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

Abstract:

Texture information from passive remote sensing images provides surrogates for habitat structure, which is relevant for modeling biodiversity across space and time and for developing effective ecological indicators. However, the applicability of this information might differ among taxa and diversity measures. We compared the ability of indicators developed from texture analysis of remotely sensed images to predict species richness and species turnover of six taxa (trees, pyraloid moths, geometrid moths, arctiinae moths, ants, and birds) in a megadiverse Andean mountain rainforest ecosystem. Partial least-squares regression models were fitted using 12 predictors that characterize the habitat and included three topographical metrics derived from a high-resolution digital elevation model and nine texture metrics derived from very high-resolution multi-spectral orthophotos. We calculated image textures derived from mean, correlation, and entropy statistics within a relatively broad moving window (102 m × 102 m) of the near infra-red band and two vegetation indices. The model performances of species richness were taxon dependent, with the lowest predictive power for arctiinae moths (4%) and the highest for ants (78%). Topographical metrics sufficiently modeled species richness of pyraloid moths and ants, while models for species richness of trees, geometrid moths, and birds benefited from texture metrics. When more complexity was added to the model such as additional texture statistics calculated from a smaller moving window (18 m × 18 m), the predictive power for trees and birds increased significantly from 12% to 22% and 13% to 27%, respectively. Gradients of species turnover, assessed by non-metric two-dimensional scaling (NMDS) of Bray-Curtis dissimilarities, allowed the construction of models with far higher predictability than species richness across all taxonomic groups, with predictability for the first response variable of species turnover ranging from 64% (birds) to 98% (trees) of the explained change in species composition, and predictability for the second response variable of species turnover ranging from 33% (trees) to 74% (pyraloid moths). The two NMDS axes effectively separated compositional change along the elevational gradient, explained by a combination of elevation and texture metrics, from more subtle, local changes in habitat structure surrogated by varying combinations of texture metrics. The application of indicators arising from texture analysis of remote sensing images differed among taxa and diversity measures. However, these habitat indicators improved predictions of species diversity measures of most taxa, and therefore, we highly recommend their use in biodiversity research.

Abstract:

To improve the current knowledge of the rainfall–runoff phenomena of tropical montane catchments, we explored the usefulness of several hydrological indicators on a nested cloud forest catchment (76.9 km2). The used metrics belong to 5 categories: baseflow mean transit time, physicochemical properties of stream water, land cover, topographic, and hydrometric parameters. We applied diverse statistical techniques for data analysis and to contrast findings. Multiple regression analysis showed that mean transit times of base flow could be efficiently predicted by sodium concentrations (higher during baseflows) and temperatures of stream water, indicating a major influence of geomorphology rather than topographic or land cover characteristics. Principal component analysis revealed that no specific subset of catchment indicators could be identified as prevailing descriptors for all catchments. The agglomerative hierarchical clustering analysis provided concomitant results, implying larger levels of dissimilarity between smaller subcatchments than between larger ones. Overall, results point out an intricate interdependence of diverse processes at surface and subsurface level indicating a high level of heterogeneity. Disregarding heterogeneity of nested or paired catchments could lead to incomplete or misleading conclusions, especially in tropical mountain regions where pronounced spatial and temporal gradients are present.

Abstract:

Trees in tropical dry forests (TDFs) have manifold drought coping strategies including succulence of different plant organs, wood anatomical traits and leaf phenology. As water availability to plants is the limiting factor for physiological activity, changes in precipitation patterns are assumed to have strong influences on tree phenology, growth and water turnover. Our objectives were to assess patterns in leaf phenology, radial stem circumference changes and sap flux responses to fluctuating moisture regimes of selected species. Based on these findings we evaluated the potential suitability as indicator species for climate change effects. The study was implemented at different elevational positions in a submontane dry forest of southern Ecuador. Annual rainfall is 600 mm with an eight months dry period; moisture availability slightly increases with altitude because of moist air coming from the Pacific. At three altitudes,we studied the tree species Ceiba trichistandra (leaf deciduous, stem succulent), Eriotheca ruizii (leaf deciduous, root succulent) and Erythrina velutina (leaf deciduous). Reversible stem swelling and shrinking was observed for all three species during the whole study period and at all positions at the altitudinal gradient. However, it was most pronounced and sensitive in the stem succulent C. trichistandra and at the lowest (driest) position. C. trichistandra flushed leaves at dry season intermittent rain events, and from dry to wet season leaf out was earlier, and in this period sap flux was high while stem circumference decreased. Length of the leaved periods of all species increased with altitude. Thus, clear differences among species, topographic positions, radial growth and tree water use patterns are revealed; especially C. trichistandra responded very sensitive to fluctuating moisture regimes with leaf phenology, sap flux and stem diameter variations, and can be regarded as a sensitive indicator for assessing climatic variations.

