The legacy of biogeographic history on the composition and structure of Andean forests.

ABSTRACT

The biogeographic origin of species may help to explain differences in average tree height and aboveground biomass (AGB) of tropical mountain forests. After the Andean uplift, small-statured trees should have been among the initial colonizers of the highlands (new cold environment) from the lowland tropics, since these species are pre-adapted to cold conditions with narrow vessels that are relatively resistant to freezing. If the descendants of these small-statured clades continue to dominate tropical highland forests, there will be a high co-occurrence of close relatives at high elevations. In other words, this scenario predicts a systematic decline in tree size, AGB, and phylogenetic diversity with elevation. In contrast, the colonization of Andean forests by some large-statured clades that originated in temperate regions may modify this expectation and promote a mixing of tropical and temperate clades, thereby increasing the phylogenetic diversity in tropical highland forests. This latter scenario predicts an increase or no change of tree size, AGB, and phylogenetic diversity with elevation. We assessed how the historical immigration of large-statured temperate-affiliated tree lineages adapted to cold conditions may have influenced the composition and structure of Andean forests. Specifically, we used 92 0.25-ha forest inventory plots distributed in the tropical Andes Mountains of Colombia to assess the relationship between the phylogenetic diversity and AGB along elevational gradients. We classified tree species as being either "tropical affiliated" or "temperate affiliated" and estimated their independent contribution to forest AGB. We used structural equation modeling to separate the direct and indirect effect of elevation on AGB. We found a hump-shaped relationship of phylogenetic diversity, AGB, and tree size with elevation. The high phylogenetic diversity found between 1,800-2,200 m above sea level (asl) was due to the mixing of highland floras containing many temperate-affiliated species, and lowland floras containing mostly tropical-affiliated species. The high AGB in highland forests, which contrasted with the expected decline of AGB with elevation, was likely due to the significant contribution of temperate-affiliated species. Our findings highlight the lasting importance of biogeographic history on the composition and structure of Andean mountain forests.