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We aim to assess net primary productivity (NPP) and carbon cycling in Andean tropical alpine grasslands (puna) and compare it with NPP of tropical montane cloud forests. We ask the following questions: (1) how do NPP and soil respiration of grasslands vary over the seasonal cycle? (2) how do burning and grazing affect puna productivity? (3) if the montane forest expands into the puna, what will be the resulting change in productivity? The study sites are located at the South-eastern Peruvian Andes; one grassland site and the forest sites are in Wayqecha biological station, and another grassland site in Manu National Park. At each grassland site, we selected a burnt and an unburnt area, installed unfenced and fenced transects in each area, and monitored above-ground productivity (NPPAG), below-ground productivity (NPPBG) and soil respiration (Rs) for 2 yr. In the forest, we monitored NPPAG, NPPBG and Rs for 2–4 yr. Grassland NPP varied between 4.6 ± 0.25 (disturbed areas) to 15.3 ± 0.9 Mg C ha-1 yr-1 (undisturbed areas) and cloud forest NPP was between 7.05 ± 0.39 and 8.0 ± 0.47 Mg C ha-1 yr-1, while soil carbon stocks were in the range of 126 ± 22 to 285 ± 31Mg C ha-1. There were no significant differences on NPP between the puna and forest sites. The most undisturbed site had significantly higher NPP than other grassland sites, but no differences were found when relating grazing and fire at other sites. There were lower residence times of above-ground biomass compared to below-ground biomass. There was a strong seasonal signal on grassland NPPAG and NPPBG, with a shift on allocation at the beginning of the austral summer. High elevation tropical grasslands can be as productive as adjacent cloud forests, but have very different carbon cycling and retention properties than cloud forests. S Online supplementary data available from stacks.iop.org/ERL/9/115011/mmedia Keywords: tropical alpine wetlands, above-ground productivity, below-ground productivity, fire, grazing, disturbances, puna

Changes to the carbon cycle in tropical forests could affect global climate, but predicting such changes has been previously limited by lack of field-based data. Here we show seasonal cycles of the complete carbon cycle for 14, 1ha intensive carbon cycling plots which we separate into three regions: humid lowland, highlands, and dry lowlands. Our data highlight three trends: (1) there is differing seasonality of total net primary productivity (NPP) with the highlands and dry lowlands peaking in the dry season and the humid lowland sites peaking in the wet season, (2) seasonal reductions in wood NPP are not driven by reductions in total NPP but by carbon during the dry season being preferentially allocated toward either roots or canopy NPP, and (3) there is a temporal decoupling between total photosynthesis and total carbon usage (plant carbon expenditure). This decoupling indicates the presence of nonstructural carbohydrates which may allow growth and carbon to be allocated when it is most ecologically beneficial rather than when it is most environmentally available.

Solar irradiance and precipitation are the most likely drivers of the seasonal variation of net primary productivity (NPP) in tropical forests. Since their roles remain poorly understood, we use litter traps, dendrometer bands and census data collected from one hectare permanent plots to quantify the seasonality of above-groundNPPcomponents and weather parameters in 13 sites distributed along a 2800-m altitudinal gradient ranging from lowland Amazonia to the high Andes. We combine canopy leaf area index and litterfall data to describe the seasonality of canopy production. We hypothesize that solar irradiance is the primary driver of canopy phenology in wetter sites, whereas precipitation drives phenology in drier systems. The seasonal rhythm of canopyNPPcomponents is in synchrony with solar irradiance at all altitudes. Leaf litterfall peaks in the late dry season, both in lowland (averaging 0.54 ± 0.08 Mg C ha y−1, n = 5) and montane forests (averaging 0.29 ± 0.04 Mg C ha y−1, n = 8). Peaks in above-ground coarse woodyNPPappears to be triggered by the onset of rainfall in seasonal lowland rain forests (averaging 0.26 ± 0.04 Mg C ha y−1, n = 5, in November), but not in montane cloud forests.

The key role of tropical forest belowground carbon stocks and fluxes is well recognised as one of the main components of the terrestrial ecosystem carbon cycle. This study presents the first detailed investigation of spatial and temporal patterns of fine root stocks and fluxes in tropical forests along an elevational gradient, ranging from the Peruvian Andes (3020 m) to lowland Amazonia (194 m), with mean annual temperatures of 11.8 degrees C to 26.4 degrees C and annual rainfall values of 1900 to 1560 mm yr(-1), respectively. Specifically, we analyse abiotic parameters controlling fine root dynamics, fine root growth characteristics, and seasonality of net primary productivity along the elevation gradient. Root and soil carbon stocks were measured by means of soil cores, and fine root productivity was recorded using rhizotron chambers and ingrowth cores. We find that mean annual fine root below ground net primary productivity in the montane forests (0-30 cm depth) ranged between 4.27 +/- 0.56 Mg C ha(-1) yr(-1) (1855 m) and 1.72 +/- 0.87 Mg C ha(-1) yr(-1) (3020 m). These values include a correction for finest roots (< 0.6 mm diameter), which we suspect are under sampled, resulting in an underestimation of fine roots by up to 31% in current ingrowth core counting methods. We investigate the spatial and seasonal variation of fine root dynamics using soil depth profiles and an analysis of seasonal amplitude along the elevation gradient. We report a stronger seasonality of NPPFineRoot within the cloud immersion zone, most likely synchronised to seasonality of solar radiation. Finally, we provide the first insights into root growth characteristics along a tropical elevation transect: fine root area and fine root length increase significantly in the montane cloud forest. These insights into belowground carbon dynamics of tropical lowland and montane forests have significant implications for our understanding of the global tropical forest carbon cycle. (Less)