Abstract:

Decomposition is a very important process. It is supporting nutrient cycles and for that reason crucial for the functionality of ecosystems. Various studies have shown that decomposition is driven by abiotic and biotic factors such as climate, seasonal weather conditions (like el Niño), soil organisms and disturbance of forests. In this study, the focus lies on investigating the impact of abiotic factors on the decomposition with a new simplified and standardised method, with tea bags as litterbags. Green tea bags were buried in two years, on different elevations between 1000 m a.s.l. and 3000 m a.s.l. in 500 m steps on disturbed and undisturbed plots in the Ecuadorian Andes. After 21 days in the soil, the decomposition rate were calculated and compared on different elevations and different disturbances in two different years. The disturbance had no effect, because the disturbed plots were fragmented and the study took place in the interior of these fragments; here no edge effects influenced the decomposition rate. As expected, the decomposition rate declined with an increase in elevation, due to changing climate conditions. In the lower, wet and warm elevations, it was faster than in the higher elevation where the climate is wetter but also colder. In the test series from 2015, the decomposition rate was higher in the upper elevations than in 2014, this could be explained by a weather phenomenon called el Niño, which took place in 2015. It leads to a slightly higher temperature and 60% more precipitation than what normally would be expected in 2015. These differences between the two years show the importance of taking seasonal and annual variations into account while investigating decomposition and the sensitivity of this important ecosystem process to changing weather and climate conditions.

Abstract:

Climate change and forest degradation are major threats to forest ecosystems. The climatic changes can affect the mineralization rate in soils and therefore change the amount of available nitrogen components for plants. Thus these changes can influence foliar C/N-ratio. Previous studies used an elevation gradient to test the influence of climatic changes on ecosystems, and showed that foliar C/N-ratio is positively correlated with elevation. Changes in foliar C/N-ratio may affect the feeding behavior of insect herbivores and therefore influence leaf area loss (LAL). LAL is used as an indicator of herbivory and therefore may consequently also be related to ant abundance because ants prey on insect herbivores. Ants in return, are also affected by the abiotic changes with increasing elevation and therefore sensitive to climate change. However, forest degradation is currently the greater threat to ecosystems. Degradation can lead to changes in the nutrient uptake of plants and therefore to changes in foliar C/N-ratio. Moreover it can reduce ant abundance as ants are sensitive to the conversion of forests. We nalyzed the causal effects of abiotic factors (elevation and degradation) on biotic factors (foliar C/N-ratio and ant activityabundance) and the relationships between the biotic factors and LAL with linear mixed-effect models in a path model. We found a significant positive correlation between foliar C/N-ratio and elevation, and a significant negative correlation between ant activity-abundance and elevation. Foliar carbon-content had a significant negative correlation with degradation. Our results revealed no significant relationships with LAL, but LAL correlated negatively with foliar C/N-ratio in the shrub layer. The results of our study point out that foliar C/N-ratio and ant activityabundance change with elevation and thus respond to climatic changes. Moreover our results indicate that foliar C/N-ratio can be connected to LAL, but that influences such as the abundance and distribution of insect herbivores along elevation should be examined closely. Our finding that foliar C/N-ratio, ant activity-abundance and LAL were unaffected by degradation indicates that the studied degraded forests are still suitable habitats for insect herbivores and ants.

