Environmental changes in biodiversity hotspot ecosystems of
South Ecuador: RESPonse and feedback effECTs (FOR2730)

Abstract:

Increased nutrient supply can have drastic effects on natural ecosystems, especially in naturally nutrient-poor ones such as most tropical rainforests. Many studies have focused on the reaction of trees to fertilization, but little is known about herbaceous plants. Ferns are a particularly common group in tropical forests, spanning all vegetation types and zones. Here, we assess how seven years of moderate addition of nitrogen (N), phosphorus (P), and N+P along an elevational gradient (1000–3000 m) have impacted richness and composition of fern and lycophyte assemblages in tropical montane rain forests growing on naturally nutrient deficient soils in the Ecuadorian Andes. We found that fertilization does not affect overall species richness, but that there were strong differences in species abundances (~60% of species), both negative and positive, that were apparently related to the systematic affiliations and ecological properties of the affected species. These diverse responses of ferns to fertilization provide insight into the sensitivity and complexity of the relationships of nutrient availability and community composition in tropical forests.

Abstract:

In the past two decades, the Amazon-exposed, tropical montane rain forests in south Ecuador experienced increasing deposition of reactive N mainly from Amazonian forest fires, episodic Ca and Mg inputs from Saharan dust, and a low but constant P deposition from unknown sources. To explore the response of this tropical, perhumid ecosystem to nutrient inputs, we established in 2007 a Nutrient Manipulation Experiment (NUMEX). Since 2008, we have applied 50 kg ha−1 year−1 of N as urea, 10 kg ha−1 year−1 of P as NaH2PO4·H2O, 50 kg ha−1 year−1 of N + 10 kg ha−1 year−1 of P and 10 kg ha−1 year−1 of Ca as CaCl2·H2O in a randomized block design at 2000 m a.s.l. in a natural forest of the south Ecuadorian Andes. Previous studies have shown that alkali and alkaline earth metals had beneficial effects on the functioning of N and P co-limited tropical forests occurring on acidic soils. Therefore, we determined the response of all major aqueous ecosystem fluxes of K, Ca, Mg and Na to nutrient amendments, to understand how increasing atmospheric deposition would affect their cycling in the future. Additions of N and P decreased K leaching from the organic layer and in the mineral soil, thus tightening K cycling. This suggests that increasing future N and P availability may result in K limitation in the long term. The leaching of Ca and Mg from the canopy increased in response to amendments of N and P and we observed an enhanced uptake of these nutrients also if Ca was amended alone. Although N was applied as urea, acidity of soil solutions and leaching of K, Ca, Mg and Na did not increase following separate N amendments. In spite of the acid soils and of its low cation-exchange competitivity, Na included in the P fertilizer was only partly leached from the organic layer. We suggest that it was probably required to cover an unmet Na demand of the soil fauna. Our results demonstrate the major role in the functioning of the tropical montane forests played by K, Ca and Mg as potential future growth-limiting elements and increasingly required nutrients in response to rising N and P availability, while they also support the importance of Na as a functional element in these ecosystems.

Abstract:

Anthropogenic atmospheric deposition can increase nutrient supply in the most remote ecosystems, potentially affecting soil biodiversity. Arbuscular mycorrhizal fungal (AMF) communities rapidly respond to simulated soil eutrophication in tropical forests. Yet the limited spatio-temporal extent of such manipulations, together with the often unrealistically high fertilization rates employed, impedes generalization of such responses. We sequenced mixed root AMF communities within a seven year-long fully factorial nitrogen (N) and phosphorus (P) addition experiment, replicated at three tropical montane forests in southern Ecuador with differing environmental characteristics. We hypothesized (i) strong shifts in community composition and species richness after long-term fertilization, (ii) site- and (iii) clade-specific responses to N versus P additions depending on local soil fertility and clade life history traits respectively. Fertilization consistently shifted AMF community composition across sites, but only reduced richness of Glomeraceae. Compositional changes were mainly driven by increases in P supply while richness reductions were observed only after combined N and P additions. We conclude that moderate increases of N and P exert a mild but consistent effect on tropical AMF communities. To predict the consequences of these shifts, current results need to be supplemented with experiments that characterize local species-specific AMF functionality.

Abstract:

The anthropogenic deposition of nitrogen (N) and phosphorus (P) into terrestrial ecosystems has an influence on the leaf morphology and leaf properties of vegetation as well as on interactions within an ecosystem. In this context, the increased availability of nutrients has an impact on herbivory. In addition to the availability of nutrients, sea level has also been shown to influence these parameters. The shown work deals with the quantification of herbivory under the influence of N- and P-fertilization and differences in certain leaf parameters (leaf area, specific leaf area, leaf toughness, leaf nitrogen content and leaf area loss) between three altitudinal levels (1000 m, 2000 m, 3000 m). The study area was located in a tropical mountain rainforest in Southern Ecuador. In a nutrient manipulation experiment (NUMEX) 48 plots within the study areas were fertilized with either N, P or NP twice a year. Per plot 50 leaves were collected, examined for various parameters and then subjected to a nutrient analysis. The aim of this study was to figure out to what extent sea level influences the leaf morphology, nitrogen content and the leaf area loss of the vegetation within the control areas and to what extent fertilization with nitrogen, phosphorus or both elements changes these leaf properties. Finally, it was investigated whether possible changes in feeding rates resulted from an increased leaf nitrogen content. Sea level had a major influence on leaf strength, which increased with increasing height gradients, and on the specific leaf area and the mean leaf area, which decreased with increasing sea level. Many of the values differed significantly between the altitudinal levels. This suggests that not only the soil becomes nutrient-poorer with rising sea level, but also that the mineralisation of the few available nutrients is slower than at lower altitudes. The climatic environmental conditions, which become more extreme as the sea level rises, also explain the changes in the leaf parameters mentioned. Fertilisation instead did not have an influence as strong as sea level on the studied parameters. There were hardly any significant differences between the plots with different fertilisation. Since tropical forests are considered nutrient-limited, the addition of nitrogen led to the formation of large and soft leaves, which was expected. The correlations between leaf nitrogen content and SLA or leaf strength were positive and significant. Leaf area loss did not correlate with nitrogen content, which was unexpected. This can be attributed to the fact that plants show highly species-specific reactions to nutrient availability, which manifest themselves in very different plastic changes in growth and defence mechanisms against herbivore species. In this work a strong influence of the sea level on the leaf parameters could be determined. An increasing influence of fertilization on the feeding rate of herbivorous species was not found.

