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

Abstract:

Understanding the response of net nitrogen (N) mineralization to climate and land-use change is important to predict the effect of environmental change on the biodiversity and nutrient supply of the tropical montane forest in Ecuador. Slow mineralization or microbial immobilization may limit the availability of N, although organic N stocks in soils are high. To determine the roles of land use and climate for N mineralization in the mineral topsoil, field incubations over 31 days were conducted with the help of soil-filled PVC cylinders in forest and pasture soils along a land-use and elevation gradient from 1000-3000 m above sea level (a.s.l.). The experiment made use of a threefold replicated, full-factorial design with two land-use types and three elevations. The incubation cylinders were closed both at the bottom and the top and therefore only permitted horizontal water and element fluxes through lateral slits. Start and end concentration of NH4-N and NO3-N were determined in 1 M KCl extracts to calculate net N mineralization as the sum of ammonification and nitrification rates. Ammonification was significantly higher under pasture than under forest, except at 3000 m a.s.l., where the highest ammonification rates were detected on the forest plots. Nitrification was, in contrast, significantly higher under forest than under pasture, with highest nitrification rates at 2000 m a.s.l. Although the climate was wetter and cooler at higher elevations, mean N mineralization in the mineral soil on the forest sites significantly increased with elevation. On pastures, N mineralization was not significantly related with elevation, which may be explained by soil management and farming intensity. The slope of the regression line of δ13C values on soil organic C concentrations in 10-cm soil layers revealed a close relationship with the N mineralization rates, indicating that the short-term field incubations reflected the long-term C mineralization regime. The shift to higher δ13C values with increasing depth of the soil profile was related to N turnover and could thus serve as predictor of N mineralization rates on the forest sites, but not on the pastures, where the vertical distribution of the δ13C values was altered by the input of organic matter from C4 grasses after land-use change. Furthermore, the relationship between the slope of the regression line of the enrichment of δ13C values on soil depth and N mineralization measured by in-situ incubation depended on the parent material. The topsoils developed from granodiorite showed lower N mineralization rates than those developed from phyllite and meta-sandstone. Different soil texture, rooting depth, nutrient availability, and different organic layers may provide possible explanations for the observed differences.

Abstract:

With increasing elevation, trees in tropical montane forests have to invest larger frac-tions of their resources into their fine roots in order to compensate for increasingly unfavorable soil conditions. It is unclear how elevation and related edaphic changes influence the variability in tree fine root traits and belowground functional diversity. We measured six fine root traits related to resource acquisition on absorptive fine roots of 288 trees from 145 species along an elevational gradient from 1000 m to 3000 m a.s.l. in tropical montane forests of the Ecuadorian Andes. We analyzed trait relation-ships with elevation and soil nutrient availability, and tested whether root functional diversity varied along these gradients. Fine roots at higher elevations and at more nutrient-poor sites were thicker, had higher tissue densities, and lower specific root length and nutrient concentrations than at lower elevations. These trends were diluted by the coexistence of tree species with a broad range of different root traits within communities particularly towards lower elevations, where root functional diversity was significantly higher. We conclude that nutrient limitation and potentially further adverse conditions at higher elevations are strong environmental filters that lead to trait convergence towards a conservative resource use strategy, whereas different trait syndromes are equally successful at lower elevations.

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:

Theory predicts positive effects of species richness on the productivity of plant communities through complementary resource use and facilitative interactions between species. Results from manipulative experiments with tropical tree species indicate a positive diversity–productivity relationship (DPR), but the existing evidence from natural forests is scarce and contradictory. We studied forest aboveground productivity in more than 80 humid tropical montane oldgrowth forests in two highly diverse Andean regions with large geological and topographic heterogeneity and related productivity to tree diversity and climatic, edaphic and stand structural factors with a likely influence on productivity. Main determinants of wood production in the perhumid study regions were elevation (as a proxy for temperature), soil nutrient (N, P and base cation) availability and forest structural parameters (wood specific gravity, aboveground biomass). Tree diversity had only a small positive influence on productivity, even though tree species numbers varied largely (6–27 species per 0.04 ha). We conclude that the productivity of highly diverse Neotropical montane forests is primarily controlled by thermal and edaphic factors and stand structural properties, while tree diversity is of minor importance.

Abstract:

Litterfall is the most easily and most frequently measured part of net primary productivity (NPP) of forests. It has been shown that litterfall accounts for about one third of total NPP and thus serves as a proxy for the total productivity of forests. Moreover, litterfall carries nutrients from the forest canopy to the soil and therefore is also a major vector of nutrient cycling. I reviewed the published literature about litterfall rates and chemical properties in Andean forests and found reports from 44 forest sites, which I evaluated together with unpublished data from 12 sites in a lower montane forest in Ecuador. I found many more reports from tropical (52 sites) than from temperate Andean forests (4 sites). In the humid tropical north Andes, litterfall showed a hump-shaped elevational distribution. It increased from premontane to lower montane forests and decreased to upper montane forests. The tropical lower montane forest had a similar productivity than tropical lowland forests. The litterfall of the temperate southern beech forests was similar to that of the tropical upper montane forests. The C/N and C/P ratios of litterfall decreased with increasing elevation, while the N/P ratios were not correlated with elevation. This illustrates that the forests become increasingly nutrient efficient with increasing elevation, while there is no indication of a general change in the kind of nutrient limitation. There was a negative correlation between litterfall and soil organic layer thickness (r = −0.61, p < 0.001) illustrating that the organic matter input via litterfall is a less important driver of organic matter accumulation on top of the mineral soil than other, mainly abiotic properties including temperature and soil waterlogging. My evaluation suggests that there are systematic relationships between abiotic conditions and litterfall, which could be used to predict litterfall in the Andes. However, the data coverage particularly of the southern Andes (Bolivia, Chile, Argentina), the Andean dry forests, and the widespread tree plantations is poor. The observed elevational influence on litterfall in the humid tropical Andes suggests that the forest productivity will likely respond to climate change driving the vegetation belts to higher elevation with an unknown overall effect on C sequestration of these forests.