Abstract:

Biodiversity monitoring is a global need due to environmental degradation and climate change. Birds are often monitored indicators for environmental health because they are easy assessable, fast reacting species, which supply important ecosystem services. But conventional biodiversity monitoring can be timeconsuming, invasive and it often needs trained specialists, thus appropriate alternatives, especially for largescale assessments, like acoustic monitoring are needed. While numerous acoustic indices were generated no studies occur about their ability to reflect conventional alpha-diversity measures such as species richness, abundance, diversity or evenness under complex conditions like a gradient of elevation and degradation. We choose bird communities along an elevation gradient in natural and degraded rainforests in a highdiverse region in the Ecuadorian Andes to investigate the association among five conventional (total abundance, species richness, Shannon Index, evenness, Gini Index) and three acoustic (Acoustic Diversity Index [ADI], Bioacoustic Index [BI], Acoustic Evenness Index [AEI]) alpha-diversity measures. Furthermore, we investigated the influence of higher resolutions (FFT window size, frequency band size) on the acoustic indices. Further, we tested a new acoustic beta-diversity measure. With help of multivariate analyzes we found acoustic diversity measures were not significantly correlated with conventional diversity measures in high-diverse rainforests. The acoustic indices were correlated between another. The values of the acoustic indices increased with higher resolutions and were strongly positively correlated with their next higher resolution. The NMDS of the conventionally assessed bird communities resulted an elevation gradient and a distinct separation between degraded and natural forest communities. The NMDS of the acoustically assessed bird communities resulted no distinct gradients. The conventional and acoustic alpha-diversity measures were not significantly associated with the elevation or habitat type. ADI, BI and AEI seem not appropriate to reflect conventional diversity measures in high-diverse ecosystems, but BI can reflect avian abundance in less diverse ecosystems. Avian acoustic activity was driven by noisy species and not by species richness or abundance. Higher resolutions of the acoustic indices did not resolute acoustic activity more exactly in this case, thus for communities where birds with long or complex calls (relevant for BI) or narrow frequency ranges (relevant for ADI/AEI) are not dominating the acoustic activity the default resolutions of the acoustic indices seem sufficient. Acoustic beta-diversity assessment seems problematic because the analyzes of frequencies does not reflect species turnover among communities due to the reason that several species can occupy the same frequencies with their calls.

Abstract:

Environmental stressors and changes in land use have led to rapid and dramatic species losses. As such, we need effective monitoring programs that alert us not only to biodiversity losses, but also to functional changes in species assemblages and associated ecosystem processes. Ants are important components of terrestrial food webs and a key group in food web interactions and numerous ecosystem processes. Their sensitive and rapid response to environmental changes suggests that they are a suitable indicator group for the monitoring of abiotic, biotic, and functional changes. We tested the suitability of the incidence (i.e. the sum of all species occurrences at 30 baits), species richness, and functional richness of ants as indicators of ecological responses to environmental change, forest degradation, and of the ecosystem process predation on herbivorous arthropods. We sampled data along an elevational gradient (1000–3000 m a.s.l.) and across seasons (wetter and drier period) in a montane rainforest in southern Ecuador. The incidence of ants declined with increasing elevation but did not change with forest degradation. Ant incidence was higher during the drier season. Species richness was highly correlated with incidence and showed comparable results. Functional richness also declined with increasing elevation and did not change with forest degradation. However, a null-model comparison revealed that the functional richness pattern did not differ from a pattern expected for ant assemblages with randomly distributed sets of traits across species. Predation on artificial caterpillars decreased along the elevational gradient; the pattern was not driven by elevation itself, but by ant incidence (or inter-changeable by ant richness), which positively affected predation. In spite of lower ant incidence (or ant richness), predation was higher during the wetter season and did not change with forest degradation and ant functional richness. We used path analysis to disentangle the causal relationships of the environmental factors temperature (with elevation as a proxy), season, and habitat degradation with the incidence and functional richness of ants, and their consequences for predation. Our results would suggest that the forecasted global warming might support more active and species-rich ant assemblages, which in turn would mediate increased predation on herbivorous arthropods. However, this prediction should be made with reservation, as it assumes that the dispersal of ants keeps pace with the climatic changes as well as a one-dimensional relationship between ants and predation within a food-web that comprises species interactions of much higher complexity. Our results also suggested that degraded forests in our study area might provide suitable habitat for epigaeic, ground-dwelling ant assemblages that do not differ in incidence, species richness, functional richness, composition, or predation on arthropods from assemblages of primary forests. Most importantly, our results suggest that the occurrence and activity of ants are important drivers of ecosystem processes and that changes in the incidence and richness of ants can be used as effective indicators of responses to temperature changes and of predation within mega-diverse forest ecosystems.