Abstract:

The tropical Andes are one of the most biodiverse hotspots on earth. Though the nutrient-limited systems are affected by anthropogenic nutrient inputs due to industry and agriculture. The NuMEx (Nutrient Manipulation Experiment) project, which was set up in 2008 and is located in southern Ecuador and aims to find out about the consequences of nutrient pollution to herbivore and plant interaction. Therefore experimental plots at three different levels of elevation were fertilised with nitrogen and phosphorus. In this study, fresh leaf samples were collected in May 2018. Plant parameters like leaf area, specific leaf area, leaf toughness and nitrogen content were analysed. Herbivory was examined trough leaf area loss measurements. Changes in these parameters along an elevation gradient from 1000 to 3000 m were analysed. Moreover, the study focussed on the influence of the nutrient addition to the sensible tropical system. A significant influence of the elevation on most of the leaf morphology parameters was shown. Specific leaf area and leaf nitrogen content showed a significant decrease along the elevation gradient, while the leaves became tougher. The mean leaf area and the leaf area loss gained no significant results; however, the leaf area tended to be smaller within rising elevation. The leaf area loss was highest at 3000 m of elevation. All elevation outcomes, except for the leaf area loss, could be explained by the harsher climatic conditions at higher elevation levels and the relationship between nitrogen content and the leaf traits. Nutrient addition results on the leaf traits were less clear. Almost no significant influences could be measured. The leaf characteristics showed mostly a clear trend though, except for the leaf area. The leaf area partially increased (mostly under mixed nitrogen and phosphorus addition) and partially decreased (nitrogen treatment) due to the nutrient addition. Specific leaf area, leaf nitrogen content and leaf area loss mostly increased due to the nutrient addition, especially nitrogen and the mixed nitrogen and phosphorus sample contributed to the increase. Leaf toughness whereas decreased. The highest decrease was caused at the plots treated with the nitrogen and phosphorus mix. The results make clear that a high leaf nitrogen content correlates with soft leaves and a high specific leaf area. An analysis of the leaf area loss and the leaf nitrogen content yielded that also these traits are correlated.

Abstract:

Global changes in nutrient deposition rates are likely to have profound effects on plant communities, particularly in the nutrient-limited systems of the tropics. We studied the effects of increased nutrient availability on the seedlings of six tree species in montane forests of southern Ecuador in situ. After five years of continued N, P, or N+P addition, naturally grown seedlings of each of the two most common species at each elevation (1000, 2000, and 3000 m asl) were harvested for analyses of leaf morphology, nutrient content, herbivory, and tissue biomass allocation. Most species showed increased foliar N and P concentrations after addition of each respective element. Leaf tissue N:P ratios of >20 in the control plants of all species suggest that P is more growth-limiting in these forests than N. Leaf morphological responses to nutrient addition were species and nutrient specific, with some species (Hedyosmum purparescens, Graffenrieda emarginata) exhibiting increased specific leaf area (SLA), and others (Graffenrieda harlingii) increased leaf area ratios (LAR). Pouteria torta (1000 m) had lower SLA and LAR after P addition. Increased herbivory was only evident in G. emarginata (after N and N+P addition). Only the species from 3000 m asl modified biomass allocation after nutrient addition. In general, N and N+P addition more strongly affected the species studied at the upper elevations, whereas P addition had a similar range of effects on the species at all elevations. We conclude that the responses of the studied tropical montane forest tree seedlings to chronic N and P addition are highly species-specific and that successful adaptation to increased nutrient availability will depend on species-specific morphological and physiological plasticity.

Abstract:

Trait response-effects are critical to forecast community structure and biomass production in highly diverse tropical forests. Ecological theory and few observation studies indicate that trees with acquisitive functional traits would respond more strongly to higher resource availability than those with conservative traits. We assessed how long-term tree growth in experimental nutrient addition plots (N, P and N+P) varied as a function of morphological traits, tree size and species identity. We also evaluated how trait-based responses affected stand scale biomass production considering the community structure. We found that tree growth depended on interactions between functional traits and the type or combination of nutrients added. Common species with acquisitive functional traits responded more strongly to nutrient addition, mainly to N+P. Phosphorous enhanced the growth rates of species with acquisitive and conservative traits, had mostly positive effects on common species and neutral or negative effects in rare species. Moreover, trees receiving N+P grew faster irrespective of their initial size relative to control or to other treatment plots. Finally, species responses were highly idiosyncratic suggesting that community processes including competition and niche dimensionality may be altered under increased resource availability. We found no statistically significant effects of nutrient additions on aboveground biomass productivity because acquisitive species had a limited potential to increase their biomass, possibly due to their generally lower wood density. In contrast, P addition increased the growth rates of species characterized by more conservative resource strategies (with higher wood density) that were poorly represented in the plant community. We provide the first long-term experimental evidence that trait-based responses, community structure, and community processes modulate the effects of increased nutrient availability on biomass productivity in a tropical forest.

