Nutrient inputs into ecosystems of the tropical mountain rainforest region are projected to further increase in the next decades. To investigate whether important ecosystem services such as nutrient cycling and matter turnover in native forests and pasture ecosystems show different patterns of response, two nutrient addition experiments have been established: NUMEX in the forest and FERPAST at the pasture. Both ecosystems already responded 1.5 years after the start of nutrient application (N, P, NP, Ca). Interestingly, most nutrients remained in the respective systems. While the pasture grass was co-limited by N and P, most tree species responded to P addition. Soil microbial biomass in the forest litter layer increased after NP fertilization pointing to nutrient co-limitation. In pasture soils, microorganisms were neither limited by N nor P. The results support the hypothesis that multiple and temporally variable nutrient limitations can coexist in tropical ecosystems.
The tropical montane forests of the E Andean cordillera in Ecuador receive episodic Sahara-dust inputs particularly increasing Ca deposition. We added CaCl2 to isolate the effect of Ca deposition by Sahara dust to tropical montane forest from the simultaneously occurring pH effect. We examined components of the Ca cycle at four control plots and four plots with added Ca (2 x 5kg ha(-1) Ca annually as CaCl2) in a random arrangement. Between August 2007 and December 2009 (four applications of Ca), we determined Ca concentrations and fluxes in litter leachate, mineral soil solution (0.15 and 0.30 m depths), throughfall, and fine litterfall and Al concentrations and speciation in soil solutions. After 1 y of Ca addition, we assessed fine-root biomass, leaf area, and tree growth. Only < 3% of the applied Ca leached below the acid organic layer (pH 3.5-4.8). The added CaCl2 did not change electrical conductivity in the root zone after 2 y. In the second year of fertilization, Ca retention in the canopy of the Ca treatment tended to decrease relative to the control. After 2 y, 21% of the applied Ca was recycled to soil with throughfall and litterfall. One year after the first Ca addition, fine-root biomass had decreased significantly. Decreasing fine-root biomass might be attributed to a direct or an indirect beneficial effect of Ca on the soil decomposer community. Because of almost complete association of Al with dissolved organic matter and high free Ca2+ : Al3+ activity ratios in solution of all plots, Al toxicity was unlikely. We conclude that the added Ca was retained in the system and had beneficial effects on some plants.
Tropical regions are facing increasing atmospheric inputs of nutrients, which will have unknown consequences for the structure and functioning of these systems. Here, we show that Neotropical montane rainforests respond rapidly to moderate additions of N (50 kg ha(-1) yr(-1)) and P (10 kg ha(-1) yr(-1)). Monitoring of nutrient fluxes demonstrated that the majority of added nutrients remained in the system, in either soil or vegetation. N and P additions led to not only an increase in foliar N and P concentrations, but also altered soil microbial biomass, standing fine root biomass, stem growth, and litterfall. The different effects suggest that trees are primarily limited by P, whereas some processes-notably aboveground productivity-are limited by both N and P. Highly variable and partly contrasting responses of different tree species suggest marked changes in species composition and diversity of these forests by nutrient inputs in the long term. The unexpectedly fast response of the ecosystem to moderate nutrient additions suggests high vulnerability of tropical montane forests to the expected increase in nutrient inputs.