Abstract:

Land-use and climate change are major threats to biodiversity and ecosystem functions. Most of the current biodiversity monitoring systems are based on periodic records of the populations of a set of threatened or popular ‘flagship’ indicator species. In contrast to the abundance-based monitoring of species, also specific indicators of processes and functional interactions in an ecosystem may become targets of a more functional monitoring which can unveil early responses of an ecosystem to environmental changes at different spatial and temporal scales. The contributions of this Special Issue present such functional indicators for assessing and predicting responses to environmental changes of ecosystem functions in a hotspot of tropical biodiversity.

Abstract:

Increasing land-use conflicts call for the development of land-use systems that reconcile agricultural production with the provisioning of multiple ecosystem services, including climate change mitigation. Agroforestry has been suggested as a global solution to increase land-use efficiency, while reducing environmental impacts and economic risks for farmers. Past research has often focused on comparing tree-crop combinations with agricultural monocultures, but agroforestry has seldom been systematically compared to other forms of land-use diversification, including a farm mosaic. This form of diversification mixes separate parcels of different land uses within the farm. The objective of this study was to develop a modelling approach to compare the performance of the agroforestry and farm mosaic diversification strategies, accounting for tree-crop interaction effects and economic and climate uncertainty. For this purpose, Modern Portfolio Theory and risk simulation were coupled with the process-based biophysical simulation model WaNuLCAS 4.0. For an example application, we used data from a field trial in Panama. The results show that the simulated agroforestry systems (Taungya, alley cropping and border planting) could outperform a farm mosaic approach in terms of cumulative production and return. Considering market and climate uncertainty, agroforestry showed an up to 21% higher economic return at the same risk level (i.e. standard deviation of economic returns). Farm compositions with large shares of land allocated to maize cultivation were also more severely affected by an increasing drought frequency in terms of both risks and returns. Our study demonstrates that agroforestry can be an economically efficient diversification strategy, but only if the design allows for economies of scope, beneficial interactions between trees and crops and higher income diversification compared to a farm mosaic. The modelling approach can make an important contribution to support land-use decisions at the farm level and reduce land-use conflicts at the landscape level.

Abstract:

Postulating that megadiverse forests are more profitable than monocultures lacks economic evidence and may put conservation incentives at risk. Letter in Response to Liang et al. published in Science on October 14

Abstract:

The effects of an increasing moisture on trees of the tropical species-rich mountain rain forest in the South Ecuadorian Andes was investigated, using the daily total water consumption (TWC) and the instantaneous water use efficiency (WUE, ratio of photosynthetic CO2 uptake per water loss by transpiration) as ecophysiological indicators. Two canopy and one sub-canopy tree species, (Vismia tomentosa, Clusiaceae, an as of yet unknown Lauracee, and Spirotheca rosea, Bombacaceae) were the experimental objects. Seasonal changes as well as a long-term (18 months) trend of increasing precipitation caused an inverse reaction of the TWC of the trees. Because of a rather unlimited water supply to the trees from a permanently high water content of the soil, transpiration followed mainly the atmospheric demand of water vapor, and increasing moisture hence reduced water loss by transpiration. It was hypothesized that in spite of the reduction in transpiratory water loss photosynthetic carbon acquisition would be not or less affected due to an increase in water use efficiency. Concomitant measurements of photosynthetic net CO2 uptake showed the expected increase of WUE in V. tomentosa and S. rosea, but no clear reaction of the Lauracee. Accompanying measurements of stem extension growth confirmed an undiminished growth of V. tomentosa and S. rosea but showed also suspended growth of the Lauracee during the wettest months. While TWC can be continuously monitored with the heat dissipation technique, WUE is determined by leaf porometry in campaigns for which access to the canopy is required. Simultaneous recordings of the gas exchange of leaves at 4 different positions in the crown of one of the experimental trees (V. tomentosa) showed the usability of the trait WUE in combination with the total daily water consumption as indicator set for assessing the response of trees to a subtly changing climate. However, not all tree species appear as likewise useful indicator trees.

