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Plant functional diversity in tropical Andean páramos

ABSTRACT Background: Tropical high mountains present extreme daily temperature variations, frequent high air evaporative demands and seasonal differences in soil water availability. Plants have adapted to these conditions through different avoidance-tolerance mechanisms. This review focuses on plant-growth forms and their adaptive strategies. Aims: This integrated review of páramo plant traits aims at contributing to understanding the functioning of plant-growth forms and their significance on ecosystem properties under environmental climate and land-use changes. Methods: Plant responses are presented along avoidance-tolerance gradients considering three main aspects: freezing resistance, water relations and gas exchange characteristics. Results from 45 herbaceous and 42 woody species along elevational gradients in the Venezuelan high Andes were analysed. Results: Leaf supercooling is the common avoidance response of woody plants to night-time freezing temperatures, while herbaceous plants tolerate frost. Trees and caulescent rosettes maintain more positive leaf water potentials under water deficit conditions compared to more tolerant herbaceous species. All plant growth-forms showed strong stomatal control under dry-season conditions. Conclusions: Páramo plant growth-forms may be separated according to an avoidance-tolerance gradient in response to water deficit and low temperature resistance. Woody growth-forms tend to avoid both freezing and water stress, while herbaceous forms tolerate frost and resist an unfavourable water status. Grasses and cushion plants are at the tolerant extreme of the gradient and coincide in that both reach the highest elevations in the páramo. Andean giant rosettes are freezing avoidant, particularly susceptible to water deficit and the most vulnerable, of all growth-forms, to changing environmental conditions.