Elevated tropical nitrogen deposition interacts with global warming via changes in forest soil trace gas emissions
Atmospheric nitrogen (N) and phosphorus (P) depositions are expected to increase in the tropics as a consequence of increasing human activities in the next decades. In the literature, it is frequently assumed that tropical montane forests are N-limited, while tropical lowland forests are P-limited. In a low-level N and P addition experiment, we determined the short-term response of N and P cycles in a north Andean montane forest on Palaeozoic shists and metasandstones at an elevation of 2100 m a.s.l. to increased N and P inputs. We evaluated experimental N. P and N + P additions (50 kg ha(-1) yr(-1) of N, 10 kg ha(-1) yr(-1) of P and 50 kg + 10 kg ha(-1) yr(-1) of N and P. respectively) and an untreated control in a fourfold replicated randomized block design. We collected litter leachate, mineral soil solution (0.15 and 0.30m depths), throughfall and litterfall before the treatment began (August 2007) until 16 months after the first nutrient application (April 2009). Less than 10 and 1% of the applied N and P. respectively, leached below the organic layer which contained almost all roots and no significant leaching losses of N and P occurred to below 0.15 m mineral soil depth. Deposited N and P from the atmosphere in dry and wet form were retained in the canopy of the control treatment using a canopy budget model. Nitrogen and P retention by the canopy were reduced and N and P fluxes in throughfall and litterfall increased in their respective treatments. The increase in N and P fluxes in throughfall after fertilization was equivalent to 2.5% of the applied N and 2% of the applied P. The fluxes of N and Pin litterfall were up to 15% and 3%, respectively, higher in the N and N + P than in the control treatments. We conclude that the expected elevated N and P deposition in the tropics will be retained in the ecosystem, at least in the short term and hence, N and P concentrations in stream water will not increase. Our results suggest that in the studied tropical montane forest ecosystem on Palaeozoic bedrock, N and P are co-limiting the growth of organisms in the canopy and organic layer. (C) 2010 Elsevier B.V. All rights reserved.
The drivers of spatial throughfall heterogeneity are still not fully understood. At an undisturbed forest site in the Ecuadorian Andes with ca. 2600 mm of annual rainfall we determined the accuracy of throughfall measurements by comparing Hellmann-type funnel gauges with troughs. At the same undisturbed and a managed, selectively-logged forest site we determined spatial variability of throughfall, temporal stability of spatial variability and the controls of spatial throughfall variability using a 4-year dataset in weekly resolution. There were no systematic differences between the collected volumes of funnel gauges and troughs. Based on the statistical distribution of annual throughfall volumes, a high number of 27 funnel-type rainfall collectors were required in the undisturbed forest and 20 in the managed forest to estimate throughfall with an error of 10% and a confidence interval of 95%. Spatial throughfall variability in the studied forests was high, markedly stable during 4 years and similar in six selected rain events suggesting that a stable canopy structure controlled throughfall variability. After mathematically eliminating the canopy influence, no meteorological variable had a significant effect on throughfall variability. We conclude that the high spatial variability of throughfall in the study forest, mainly controlled by a long-term stable canopy structure, contributes to the creation of different ecological niches which are a prerequisite for the enormous biological diversity of the north Andean forests.
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Tropical nitrogen (N) deposition is projected to increase substantially within the coming decades. Increases in soil emissions of the climate-relevant trace gases NO and N2O are expected, but few studies address this possibility. We used N addition experiments to achieve N-enriched conditions in contrasting montane and lowland forests and assessed changes in the timing and magnitude of soil N-oxide emissions. We evaluated transitory effects, which occurred immediately after N addition, and long-term effects measured at least 6 weeks after N addition. In the montane forest where stem growth was N limited, the first-time N additions caused rapid increases in soil N-oxide emissions. During the first 2 years of N addition, annual N-oxide emissions were five times (transitory effect) and two times (long-term effect) larger than controls. This contradicts the current assumption that N-limited tropical montane forests will respond to N additions with only small and delayed increases in soil N-oxide emissions. We attribute this fast and large response of soil N-oxide emissions to the presence of an organic layer (a characteristic feature of this forest type) in which nitrification increased substantially following N addition. In the lowland forest where stem growth was neither N nor phosphorus (P) limited, the first-time N additions caused only gradual and minimal increases in soil N-oxide emissions. These first N additions were completed at the beginning of the wet season, and low soil water content may have limited nitrification. In contrast, the 9- and 10-year N-addition plots displayed instantaneous and large soil N-oxide emissions. Annual N-oxide emissions under chronic N addition were seven times (transitory effect) and four times (long-term effect) larger than controls. Seasonal changes in soil water content also caused seasonal changes in soil N-oxide emissions from the 9- and 10-year N-addition plots. This suggests that climate change scenarios, where rainfall quantity and seasonality change, will alter the relative importance of soil NO and N2O emissions from tropical forests exposed to elevated N deposition.