Abstract:

Human influence and its impacts are perceptible in all ecosystems resulting in land transformation, changes in global biogeochemistry, climate change, and loss of biological diversity. Mapping the spatial and temporal patterns of human influence is essential to address land use management and conservation programs. In this study, we tailored the Human Footprint index (HF) developed at global level to evaluate the spatial and temporal patterns of human pressure in South Ecuador for 1982, 1990 and 2008. Landscape and ecosystem levels were analyzed to identify the contribution of different human proxies to the HF.We also used the HF to evaluate the effectiveness of protected areas to reduce human pressure in the surrounding landscape. We found that levels of human pressure increased and the wildest areas decreased since 1982. We identified important “hotspots of changes” in the seasonally dry forests in the western part and the premontane evergreen forest in the eastern part of the study area. Our results show that each human proxy contributes in a different way to the observed values of HF in the studied ecosystems. Finally, we found that Podocarpus NP, the most important protected area in our study region, seems to be partially effective in reducing human pressure inside and in the buffer zones where only a low increase in HF was detected. However, the HF values observed in the surrounding landscape were higher than those observed in the buffer zone and inside the protected area. We demonstrated that HF could be a useful regional evaluation tool to facilitate conservation planning.

Abstract:

In einem tropischen Bergregenwaldgebiet in Ecuador, haben verschiedene Forschungsarbeiten gezeigt, dass auch Mikronährelemente einen Einfluss auf die mikrobielle Aktivität haben können. Mittels eines Inkubationsversuches mit einer Streuauflagenprobe aus dem Untersuchungsgebiet, wurde analysiert ob eine multiple Mikronährlimitierung von Mikroorganismen vorliegt. Dabei wurde die Streuprobe in einem sechs Tage dauernden Inkubationsversuch täglich mit verschiedenen Mikronährelementen (Cu, Mo, Mn, Zn, P)gedüngt. Danach wurde die Nährlösung abgesaugt und die daraus entstandenen Eluate mithilfe eines Flammen-Atomabsorptionsspektrometer auf die verbliebenen Metallgehalte analysiert. Zuverlässige Werte konnten nur für Cu, Mn und Zn gemessen werden, die Werte für Mo und P lagen unter der Nachweis- oder Bestimmungsgrenze. Bei der Kupfermessung konnte die höchste Kupferkonzentration in der Cu-gedüngten Probe gemessen werden, doch es wurde schon weniger Cu ausgewaschen, wenn zusätzlich mit einem anderen Mikronährelement gedüngt wurde als in den Kontrolldüngungen. Für Mn wurde in der Kontrolllösung die höchste Mn-Konzentration gemessen, in allen anderen Proben mit zusätzlicher Düngung wurde Mn stärker zurückgehalten. Die gemessene Zn-Konzentration bei Düngung mit Cu, Mn, Mo und P lag fast bei null. Zink wurde sehr stark zurückgehalten. Auch in der Zink-Gedüngten Probe wurde Zn in den ersten drei Tagen stark zurückgehalten und wurde erst danach stärker ausgewaschen als in den Kontrollproben. Aufgrund der Resultate ist zu erkennen, dass auch Mikronährelemente eine limitierende Wirkung auf Mikroorganismen haben können und die Mikroorganismen im untersuchten Gebiet durch verschiedene Mikronährelemente limitiert sind.

