RESUMEN
Patterns of species diversity have been associated with changes in climate across latitude and elevation. However, the ecological and evolutionary mechanisms underlying these relationships are still actively debated. Here, we present a complementary view of the well-known tropical niche conservatism (TNC) hypothesis, termed the multiple zones of origin (MZO) hypothesis, to explore mechanisms underlying latitudinal and elevational gradients of phylogenetic diversity in tree communities. The TNC hypothesis posits that most lineages originate in warmer, wetter, and less seasonal environments in the tropics and rarely colonize colder, drier, and more seasonal environments outside of the tropical lowlands, leading to higher phylogenetic diversity at lower latitudes and elevations. In contrast, the MZO hypothesis posits that lineages also originate in temperate environments and readily colonize similar environments in the tropical highlands, leading to lower phylogenetic diversity at lower latitudes and elevations. We tested these phylogenetic predictions using a combination of computer simulations and empirical analyses of tree communities in 245 forest plots located in six countries across the tropical and subtropical Andes. We estimated the phylogenetic diversity for each plot and regressed it against elevation and latitude. Our simulated and empirical results provide strong support for the MZO hypothesis. Phylogenetic diversity among co-occurring tree species increased with both latitude and elevation, suggesting an important influence on the historical dispersal of lineages with temperate origins into the tropical highlands. The mixing of different floras was likely favored by the formation of climatically suitable corridors for plant migration due to the Andean uplift. Accounting for the evolutionary history of plant communities helps to advance our knowledge of the drivers of tree community assembly along complex climatic gradients, and thus their likely responses to modern anthropogenic climate change.
RESUMEN
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.