Tussocks facilitate their neighbours mainly by ameliorating extreme temperatures in tropical high mountains

Facilitation by tussocks is common in high-altitude tropical environments. It is thought that facilitation results from stress amelioration, but it is unclear which of the many stressors acting in these environments is ameliorated. We aimed at determining the relative importance of different stressors as drivers of facilitation by the tussock Festuca tolucensis in Mexico. We employed eight experimental treatments to manipulate five stressors in the field: minimum temperatures by using electric radiators that kept plants warm; maximum temperatures by means of reflective sand that precluded temperature build-up during the day; UV radiation by using screens opaque to UV; poor soil properties by comparing soils from beneath tussocks and from bare ground; and low water availability by adding vermiculite to the soil. The performance (survival and growth) of Mexerion sarmentosum (a plant usually associated with Festuca) in these treatments was compared to that recorded under tussocks and in bare ground. Amelioration of extreme temperatures had the largest positive effects on Mexerion survival. UV radiation and increased soil humidity did not affect survival, although humidity increased growth rates. Nevertheless, tussocks reduced the growth of Mexerion, which is consistent with observations of competition between plants and soil microorganisms favoured by tussocks. Our results highlight the importance of the extreme daily fluctuations in temperature that characterise tropical mountains as fundamental drivers of their dynamics.