Abstract:

Deforestation and land use change have led to a strong reduction of tropical forest cover during the last decades. Climate change will amplify the pressure to the remaining refuges in the next years. In addition, tropical regions are facing increasing atmospheric inputs of nutrients, which will have unknown consequences for the structure and functioning of these systems, no matter if they are within protected areas or not. Even remote areas are expected to receive rising amounts of nutrients. The effects of higher rates of atmospheric nutrient deposition on the biological diversity and ecosystem functioning of tropical ecosystems are poorly understood and our knowledge of nutrient fluxes and nutrient limitation in tropical forest ecosystems is still limited. Yet, it will be of paramount importance to know the effects of increased nutrient availability to conserve these ecosystems with their biological and functional diversity. During the last years, research efforts have more and more focused on the understanding of the role of nutrients in tropical ecosystems and several coordinated projects have been established that study the effects of experimental nutrient addition. This Research Topic combines results from experiments and from observational studies with the aim to review and conclude on our current knowledge on the role of additional nutrients in ecosystems.

Abstract:

Ecosystem services, provided from tropical forests, are indispensable for human beings. Coherencies in the system, their networks, drivers and various underlying pathways are not completely understood yet. Elucidation on directions of key nutrients and changes in organisms delivers the opportunity to get an overview about these relations. Leaf functional traits are one important component to uncover those cascades and organizations. With their fast respond to environmental conditions, changes can be detected. Since, tropical regions suffer from increasing atmospheric inputs of nitrogen (N) and phosphorus (P), it is a need to investigate these progressions and predict future scenarios. The alarming fact, that these anthropogenic caused input have unknown consequences for the structure and functioning of tropical forests leads to a crucial study topic. Furthermore, questions should be answered if these increased inputs have a comparable influence on different elevations, based on identification the limiting nutrients. To forecast these impacts the Nutrient Manipulation Experiment (NUMEX) has been conducted since 2008. It gained to show, that Neotropical montane forests respond rapidly to moderate nutrient additions of N (50 kg ha-1 yr-1) and P (10 kg ha-1 yr-1). Within the present study, the eight most abundant tree species from the NUMEX sites were analyzed to their leaf functional traits from three elevations (altitude above sea level) in an Ecuadorian montane forest. The premontane forest in Bombuscaro (1,000 m) contains the species Clarisia racemosa and Pouteria torta. Further, in San Francisco (2,000 m), Alchornea lojaensis, Graffenrieda emarginata, Hieronyma fendleri, and Myrcia sp., were selected. In Cajanuma (3,000 m), the upper montane forest contains Hedyosmum purpurascens and Weinmannia loxensis. The fertilization effects were calculated, using a mixed effect model, including study year and treatment as fixed effects and block and individual as random effects. The results show, that foliar N decrease over the years, whereas, Foliar P conversely increases. Further, the relation of nitrogen and phosphorus (N:P) is decreasing during the study years. Upwards trends for ?15N are best replicated for the San Francisco site. With the leaf trait nutrient resorption efficiency (NuR) for N and P a decline over the study years is proceeding. In addition, leaf area (LA) and specific leaf area (SLA) tend to decrease. In conclusive processes over the study years might be the result of microclimatic events. The sites differ in that San Francisco and Cajanuma respond more similar in some leaf functional traits, compared to them from Bombuscaro. It can be deduced that changes in the analyzed leaf traits over the last eight years between the sites and species are not significantly different. Summarizing, results represent the emphasis on the considerable influence of nutrient addition with notably changing foliar nutrient and show that the predicted nutrient deposition will probably change the ecosystem dynamics sustained.

Abstract:

Tropical montane forests of the Andes belong to the hotspots of biodiversity. But these nutrient poor ecosystems are simultaneously threatened by increased element inputs of nitrogen and phosphor in nutrient cycles. NUMEX-Projekt simulates the expected nutrient inputs in the for-ests and quantifies the changes of this ecosystem. In line of this study litter samples were collected by littertraps during a working period from February till May 2016. Leaf litter produc-tion, parameters of leaf morphology (leaf area and specific leaf area) and litter nutrients of nitrogen and phosphor were determined. Additionally, the parameters of herbivory (leaf mass loss and leaf area loss) were calculated by measuring holes area. The variation of these pa-rameters along an altitudinal gradient from 1.000 m.a.s.l. to 3.000 m.a.s.l. was investigated. Besides the variation after nitrogen and/or phosphor addition was explored. Furthermore, the extend of nutrients, lost through herbivory, was identified for the stand level. Terminatory the influence of two soil parameters (C/N ratio and Presin) was discussed. There is a significant influence of altitude (p<0,05) on leaf morphology (leaf area: 1.000 m.a.s.l: 26,4±1,6 cm², 3.000 m.a.s.l: 7,4±1,0 cm², specific leaf area: 1.000 m.a.s.l: 99,7±4,1 cm²·g-1, 3.000 m.a.s.l: 56,1±6,6 cm²·g-1), leaf nutrients of nitrogen (1.000 m.a.s.l: 15,3±0,5 g·kg-1, 3.000 m.a.s.l: 6,6±0,7 g·kg-1) and phosphor (1.000 m.a.s.l: 0,54±0,03 g·kg-1, 3.000 m.a.s.l: 0,25±0,03 g·kg-1) and also on the parameters of herbivory: holes area (1.000 m.a.s.l: 1,8±0,2 cm², 3.000 m.a.s.l: 0,3±0,1 cm²) and leaf area loss (1.000 m.a.s.l: 6,8±0,4 %, 3.000 m.a.s.l: 3,6±0,6 %). Nutrient addition did not lead to distinct results according to the variation of measured param-eters. NP fertilization had a positive effect on leaf area and P fertilization a positive effect on specific leaf area. Leaf nutrients showed various answers. For this parameter, NP fertilization causes significantly raised nitrogen and phosphor concentrations in litter in each site. Leaf litter production in 1.000 m.a.s.l site significantly decreased through NP fertilization. Referring to herbivory only a little number of significant effects were detected. Holes area increased on 2.000 m.a.s.l site by NP addition while leaf area loss in 1.000 m.a.s.l site – as well as leaf area production – decreased. On stand level, there was no variation of leaf area loss. Soil parameters have a higher impact on measured parameters on stand level. Specific leaf area and leaf area loss correlate positively with soil C/N of upper mineral soil. Correlations of these parameters with plant available phosphor Presin is negative. Nutrient losses on stand level decrease with altitude. There was a negative effect of NP addition on nitrogen loss in 1.000 m.a.s.l site and a positive effect of NP addition on phosphor loss in all sites.

