The abiotic and biotic drivers of rapid diversification in Andean bellflowers (Campanulaceae).

The tropical Andes of South America, the world's richest biodiversity hotspot, are home to many rapid radiations. While geological, climatic, and ecological processes collectively explain such radiations, their relative contributions are seldom examined within a single clade. We explore the contribution of these factors by applying a series of diversification models that incorporate mountain building, climate change, and trait evolution to the first dated phylogeny of Andean bellflowers (Campanulaceae: Lobelioideae). Our framework is novel for its direct incorporation of geological data on Andean uplift into a macroevolutionary model. We show that speciation and extinction are differentially influenced by abiotic factors: speciation rates rose concurrently with Andean elevation, while extinction rates decreased during global cooling. Pollination syndrome and fruit type, both biotic traits known to facilitate mutualisms, played an additional role in driving diversification. These abiotic and biotic factors resulted in one of the fastest radiations reported to date: the centropogonids, whose 550 species arose in the last 5 million yr. Our study represents a significant advance in our understanding of plant evolution in Andean cloud forests. It further highlights the power of combining phylogenetic and Earth science models to explore the interplay of geology, climate, and ecology in generating the world's biodiversity.

Coloration of the Chilean Bellflower, Nolana paradoxa, interpreted with a scattering and absorbing layer stack model.

MAIN CONCLUSION: An absorbing-layer-stack model allows quantitative analysis of the light flux in flowers and the resulting reflectance spectra. It provides insight in how plants can optimize their flower coloration for attracting pollinators. The coloration of flowers is due to the combined effect of pigments and light-scattering structures. To interpret flower coloration, we applied an optical model that considers a flower as a stack of layers, where each layer can be treated with the Kubelka-Munk theory for diffusely scattering and absorbing media. We applied our model to the flowers of the Chilean Bellflower, Nolana paradoxa, which have distinctly different-colored adaxial and abaxial sides. We found that the flowers have a pigmented, strongly scattering upper layer, in combination with an unpigmented, moderately reflecting lower layer. The model allowed quantitative interpretation of the reflectance and transmittance spectra measured with an integrating sphere. The absorbance spectrum of the pigment measured with a microspectrophotometer confirmed the spectrum derived by modeling. We discuss how different pigment localizations yield different reflectance spectra. The absorbing layer stack model aids in understanding the various constraints and options for plants to tune their coloration.