The net primary productivity, carbon (C) stocks and turnover rates (i.e. C dynamics) of tropical forests are an important aspect of the global C cycle. These variables have been investigated in lowland tropical forests, but they have rarely been studied in tropical montane forests (TMFs). This study examines spatial patterns of above- and belowground C dynamics along a transect ranging from lowland Amazonia to the high Andes in SE Peru. Fine root biomass values increased from 1.50 Mg C ha−1 at 194 m to 4.95 ± 0.62 Mg C ha−1 at 3020 m, reaching a maximum of 6.83 ± 1.13 Mg C ha−1 at the 2020 m elevation site. Aboveground biomass values decreased from 123.50 Mg C ha−1 at 194 m to 47.03 Mg C ha−1 at 3020 m. Mean annual belowground productivity was highest in the most fertile lowland plots (7.40 ± 1.00 Mg C ha−1 yr−1) and ranged between 3.43 ± 0.73 and 1.48 ± 0.40 Mg C ha−1 yr−1 in the premontane and montane plots. Mean annual aboveground productivity was estimated to vary between 9.50 ± 1.08 Mg C ha−1 yr−1 (210 m) and 2.59 ± 0.40 Mg C ha−1 yr−1 (2020 m), with consistently lower values observed in the cloud immersion zone of the montane forest. Fine root C residence time increased from 0.31 years in lowland Amazonia to 3.78 ± 0.81 years at 3020 m and stem C residence time remained constant along the elevational transect, with a mean of 54 ± 4 years. The ratio of fine root biomass to stem biomass increased significantly with increasing elevation, whereas the allocation of net primary productivity above- and belowground remained approximately constant at all elevations. Although net primary productivity declined in the TMF, the partitioning of productivity between the ecosystem subcomponents remained the same in lowland, premontane and montane forests.

Severe drought in a tropical forest ecosystem suppresses photosynthetic carbon uptake and plant maintenance respiration, but growth is maintained, suggesting that, overall, less carbon is available for tree tissue maintenance and defence, which may cause the subsequent observed increase in tree mortality. The underlying mechanisms that determine the response of tropical forest ecosystems to drought remain poorly understood. Based on observations from a network of intensively measured forest plots in the Amazon basin, this study shows that severe drought suppresses photosynthetic carbon uptake and plant maintenance respiration. Plant growth is maintained however, suggesting that less carbon is available for tree tissue maintenance and defence — which may explain the observed increase in tree mortality that follows a drought. In 2005 and 2010 the Amazon basin experienced two strong droughts1, driven by shifts in the tropical hydrological regime2 possibly associated with global climate change3, as predicted by some global models3. Tree mortality increased after the 2005 drought4, and regional atmospheric inversion modelling showed basin-wide decreases in CO2 uptake in 2010 compared with 2011 (ref. 5). But the response of tropical forest carbon cycling to these droughts is not fully understood and there has been no detailed multi-site investigation in situ. Here we use several years of data from a network of thirteen 1-ha forest plots spread throughout South America, where each component of net primary production (NPP), autotrophic respiration and heterotrophic respiration is measured separately, to develop a better mechanistic understanding of the impact of the 2010 drought on the Amazon forest. We find that total NPP remained constant throughout the drought. However, towards the end of the drought, autotrophic respiration, especially in roots and stems, declined significantly compared with measurements in 2009 made in the absence of drought, with extended decreases in autotrophic respiration in the three driest plots. In the year after the drought, total NPP remained constant but the allocation of carbon shifted towards canopy NPP and away from fine-root NPP. Both leaf-level and plot-level measurements indicate that severe drought suppresses photosynthesis. Scaling these measurements to the entire Amazon basin with rainfall data, we estimate that drought suppressed Amazon-wide photosynthesis in 2010 by 0.38 petagrams of carbon (0.23–0.53 petagrams of carbon). Overall, we find that during this drought, instead of reducing total NPP, trees prioritized growth by reducing autotrophic respiration that was unrelated to growth. This suggests that trees decrease investment in tissue maintenance and defence, in line with eco-evolutionary theories that trees are competitively disadvantaged in the absence of growth6. We propose that weakened maintenance and defence investment may, in turn, cause the increase in post-drought tree mortality observed at our plots.