Abstract:

Growth limitation induced by Al toxicity is believed to commonly occur in tropical forests, although a direct proof is frequently lacking. To test for the general assumption of Al toxicity, Al, Ca, and Mg concentrations in precipitation, throughfall, stemflow, organic layer leachate, mineral soil solutions, stream water, and the leaves of 17 native tree species were analyzed. We calculated Al fluxes and modeled Al speciation in the litter leachate and mineral soil solutions. We assessed potential Al toxicity based on soil base saturation, Al concentrations, Ca:Al and Mg:Al molar ratios and Al speciation in soil solution as well as Al concentrations and Ca:Al andMg:Al molar ratios in tree leaves. High Al fluxes in litterfall (8.77±1.3 to 14.2±1.9 kg ha?1 yr?1, mean ± SE) indicated a high Al circulation through the ecosystem. The fraction of exchangeable and potentially plant-available Al in mineral soils was high, being a likely reason for a low root length density in the mineral soil. However, Al concentrations in all solutions were consistently below critical values and Ca:Al molar and the Ca2+:Alinorganic molar ratios in the organic layer leachate and soil solutions were above 1, the suggested threshold for Al toxicity. Except for two Al-accumulating and one non-accumulating tree species, the Ca:Al molar ratios in tree leaves were above the Al toxicity threshold of 12.5. Our results demonstrate high Al cycling through the vegetation partly because of the presence of some Al accumulator plants. However, there was little indication of an Al toxicity risk in soil and of acute Al toxicity in plants likely reflecting that tree species are well adapted to the environmental conditions at our study site and thus hardly prone to Al toxicity.

Abstract:

The forests of South Ecuador have high conservation value because they are highly threatened, but also possess high diversity and endemism levels. However, as the process of conservation is inherently spatial, the little available information about the factors that promote vulnerability of these forests as well as the spatial distribution of the conservation targets have produced some conservation pitfalls. The main objective of this dissertation was to generate spatial information about the threats and biological values occurring in this region to prioritize areas for conservation. The second chapter analyzes the deforestation and fragmentation patterns in the region since the 1970s using aerial photographs and satellite images to identify areas with remaining forest, fronts of deforestation, annual deforestation rates and the dynamics in the composition and configuration of the landscape of South Ecuador. We recorded annual deforestation rates of 0.75% (1976 – 1989) and 2.86% (1989 – 2008) for two consecutive survey periods. Also, we found that South Ecuador is experiencing an ongoing fragmentation process due to an increase in the number of patches, a decrease in mean patch size and an increase in the isolation of forest fragments. This study also contributes to a better understanding of forest change dynamics in the tropics. We found that substantial portions of natural forests are being degraded or converted into pastures and that main fronts of deforestation are located in the lowest areas in the premontane evergreen forest. The third chapter describes the temporal and spatial patterns of human pressure, as it is one of the main factors that influence the effectiveness of conservation strategies. At local scale, we adapted at local scale the Human Footprint Index (HF) developed by Sanderson et al. (2002), to evaluate spatial changes in HF during a 26 year period at both landscape and ecosystem levels. This information allowed us to identify “hotspots of change” and the wildest areas remaining in order to evaluate how different human proxies contribute to HF and to demonstrate how effective the most important protected areas have been in reducing human pressure inside and outside their boundaries. The findings show a noticeable 4 increase in human pressure levels in South Ecuador and a progressive reduction in the wildest areas. We also identified that the important “hotspots of changes” are located in the western region and the Rio Zamora river basin. The most impacted vegetation types were seasonally dry forest and shrubland. Here, population density is the human proxy with the highest contribution to the observed patterns. Finally, we found that Podocarpus National Park has been partially effective in reducing human pressure inside and outside its borders. HF levels have increased inside and outside the boundaries of the protected area, but the human pressure was always lower than that observed in the surrounding landscape. The fourth chapter analyzes the patterns of alpha and beta diversity of endemic plant species to evaluate the congruence of both patterns and to identify areas with the highest diversity of endemic plants in order to prioritize areas for conservation. We found that hotspots of alpha diversity are concentrated along the Andes, but this diversity was only slightly congruent with beta diversity patterns of endemic plants mostly concentrated in the western and eastern escarpments of the Andes, and in the Coastal and Amazon cordilleras. We also found that approximately 40% of the areas with the highest alpha and beta diversities have already disappeared due to deforestation and that only 30% is under protection in Ecuador. Thus, we propose 12 potential areas with a high priority of conservation mostly located in South Ecuador to improve the representativeness and complementarity of the current reserve network. Finally, the fifth chapter synthesizes the principal findings of this thesis highlighting the implications for conservation and suggesting potential areas to be preserved based on human pressure levels, remaining forest and alpha and beta diversity patterns of endemic plants.