Abstract:

Anthropogenic activities have produced changes in natural ecosystems worldwide. In tropical regions in South America, industrialization of cities and forest clearance via burning are the main activities releasing pollutants into the atmosphere and inducing changes in nutrient deposition patterns and climate of primary forests. Nitrogen (N) and phosphorus (P) are considered the main limiting nutrients of plant growth since their availability is vital for net primary productivity. Therefore, any change in N or P availability in soils would likely alter important mechanisms of forests dynamics such as growth and survival. Changes in soil pH (acidity), nutrient cycles and altered nutrient stocks affect N and P availability and affect various physiological processes of trees. Although low levels of nutrient deposition have been reported for montane forests in southern Ecuador (ca. 5 kg ha–1 for N, 0.49 kg ha–1 for P), even these levels are expected to lead to changes in forest structure and dynamics over the long term. The responsiveness of forest to changes in resource availability varies with forest age and successional status, as well as life stage of the plant. Young plants (tree seedlings) should be more responsive to changes in nutrient availability than mature forest trees. Further, any demographic response is more likely to be visible in seedlings before mature trees because of the more rapid dynamics of seedlings. Therefore, I studied the regeneration dynamics of montane forest to understand which nutrient-related processes are involved in the growth and establishment of seedlings at both the individual and community levels. The Ecuadorian NUtrient Manipulation EXperiment (NUMEX) has been designed to study the response of montane forest to moderate fertilization. The experiment has been set up over an elevation range across three main study sites (1000 m. a.s.l: Bombuscaro; 2000 m. a.s.l: San Francisco; 3000 m. a.s.l: Cajanuma) within the Podocarpus National Park and San Francisco Reserve. The factorial experiment consists of four blocks containing four experimental plots (N, P, NP and control) in every study site (16 plots per elevation). Fertilization has been done since 2008, adding moderate quantities of fertilizer (50 kg ha–1 y–1 of N and/or 10 kg ha–1 y–1 of P). Three different approaches were used to assess the seedling community and common species responses to fertilization. First, the seedling community was monitored in natural forest at 2000 m over three consecutive years (2011 – 2013). In 192 monitoring subplots (1m2 each) in San Francisco, all seedlings and saplings were mapped and tagged. Measurements of height, diameter, herbivory and leaf production were recorded for all individuals, and the number of recruited and dead seedlings was counted every year. Second, allocation patterns and stoichiometry of seedlings of the six most common species were determined across the elevation gradient. Naturally occurring seedlings of the following species were harvested: Clarisia racemosa and Pouteria torta in Bombuscaro, Graffenrieda emarginata and Palicourea angustifolia in San Francisco and Grafferieda harlingii and Hedyosmum purpuracens in Cajanuma. Morphology (biomass allocation, herbivory and foliar areas such as SLA, LAR and LA) and foliar stoichiometry (nutrient contents and N:P ratios) were recorded and analyzed. Third, a reciprocal transplantation experiment (STE) within the NUMEX experimental plots monitored seedlings of the most common species in Bombuscaro, (Pouteria torta), to assess specific responses in seedling performance over one year. These three levels of analysis showed differential responses of the species community and common species to fertilization. Contrary to lowland forests, the community seedlings in this montane forest showed a moderate response. The density of seedling individuals decreased following nutrient addition, as a result of lower recruitment in treatments plots, but mortality was unaffected. Consequently, density-dependent mechanisms activated by additional N uptake did not show evidence of being a main driver to changes in species density. The community of seedlings did not show significant growth in response to fertilization although seedlings were higher in the N treatment. However, N addition increased herbivory over all seedlings and plots, which could have masked other seedlings growth responses. Common species seem to be well adapted to the relatively poor soils since these species were not favored by nutrient addition showing no change in growth or leaf traits. Over the long-term, common species might lose their dominance by being less competitive than faster growing species. However, it would need a longer period of monitoring under continued fertilization to produce visible shifts in community composition. Species-specific responses are hard to determine in such species rich communities. Therefore, analyses of the six common species were conducted to complement the community-level study. Leaf morphology and foliar stoichiometry were assessed from harvested seedling from the experimental plots. Both N and P had effects on seedling traits, suggesting co-limitation of N and P in montane tree species in all elevations. However, foliar N:P ratios and the more frequently response to P addition indicated this nutrient might be more limiting than N along the gradient. Responsiveness of the studied species varied between the six species. Stronger foliar P concentration compared with N in all species but Pouteria torta suggested higher P consumption in montane forest species, similar to the responses of several other tropical plant species after P addition. Increased herbivory was only evident in the opposite responses of G. emarginata (N and N+P addition) and P. angustifolia (P addition) at 2000 m suggesting that plant selection by herbivores is driven by resource quality. Most of the species had higher belowground biomass in root fractions following fertilization, except for G. harlingii at 3000 m, the only species that showed significant increase in aboveground biomass after nutrient addition. Pouteria torta seedlings showed no significant changes in species performance after fertilization one year after establishment of the STE. Pouteria seems to be well-adapted to poor soils, since N and P addition did not alter foliar nutrient concentrations. Nevertheless, N and N+P addition significantly increased mortality and diameter growth rates. P addition resulted in higher leaf area loss and shifted carbon allocation to root growth. These responses indicated consequences in the competitive strength in the understory and recruitment success of Pouteria torta. However, the ambiguous response in some attributes (growth and herbivory) not related with mortality made it difficult to predict the future abundance of this species in long term. In conclusion, complementary studies demonstrated that specific nutrient limitation for N or P in montane species seem not to be a rule in rich species ecosystems although nutrient addition did alter some pattern of growth and survival. Thus, nutrient fertilization might affect various mechanisms and dynamics of plant communities, the balance of which will only play out over long time scales.