Abstract:

Tropical forests harbor diverse ecological communities of plants and animals that are organized in complex interaction networks. The diversity and structure of plant–animal interaction networks may change along elevational gradients and in response to human-induced habitat fragmentation. While previous studies have analyzed the effects of elevation and forest fragmentation on species interaction networks in isolation, to our knowledge no study has investigated whether the effects of forest fragmentation on species interactions may differ along elevational gradients. In this study, we analyzed main and interaction effects of elevation and forest fragmentation on plant–frugivore interaction networks at plant and bird species level. Over a period spanning two years, we recorded plant–frugivore interactions at three elevations (1000, 2000 and 3000 m a.s.l.) and in two habitat types (continuous and fragmented forest) in tropical montane forests in southern Ecuador. We found a consistent effect of elevation on the structure of plant–frugivore networks. We observed a decrease in the number of effective bird partners of plants and, thus, a decline in the redundancy of bird species with increasing elevation. Furthermore, bird specialization on specific plant partners increased towards high elevations. Fragmentation had a relatively weak effect on the interaction networks for both plant and bird species, but resulted in a significant increase in bird specialization in fragmented forests at high elevations. Our results indicate that forest fragmentation may have stronger effects on plant–frugivore interaction networks at high compared to low elevations because bird species richness declined more steeply towards high elevations than plant species richness. We conclude that conservation efforts should prioritize the maintenance of consumer diversity, for instance by maintaining stretches of continuous forest. This applies in particular to species-poor communities, such as those at high elevations, as the ecological processes in these communities seem most sensitive towards forest fragmentation.

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:

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:

Ant assemblages are sensitive to abiotic changes in the environment, therefore they are widely used as indicators of environmental changes. Previous studies demonstrated that abiotic changes with elevation and increased anthropogenic disturbance not only reduce species richness of ant assemblages, but also modify their trophic composition and nutrient use. In tropical ecosystems where nutrient availability may vary between dry and wet seasons, seasonal or interactive effects might play an important but still neglected role in shaping these patterns. Here I used standardized bait experiments in natural and disturbed sites along an elevation gradient in a tropical montane rainforest during the wet and dry season. In order to analyze the single and interactive effects of elevation, disturbance and season on species richness, nutrient use and trophic composition of ants, I used linear mixed effect models. Additionally, I used principal component analysis (PCA) to assess whether morphological traits of ants are linked to their nutrient use. Species richness decreased monotonically along the elevation gradient, with a stronger decline in the dry season. Forest disturbance had no significant effect on species richness. The relative use of most nutrients decreased with increasing elevation. Forest disturbance only affected the relative use of lipids by decreasing it compared to natural forests. However, my results revealed complex interactive effects of elevation, disturbance and season on species richness and the use of nutrients by ant assemblages. Furthermore, I found a shift from predominantly omnivore species to more predatory species with increasing elevation. PCA revealed a preference of lipid baits by species with morphological traits associated with predatory taxa. My findings highlight the importance of seasonality and mixed effects on the composition of ant assemblages and their nutrient use in a tropical montane forest. Additionally, these results highlight the value of disturbed forests within my study area, since they support similar species richness and trophic composition of ant assemblages compared to natural forests.

Abstract:

Phylogenetic niche conservatism (PNC) is the tendency of species within a clade to retain ancestral traits and to persist in their primary ecological niches on geological time scales. It links evolutionary and ecological processes and has been hypothesized to explain patterns of species richness and the composition of species assemblages. Decreasing patterns of species richness along latitudinal gradients were often explained by the combination of ancient tropical climates, trait retention of tropical lineages and environmental filtering. PNC also predicts decreasing phylodiversity and family age with decreasing tropicality and has been invoked to explain these patterns along climatic gradients across latitudinal as well as elevational gradients. However, recent studies on tree assemblages along latitudinal and elevational gradients in South America found patterns contradicting the PNC framework. Our study aims to shed light on these contradictions using three different metrics of the phylogenetic composition that form a gradient from recent evolutionary history to deep phylogenetic relationships. We analyzed the relationships between elevation and taxonomic species richness, phylodiversity and family age of tree assemblages in Andean rainforests in Ecuador. In contrast to predictions of the PNC we found no associations of elevation with species richness of trees and increasing clade level phylodiversity and family age of the tree assemblages with elevation. Interestingly, we found that patterns of phylodiversity across the studied elevation gradient depended especially on the deep nodes in the phylogeny. We therefore suggest that the dispersal of evolutionarily old plant lineages with extra-tropical origins influences the recent composition of tree assemblages in the Andes. Further studies spanning broader ecological gradients and using better resolved phylogenies to estimate family and species ages are needed to obtain a deeper mechanistic understanding of the processes that drive the assembly of tree communities along elevational gradients.