis; however, they do not fully support the ambient energy hypothesis. Results from testing the non-directional rescue effect showed that the inflow intensity of species from both directions (high and low elevations) affected species richness. And we found that the species with intermediate range-size, rather than narrowest or widest range-size were the main cause of a mid-peak of species richness and the non-directional rescue effect. Additionally, the richness of species with intermediate range-size was highly related to minimum temperature, habitat heterogeneity, or primary productivity. Although altitudinal range-size distribution results were similar to the phenomenon of altitudinal Rapoport's rule, the mid-peak pattern of species richness could not be explained by the underlying mechanism of Rapoport's-rescue effect; however, the non-directional rescue effect could explain a mid-peak pattern of species richness along altitudinal gradient.