Abstract:

In the past two decades, the tropical montane rain forests in south Ecuador experienced increasing deposition of reactive nitrogen mainly originating from Amazonian forest fires, while Saharan dust inputs episodically increased deposition of base metals. Increasing air temperature and unevenly distributed rainfall have allowed for longer dry spells in a perhumid ecosystem. This might have favored mineralization of dissolved organic matter (DOM) by microorganisms and increased nutrient release from the organic layer. Environmental change is expected to impact the functioning of this ecosystem belonging to the biodiversity hotspots of the Earth. In 2007, we established a nutrient manipulation experiment (NUMEX) to understand the response of the ecosystem to moderately increased nutrient inputs. Since 2008, we have continuously applied 50 kg ha-1 a-1 of nitrogen (N), 10 kg ha-1 a-1 of phosphorus (P), 50 kg + 10 kg ha-1 a-1 of N and P and 10 kg ha-1 a-1 of calcium (Ca) in a randomized block design at 2000 m a.s.l. in a natural forest on the Amazonia-exposed slopes of the south Ecuadorian Andes. Nitrogen concentrations in throughfall increased following N+P additions, while separate N amendments only increased nitrate concentrations. Total organic carbon (TOC) and dissolved organic nitrogen (DON) concentrations showed high seasonal variations in litter leachate and decreased significantly in the P and N+P treatments, but not in the N treatment. Thus, P availability plays a key role in the mineralization of DOM. TOC/DON ratios were narrower in throughfall than in litter leachate but their temporal course did not respond to nutrient amendments. Our results revealed an initially fast, positive response of the C and N cycling to nutrient additions which declined with time. TOC and DON cycling only change if N and P supply are improved concurrently, while NO3-N leaching increases only if N is separately added. This indicates co-limitation of the microorganisms by N and P. The current increasing reactive N deposition will increase N export from the root zone, while it will only accelerate TOC and DON turnover if P availability is simultaneously increased. The Saharan dust-related Ca deposition has no impact on TOC and DON turnover.

Abstract:

Atmospheric nutrient deposition and climate change are expected to endanger the diversity of tropical forest ecosystems. Nitrogen (N) deposition might influence nutrient fluxes beyond the N cycle by a concomitant increased demand for other nutritional elements such as phosphorus (P). Organisms might respond to the increased P demand by enhanced activity of enzymes involved in releasing inorganic P from organic matter (OM). Our aims were to assess the effect of i) climate shifts (approximated by an altitudinal gradient), and ii) nutrient addition (N, P, N+P) on phosphatase activity (PA) in organic layer and mineral soil of a tropical montane rainforest in Southern Ecuador. A nutrient manipulation experiment (NUMEX) was set up along an altitudinal gradient (1000, 2000, and 3000 m a.s.l.). We determined PA and inorganic and total P concentrations. PA at 1000 m was significantly lower (mean ± standard error: 48 ± 20 µmol p-NP g-1 dm h-1) as compared to 2000 m and 3000 m (119 ± 11 and 137 ± 19, respectively). One explanation might be that very rapid decomposition of OM at 1000 m results in very thin organic layers reducing the stabilization of enzymes and thus, resulting in leaching loss of enzymes under the humid tropical climate. We found no effect of N addition on PA neither in the organic layer nor in mineral soil, probably because of the low nutrient addition rates that showed ambiguous results so far on productivity measures as a proxy for P demand. In the organic layers of P and N+P treatments, we found decreased PA and increased concentrations of inorganic P. This indicates that the surplus of inorganic P reduced the biosynthesis of phosphatase enzymes. PA in megadiverse montane rainforests is likely to be unaffected by increased atmospheric N deposition but reduced upon atmospheric P deposition.

Abstract:

Tropical ecosystems have an important role in global change scenarios, in part because they serve as a large terrestrial carbon pool. Carbon protection is mediated by soil aggregation processes, whereby biotic and abiotic factors influence the formation and stability of aggregates. Nutrient additions may affect soil structure indirectly by simultaneous shifts in biotic factors, mainly roots, and fungal hyphae, but also via impacts on abiotic soil properties. Here, we tested the hypothesis that soil aggregation will be affected by nutrient additions primarily via changes in arbuscular mycorrhizal fungal (AMF) hyphae and root length in a pristine tropical forest system. Therefore, the percentage of water-stable macroaggregates (> 250 ?m) (WSA) and the soil mean weight diameter (MWD) was analyzed, as well as nutrient contents, pH, root length, and AMF abundance. Phosphorus additions significantly increased the amount of WSA, which was consistent across two different sampling times. Despite a positive effect of phosphorus additions on extra-radical AMF biomass, no relationship between WSA and extra-radical AMF nor roots was revealed by regression analyses, contrary to the proposed hypothesis. These findings emphasize the importance of analyzing soil structure in understudied tropical systems, since it might be affected by increasing nutrient deposition expected in the future.

