Biotic diversification in the Guayana Highlands: a proposal
Until recently, the high degree of diversity and endemism of the Guayana Highlands was explained within the frame of the refuge theory. Although this hypothesis is unsupported by recent palaeoecological evidence, no new diversification model has been proposed. This paper is a proposal based on the latest palynological findings that indicate a downward biotic migration of c. 1100 m altitude during glacials, and the subsequent interglacial upward shift, in response to colder and warmer climates, respectively. Therefore, during glacials, biotic mixing is expected in the lowlands, thus promoting sympatric speciation, hybridization and polyploidy. At the mountaintops, unknown cold-adapted taxa and páramo-like(?) communities are expected to have occurred, and vicariance prevailed. In the interglacials, many taxa have had the opportunity for ascending to the mountains again, allowing genetic interchange among their slopes and summits, while others would have been adapted to lowlands. The interglacial highland communities, where vicariance still predominated, experienced some extinction owing to habitat loss by upland displacement. According to this model, the successive alternation of glacials and interglacials resulted in a net increase of diversity and endemism, favoured by the complex topography and habitat heterogeneity, which allowed high niche diversification. This model has some similarities with the Andean and Amazon modes of diversification, but the special topographical characteristics of the Guayana region made it different in other fundamental aspects. The Guayana Highlands would have acted as a ‘biodiversity pump’ for the surrounding inner and coastal lowlands, due to the repeated speciation and further spreading events, as a response to climate. Several working hypotheses are suggested in relation to the proposed model. The use of coordinated international multiproxy projects combining palaeoecology and genetic analysis of modern taxa is strongly encouraged for exploring these ideas.