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:

Tropische und subtropische Trockenwälder zählen zu den bedrohtesten tropischen Ökosystemen (MILES et al. 2006). Sie bieten für mehr Menschen Siedlungsraum als humide Waldgebiete und sind dadurch besonders stark dem menschlichen Einfluss ausgesetzt. Die vorliegende Arbeit beschäftigt sich mit der Entwicklung der Baumverjüngung in der tumbesischen Trockenwaldregion Südecuadors und Nordperus und dem Einfluss der dort üblichen Waldweide. In Reservaten in der Provinz Loja (Ecuador) und im angrenzenden Nationalpark Cerros de Amotape auf peruanischer Seite wurde in 288 Parzellen (je 4 m²) die Baumregeneration registriert. Die Parzellen waren im Trockenwald mit den Vegetationsformen halb-laubabwerfend und laubabwerfend verteilt; zudem wurde die Hälfte der Parzellen eingezäunt, um den Einfluss der Beweidung auf die Regeneration zu erfassen. Die Aufnahmen fanden in einem Zeitraum von zwei Jahren zweimal während der Trockenzeit und zweimal kurz nach der Regenzeit statt, beginnend mit einer Aufnahme in der Trockenzeit 2014. Es wurden die Individuen markiert und die Artnamen dokumentiert, Höhen und abgestorbene Individuen wurden aufgenommen. Aus den Daten wurden die Höhenzuwächse, die Mortalitätsraten sowie die Shannon- und Evennessindices berechnet. Insgesamt wurden 93 Arten aus 36 verschiedenen Familien aufgenommen. Die Ergebnisse zeigten für den Erhebungszeitraum eine höhere Individuenanzahl während der Regenzeit, die Anzahl der Arten war im Vergleich zur Trockenzeit jedoch geringer. Die Diversität unterschied sich nicht signifikant zwischen den Jahreszeiten. Bei Betrachtung der Aufnahmen stach das zweite Monitoring mit besonders hohen Individuenanzahlen heraus. Zwischen den Waldtypen gab es einen signifikanten Unterschied in der Höhe der Verjüngung, sowie im Höhenzuwachs von der dritten auf die vierte Aufnahme. Die offenen und eingezäunten Flächen unterschieden sich ebenso in den Höhen und dem Höhenzuwachs signifikant. Die Diversität unterschied sich mit und ohne Zäunung und bei den beiden Waldtypen nicht eindeutig. Die Ergebnisse können unter anderem durch die Struktur des Altbestandes, dessen Einfluss auf die Strahlung, die Temperatur und die Feuchtigkeit, und wiederum deren Einfluss auf die Verjüngung erklärt werden. Lokale und regionale floristische Diversität und Zusammenstellung, Wachstumsformen, Phänologie und Demographie der Verjüngungspflanzen sind in weiten Formen besonders abhängig von der Hydrologie. Beweidung andererseits lenkt die Ausbreitung Verbiss resistenterer Arten, welche durch Reduzierung der Weidetiere zur Walderhaltung beitragen können. Diese Studie soll als lokale Forschungsarbeit zum besseren Verständnis des Ökosystems Trockenwald beitragen. Die gesammelten Erkenntnisse sollen später der Entwicklung nachhaltiger Landnutzungssysteme dienen, welche den Fortbestand dieses gefährdeten Habitats sichern.