Abstract:

Studies in temperate systems provide evidence that the abundance of arbuscular mycorrhizal fungal (AMF) depends on soil nutrient availability, which is mainly explained in the context of resource stoichiometry and differential plant biomass allocation. We applied this concept to an understudied ecosystem – tropical montane forest – analyzing root and AMF abundance along an elevational gradient with decreasing nutrient availability, combined with responses to nitrogen (N) versus phosphorus (P) additions. At three sites from 1000 to 3000 m above sea-level we analyzed fine root length, AMF root colonization as well as extraradical AMF biomass (neutral lipid fatty acid 16:1?5, hyphal length and spore counts) in a nutrient manipulation experiment. We found a significant increase in root length as well as intra- and extraradical AMF abundance with elevation. Overall, P additions significantly increased, whereas N additions decreased AMF abundance, with differential though nonsystematic changes along the elevational gradient. Strongest effects were clearly observed at the intermediate site. These findings suggest a general dependency of roots and AMF on nutrient availability, though responses to N and P additions differed from previous studies in temperate systems. In the context of future nutrient depositions, results suggest diverging responses of AMF abundance depending on site characteristics.

Abstract:

Nutrient deposition to tropical forests is increasing, which could affect soil fluxes of nitrous oxide (N2O), a powerful greenhouse gas. We assessed the effects of 35–56 months of moderate nitrogen (N) and phosphorus (P) additions on soil N2O fluxes and net soil N-cycling rates, and quantified the relative contributions of nitrification and denitrification to N2O fluxes. In 2008, a nutrient manipulation experiment was established along an elevation gradient (1000, 2000, and 3000 m) of montane forests in southern Ecuador. Treatments included control, N, P, and N+P addition (with additions of 50 kg N ha?1 yr?1 and 10 kg P ha?1 yr?1). Nitrous oxide fluxes were measured using static, vented chambers and N cycling was determined using the buried bag method. Measurements showed that denitrification was the main N2O source at all elevations, but that annual N2O emissions from control plots were low, and decreased along the elevation gradient (0.57 ± 0.26–0.05 ±0.04 kg N2O-N ha?1 yr?1). We attributed the low fluxes to our sites' conservative soil N cycling as well as gaseous N losses possibly being dominated by N2. Contrary to the first 21 months of the experiment, N addition did not affect N2O fluxes during the 35–56 month period, possibly due to low soil moisture contents during this time. With P addition, N2O fluxes and mineral N concentrations decreased during Months 35–56, presumably because plant P limitations were alleviated, increasing plant N uptake. Nitrogen plus phosphorus addition showed similar trends to N addition, but less pronounced given the counteracting effects of P addition. The combined results from this study (Months 1–21 and 35–56) showed that effects of N and P addition on soil N2O fluxes were not linear with time of exposure, highlighting the importance of long-term studies.

Abstract:

Nitrogen deposition to tropical forests is predicted to increase in future in many regions due to agricultural intensification. We conducted a seedling transplantation experiment in a tropical premontane forest in Ecuador with a locally abundant late-successional tree species (Pouteria torta, Sapotaceae) aimed at detecting species-specific responses to moderate N and P addition and to understand how increasing nutrient availability will affect regeneration. From locally collected seeds, 320 seedlings were produced and transplanted to the plots of the Ecuadorian Nutrient Manipulation Experiment (NUMEX) with three treatments (moderate N addition: 50 kg N ha?1 year?1, moderate P addition: 10 kg P ha?1 year?1 and combined N and P addition) and a control (80 plants per treatment). After 12 months, mortality, relative growth rate, leaf nutrient content and leaf herbivory rate were measured. N and NP addition significantly increased the mortality rate (70 vs. 54% in the control). However, N and P addition also increased the diameter growth rate of the surviving seedlings. N and P addition did not alter foliar nutrient concentrations and leaf N:P ratio, but N addition decreased the leaf C:N ratio and increased SLA. P addition (but not N addition) resulted in higher leaf area loss to herbivore consumption and also shifted carbon allocation to root growth. This fertilization experiment with a common rainforest tree species conducted in old-growth forest shows that already moderate doses of added N and P are affecting seedling performance which most likely will have consequences for the competitive strength in the understory and the recruitment success of P. torta. Simultaneous increases in growth, herbivory and mortality rates make it difficult to assess the species' overall performance and predict how a future increase in nutrient deposition will alter the abundance of this species in the Andean tropical montane forests.

Abstract:

The increasing anthropogenic influence on the environment on a global scale has led to a shift in nutrient cycles which are not fully understood yet. These shifts could alter the properties of ecosystems and therefore change habitats and species composition. Therefore it is crucial to understand nutrient cycles and how ecosystems react to these nutrient availability shifts. Especially the phosphorus and nitrogen cycles are highly important because of their direct influence on plant growth as primary macronutrients. Even in some so far undisturbed regions, a human caused rise in atmospheric input of phosphorus-containing particles is expected. One of the reasons therefor is land-use change in rural areas with associated combustions, from which the particles are transported downwind. This study simulates the prognosticated increased nutrient input and focuses on the following fate of the phosphorus in soil. For this, in 2008 a nutrient manipulation experiment was established to fertilize an old-growth tropical montane forest with moderate phosphorus (10 kg P ha-1 yr-1) and/or nitrogen additions (50 kg N ha-1 yr-1). This experimental setup was conducted for the first time at three altitudes (1000m a.s.l., 2000m a.s.l., 3000m a.s.l.) to compare the effects along an altitudinal gradient. After seven years the total phosphorus and bioavailable phosphorus concentrations of these locations were investigated in litterfall, the organic layer and in mineral soil to see if the added phosphorus is retained and enriched. The results showed that the application of the phosphorus fertilizer resulted in increased total and bioavailable phosphorus concentrations mainly in the organic layer. The phosphorus increase in mineral soil was not significant and indicated that the added phosphorus is not leached, but retained in the ecosystems. There were no effects on phosphorus concentrations after nitrogen addition. The altitudinal gradient referred to a pronounced difference between 1000m and the other altitudes (2000m and 3000m), due to the accumulated organic matter at the higher altitudes. Overall this study shows that the sustainable anthropogenic phosphorus increase may cause a change in the ecosystem’s characteristics and nutrient cycles.

