Tropical plant species are expected to have high heat tolerance reflecting phenotypic adjustments to warm regions or their evolutionary adaptation history. However, tropical highland specialists adapted to the colder temperatures found in the highlands, where short and prostrated vegetation decouples plants from ambient conditions, could exhibit different upper thermal limits than those of their lowland counterparts. Here we evaluated leaf heat tolerance of 21 tropical alpine paramo species to determine: 1) whether species with restricted distribution (i.e., highland specialists) have lower heat tolerance and are more vulnerable to warming than species with widespread distribution; 2) whether different growth forms have different heat tolerance; and 3) whether species height (i.e., microhabitat) influences its heat tolerance. We quantified heat tolerance by evaluating T50, which is the temperature that causes a reduction in 50% of initial Fv/Fm values and reflects an irreversible damage to the photosynthetic apparatus. Additionally, we estimated the thermal safety margins as the difference between T50 and the maximum leaf temperature registered for the species. All species presented high T50 values ranging between 45.4°C and 53.9°C, similar to those found for tropical lowland species. Heat tolerance was not correlated with species distributions or plant height, but showed a strong relationship with growth form, with rosettes having the highest heat tolerance. Thermal safety margins ranged from 12.1 to 31.0°C. High heat tolerance and broad thermal safety margins suggest low vulnerability of paramo species to warming as long as plants are capable of regulating the leaf temperature within this threshold. Whether paramo plants would be able to regulate leaf temperature if drought episodes become more frequent and transpirational cooling is compromised is the next question that needs to be answered.

Páramos, tropical alpine ecosystems, host one of the world’s most diverse alpine floras, account for the largest water reservoirs in the Andes, and some of the largest soil carbon pools worldwide. It is of global importance to understand the future of this extremely carbon-rich ecosystem in a warmer world and its role on global climate feedbacks. This study presents the result of the first in situ warming experiment in two Colombian páramos using Open-Top Chambers. We evaluated the response to warming of several ecosystem carbon balance-related processes, including decomposition, soil respiration, photosynthesis, plant productivity, and vegetation structure after 3 years of warming. We found that OTCs are an efficient warming method in the páramo, increasing mean air temperature by 1.7°C and mean daytime temperature by 3.4°C. The maximum air temperature differences between OTC and control was 23.1°C. Soil temperature increased only by 0.1°C. After 3 years of warming using 20 OTC (10 per páramo) in a randomized block design, we found no evidence that warming increased CO2 emissions from soil respiration, nor did it increase decomposition rate, photosynthesis or productivity in the two páramos studied. However, total C and N in the soil and vegetation structure are slowly changing as result of warming and changes are site dependent. In Sumapaz, shrubs, and graminoids cover increased in response to warming while in Matarredonda we observed an increase in lichen cover. Whether this change in vegetation might influence the carbon sequestration potential of the páramo needs to be further evaluated. Our results suggest that páramos ecosystems can resist an increase in temperature with no significant alteration of ecosystem carbon balance related processes in the short term. However, the long-term effect of warming could depend on the vegetation changes and how these changes alter the microbial soil composition and soil processes. The differential response among páramos suggest that the response to warming could be highly dependent on the initial conditions and therefore we urgently need more warming experiments in páramos to understand how specific site characteristics will affect their response to warming and their role in global climate feedbacks.

The protocol is based on Krause (2010) with some modifications proposed by the Feeley Lab in 2016. This protocol is for finding heat tolerance or the critical thermal maximum temperature of plant leaves. Plants can undergo heat damage that can be repaired as long as the critical value is not exceed. This value is called critical thermal maximum (CT max) or heat tolerance (T 50) and is the highest temperature a leaf can take before and irreversible damage occurs, and when exceeded leads to necrosis. The way we measure this is using chlorophyll fluorescence, more specifically, the maximum efficiency of PSII value or F v /F m , which optimal values are around 0.83 in non stressed leaves. F v /F m can be obtained with a fluorometer or with an Irga, in both cases using dark adaptation clips. Plant collection is recommended to be done on the day of the procedure or, as long as leaves are kept well hydrated, the material can be even measured a day after the collection. The total time required for the measurement to be compleated is 2 days (including a 24h wait). Apart from the fluorometer, the equipment needed includes a water heating device (stable water temperature is imperative), such as a water bath or if possible a Sous Vide (recommended to us by Ken Feeley; works great!).