Abstract:

Large areas in the tropics receive elevated atmospheric nutrient inputs. Presently, little is known on how nitrogen (N) cycling in tropical montane forest soils will respond to such increased nutrient inputs. We assessed how gross rates of mineral N production (N mineralization and nitrification) and microbial N retention (NH4+ and NO3- immobilization and dissimilatory NO3- reduction to NH4+ [DNRA]) change with elevated N and phosphorus (P) inputs in montane forest soils at 1000-, 2000-, and 3000-m elevations in south Ecuador. At each elevation, four replicate plots (20320 m each) of control, N (added at 50 kg N ha-1yr-1), P (added at 10 kg P ha-1 yr-1), and combined N x P additions have been established since 2008. We measured gross N cycling rates in 2010 and 2011, using 15N pool dilution techniques with in situ incubation of intact soil cores taken from the top 5 cm of soil. In control plots, gross soil-N cycling rates decreased with increase in elevation, and microbial N retention was tightly coupled with mineral N production. At 1000 m and 2000 m, four-year N and combined N þ P additions increased gross mineral N production but decreased NH4+ and NO3- immobilization and DNRA compared to the control. At 3000 m, four-year N and combined N x P additions increased gross N mineralization rates and decreased DNRA compared to the control; although NH4+ and NO3- immobilization in the N and NxP plots were not different from the control, these were lower than their respective mineral N production. At all elevations, decreased microbial N retention was accompanied by decreased microbial biomass C and C:N ratio. P addition did not affect any of the soil-N cycling processes. Our results signified that four years of N addition, at a rate expected to occur at these sites, uncoupled the soil-N cycling processes, as indicated by decreased microbial N retention. This fast response of soil-N cycling processes across elevations implies that greater attention should be paid to the biological implications on montane forests of such uncoupled soil-N cycling.

Abstract:

Although often overlooked in forest research, the canopy can play an important role in forest nutrient cycling. Since the canopy is spatially isolated from the forest floor, nutrient cycling in the two areas may differ as terrestrial nutrients accumulate. We measured rates of free-living N2 fixation along an elevation gradient (1,000, 2,000 and 3,000 m) of tropical montane canopy soils, compared these to rates measured in the top 5 cm of forest floor soils (excluding fresh litter), and assessed the effects of elevated nutrient inputs to the forest floor. N2 fixation was measured using the acetylene reduction assay. Measurements occurred in the field, in the wet and dry seasons, using intact cores of soil. The forest floor had been fertilized biannually with moderate amounts of nitrogen (N) and phosphorus (P) for 4 years; treatments included control, N, P and N x P. N2 fixation rates exhibited little variation with Elevation but were higher in the dry season than the wet season. Fixation was inhibited in forest floor N plots compared to control and P plots, and stimulated in canopy P plots compared to control. At 2,000 m, the canopy contributed 12 % of measured canopy and forest floor N2 fixation (1.2 kg N ha-1 year-1). Results suggest that N2 fixation is an active process in canopy soils, which is variable across seasons and sensitive to changes in terrestrial nutrient availability. Long-term terrestrial accumulation of N and/or P has the potential to significantly change the dynamics of soil N cycling in these canopies.

Abstract:

Ecosystems worldwide face increasing nutrient depositions mainly caused by anthropogenic processes. In particular, tropical ecosystems react sensitively to altering nutrient supply. The deposition of nutrients might influence the nutrient cycles, primarily of N and P in tropical montane rainforests. Increased nutrient supply leads to an enhanced biomass production and therefore other nutrients become limited for plants and microorganisms. For this reason, the aim of this thesis is to study the response of phosphatase activity (PA) on moderate fertilization along an altitudinal gradient in a tropical montane rainforest in South Ecuador. The experiment was conducted on the NUMEX study sites including three different elevation levels 1000, 2000 and 3000m a.s.l. The different plots were treated with N, P, N+P to simulate increased nutrient depositions and one control plot. Further, organic layer and mineral soil was sampled and phosphomono- and phosphodiesterase activity (PMEA and PDEA) were determined. The N fertilized plots showed only small effects compared to the control; presumably due to low amounts of added fertilizer. PA in the P addition plots showed reduced activity compared to the control with significant results of PMEA in the organic layer of the study sites on 2000 and 3000m a.s.l. The reason might be sufficient quantities of inorganic P which suppresses the production of phosphatases. Further, PA in N+P plots showed lower PA compared to the control than in the P addition plots. This effect could be caused through the dominating inhibitory effect of P in contrast to the stimulating effect of N on PA. Altitudinal differences were observed comparing the control plots at different elevation levels (1000, 2000 and 3000m a.s.l.). The results for the organic layer showed increasing PA along the altitudinal gradient with the lowest PA at 1000m a.s.l. The findings are contrary to the present literature which states that lowland tropical forests are characterized by high decomposition rates coming up with high PA. The findings cannot support this hypothesis; therefore more research is needed in the studied area in South Ecuador.

Abstract:

Although the canopy can play an important role in forest nutrient cycles, canopy-based processes are often overlooked in studies on nutrient deposition. In areas of nitrogen (N) and phosphorus (P) deposition, canopy soils may retain a significant proportion of atmospheric inputs, and also receive indirect enrichment through root uptake followed by throughfall or recycling of plant litter in the canopy. We measured net and gross rates of N cycling in canopy soils of tropical montane forests along an elevation gradient and assessed indirect effects of elevated nutrient inputs to the forest floor. Net N cycling rates were measured using the buried bag method. Gross N cycling rates were measured using 15N pool dilution techniques. Measurements took place in the field, in the wet and dry season,using intact cores of canopy soil from three elevations (1000, 2000 and 3000 m). The forest floor had been fertilized biannually with moderate amounts of N and P for 4 years; treatments included control, N, P, and N + P. In control plots, gross rates of NH4+ transformations decreased with increasing elevation; gross rates of NO3- transformations did not exhibit a clear elevation trend, but were significantly affected by season. Nutrient-addition effects were different at each elevation, but combined N + P generally increased N cycling rates at all elevations. Results showed that canopy soils could be a significant N source for epiphytes as well as contributing up to 23% of total (canopy + forest floor) mineral N production in our forests. In contrast to theories that canopy soils are decoupled from nutrient cycling in forest floor soil, N cycling in our canopy soils was sensitive to slight changes in forest floor nutrient availability.Long-term atmospheric N and P deposition may lead to increased N cycling, but also increased mineral N losses from the canopy soil system.

Abstract:

Tank bromeliads are common epiphytic plants throughout neotropical forests that store signi?cant amounts of water in phytotelmata (tanks) formed by highly modi?ed leafs. Methanogenic archaea in these tanks have recently been identi?ed as a signi?cant source of atmospheric methane. We address the effects of environmental drivers (temperature, tank water content, sodium phosphate [P], and urea [N] addition) on methane production in anaerobically incubated bromeliad slurry and emissions from intact bromeliad tanks in montane Ecuador. N addition ? 1 mg g
1 had a signi?cantly positive effect on headspace methane concentrations in incubation jars while P addition did not affect methane production at any dosage (? 1 mg g
1 ). Tank bromeliads (Tillandsia complanata) cultivated in situ showed signi?cantly increased ef?uxes of methane in response to the addition of 26 mg N addition per tank but not to lower dosage of N or any dosage of P (? 5.2 mg plant
1 ). There was no signi?cant interaction between N and P addition. The brevity of the stimulatory effect of N addition on plant methane ef?uxes (1–2 days) points at N competition by other microorganisms or bromeliads. Methane ef?ux from plants closely followed within-day temperature ?uctuations over 24 h cycles, yet the dependency of temperature was not exponential as typical for terrestrial wetlands but instead linear. In simulated drought, methane emission from bromeliad tanks was maintained with minimum amounts of water and regained after a short lag phase of approximately 24 h. Our results suggest that methanogens in bromeliads are primarily limited by N and that direct effects of global change (increasing temperature and seasonality, remote fertilization) on bromeliad methane emissions are of moderate scale.

Abstract:

Increased nitrogen (N) depositions expected in the future endanger the diversity and stability of ecosystems primarily limited by N, but also often co-limited by other nutrients like phosphorus (P). In this context a nutrient manipulation experiment (NUMEX) was set up in a tropical montane rainforest in southern Ecuador, an area identified as biodiversity hotspot. We examined impacts of elevated N and P availability on arbuscular mycorrhizal fungi (AMF), a group of obligate biotrophic plant symbionts with an important role in soil nutrient cycles. We tested the hypothesis that increased nutrient availability will reduce AMF abundance, reduce species richness and shift the AMF community toward lineages previously shown to be favored by fertilized conditions. NUMEX was designed as a full factorial randomized block design. Soil cores were taken after 2 years of nutrient additions in plots located at 2000 m above sea level. Roots were extracted and intraradical AMF abundance determined microscopically; the AMF community was analyzed by 454-pyrosequencing targeting the large subunit rDNA. We identified 74 operational taxonomic units (OTUs) with a large proportion of Diversisporales. N additions provoked a significant decrease in intraradical abundance, whereas AMF richness was reduced significantly by N and P additions, with the strongest effect in the combined treatment (39% fewer OTUs), mainly influencing rare species. We identified a differential effect on phylogenetic groups, with Diversisporales richness mainly reduced by N additions in contrast to Glomerales highly significantly affected solely by P. Regarding AMF community structure, we observed a compositional shift when analyzing presence/absence data following P additions. In conclusion, N and P additions in this ecosystem affect AMF abundance, but especially AMF species richness; these changes might influence plant community composition and productivity and by that various ecosystem processes.

Abstract:

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 × 5 kg 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.

Abstract:

Zu Beginn der Untersuchungen wurden im ersten Schritt die freien Aluminiumkonzentrationen mit Hilfe der Donnan Membran Technik bestimmt. Anschließend wurden im zweiten Schritt die gemessenen freien Aluminiumkonzentrationen mit den modellierten Aluminiumkonzentrationen verglichen. Hierbei stellte sich heraus, dass die Werte vergleichbar sind und somit das Programm Visuell Minteq zur Modellierung der Aluminiumspeziierung eingesetzt werden konnte. Es wurde weiter festgestellt, dass der Anteil des freien Aluminiums an der gesamten Aluminiumkonzentration mit der Bodentiefe zunimmt. Der Anteil des freien Aluminiums an der gesamten Al-Konzentration betrug in der Bodenlösung unterhalb der organischen Auflage in allen untersuchten Plots lediglich 0% bis 0,96% (Mittelwert 0,0451%), während er im Mineralboden, bedingt durch geringe DOC-Werte, zwischen 0% und 29,11% (15 cm Tiefe) sowie zwischen 0% und 36,88% (30 cm Tiefe) schwankte. Der dritte Schritt galt der Untersuchung des Ca/Al Verhältnisses. Das Ergebnis zeigte hier, dass das freie molare Ca2+/Al3+ Verhältnis durch die Calciumdüngung nur geringfügig beeinflusst wird und über dem von Cronan & Grigal (1995) als kritisch betrachtetem Wert von <1 liegt. Calcium tritt daher vermutlich als wachstumslimitierender Faktor auf. Eine direkte Aluminiumtoxizität ist in den untersuchten Bodenlösungen nicht bzw. nur in geringem Maße festzustellen gewesen