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

Abstract:

Ecosystems worldwide are threatened by the increasing impact of land use and climate change. To protect their diversity and functionality, spatially explicit monitoring systems are needed. In remote areas, monitoring is difficult and recurrent field surveys are costly. By using Lidar or themore cost-effective and repetitive optical satellite data, remote sensing could provide proxies for habitat structure supporting measures for the conservation of biodiversity. Here we compared the explanatory power of both, airborne Lidar and optical satellite data in modeling the spatial distribution of biodiversity of birds across a complex tropical mountain forest ecosystem in southeastern Ecuador. Weused data fromfield surveys of birds and chose three measures as proxies for different aspects of diversity: (i) Shannon diversity as a measure of ?-diversity that also includes the relative abundance of species, (ii) phylodiversity as a first proxy for functional diversity, and (iii) community composition as a proxy for combined ?- and ?-diversity.We modeled these diversity estimates using partial least-square regression of Lidar and optical texturemetrics separately and compared themodels using a leave-one-out validated R2 and rootmean square error. Bird community informationwas best predicted by both remote sensing datasets, followed by Shannon diversity and phylodiversity. Our findings reveal a high potential of optical texture metrics for predicting Shannon diversity and ameasure of community composition, but not for modeling phylodiversity. Generalizing from the investigated tropicalmountain ecosystem,we conclude that texture information retrieved frommultispectral data of operational satellite systems could replace costly airborne laser-scanning formodeling certain aspects of biodiversity.

Abstract:

Understanding of evapotranspiration (ET) processes over Andean mountain environments is crucial, particularly due to the importance of these regions to deliver water-related ecosystem services. In this context, the detection of spatio-temporal changes in ET remains poorly investigated for specific Andean ecosystems, like the páramo. To overcome this lack of knowledge, we implemented the energy-balance model METRIC with Landsat 7 ETM+ and MODIS-Terra imagery for a páramo catchment. The implementation contemplated adjustments for complex terrain in order to obtain daily, monthly and annual ET maps (between 2013 and 2014). In addition, we compared our results to the global ET product MOD16. Finally, a rigorous validation of the outputs was conducted with residual ET from the water balance. ET retrievals from METRIC (Landsat-based) showed good agreement with the validation-related ET at monthly and annual steps (mean bias error <8 mm·month?1 and annual deviation <17%). However, METRIC (MODIS-based) outputs and the MOD16 product were revealed to be unsuitable for our study due to the low spatial resolution. At last, the plausibility of METRIC to obtain spatial ET retrievals using higher resolution satellite data is demonstrated, which constitutes the first contribution to the understanding of spatially-explicit ET over an alpine catchment in the neo-tropical Andes.

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:

Ecosystems worldwide are threatened by the increasing impact of land use and climate change. To protect their diversity and functionality, spatially explicit monitoring systems are needed. In remote areas, monitoring is difficult and recurrent field surveys are costly. By using Lidar or themore cost-effective and repetitive optical satellite data, remote sensing could provide proxies for habitat structure supporting measures for the conservation of biodiversity. Here we compared the explanatory power of both, airborne Lidar and optical satellite data in modeling the spatial distribution of biodiversity of birds across a complex tropical mountain forest ecosystem in southeastern Ecuador. Weused data fromfield surveys of birds and chose three measures as proxies for different aspects of diversity: (i) Shannon diversity as a measure of ?-diversity that also includes the relative abundance of species, (ii) phylodiversity as a first proxy for functional diversity, and (iii) community composition as a proxy for combined ?- and ?-diversity.We modeled these diversity estimates using partial least-square regression of Lidar and optical texturemetrics separately and compared themodels using a leave-one-out validated R2 and rootmean square error. Bird community informationwas best predicted by both remote sensing datasets, followed by Shannon diversity and phylodiversity. Our findings reveal a high potential of optical texture metrics for predicting Shannon diversity and ameasure of community composition, but not for modeling phylodiversity. Generalizing from the investigated tropical mountain ecosystem, we conclude that texture information retrieved frommultispectral data of operational satellite systems could replace costly airborne laser-scanning formodeling certain aspects of biodiversity.

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 theory of portfolio selection has often been applied to help improving decisions on the environment. The information required to apply this theory includes data on covariance of the uncertain returns between all combinations of the economic options and normally distributed returns. As it may be problematic to fulfil all data requirements and assumptions, the paper proposes a variant of robust portfolio optimization as an alternative. It considers future uncer-tainties in a non-stochastic fashion by means of possible deviations from the nominal return of land-use alternatives. The maximization of the economic return of the land-use portfolio is conditional to meeting an inclusive set of constraints. These demand that the same, whenever possible high, proportion of a required return threshold is achieved by means of the robust solution for each deviation scenario considered. The paper compares the land-use portfolios derived non-stochastically with portfolios generated by classical stochastic mean-variance op-timization. Based on data for eight agricultural crops typical for the Ecuadorian lowlands the results show that, depending on the deviation scenario assumed, the robust portfolios show a greater Shannon index compared to classical portfolios. For the same standard deviation of returns (derived through the classical sum of all covariances) the robust portfolios show no more than 2-5% loss in economic return in most cases. Occasionally, the loss has been higher, up to 20%. In this case the Shannon index was about 2.5 times higher compared with that of the conventional portfolio. The highly diverse portfolio achieved a much better protection against low relative performance. The results obtained show that the non-stochastic derivation of land-use portfolios is a good alternative to the classical stochastic model, whenever eco-nomic information is scarce.

Abstract:

Deforestation and fragmentation are major components of global change; both are contributing to the rapid loss of tropical forest area with important implications for ecosystem functioning and biodiversity conservation. The forests of South Ecuador are a biological 'hotspot' due to their high diversity and endemism levels. We examined the deforestation and fragmentation patterns in this area of high conservation value using aerial photographs and Aster satellite scenes. The registered annual deforestation rates of 0.75% (1976–1989) and 2.86% (1989–2008) for two consecutive survey periods, the decreasing mean patch size and the increasing isolation of the forest fragments show that the area is under severe threat. Approximately 46% of South Ecuador's original forest cover had been converted by 2008 into pastures and other anthropogenic land cover types. We found that deforestation is more intense at lower elevations (premontane evergreen forest and shrubland) and that the deforestation front currently moves in upslope direction. Improved awareness of the spatial extent, dynamics and patterns of deforestation and forest fragmentation is urgently needed in biologically diverse areas like South Ecuador.

Abstract:

With respect to sampling for regression-based digital soil mapping (DSM), the above all aim is to ensure that the spatial variability of the soil is well-captured without introducing any bias, while the design remains feasible according to operational constraints such as accessibility, man power and cost. Representativeness of the sample concerning the population to be sampled needs to be guaranteed in any regression-based modelling approach. Four selected sampling designs were adapted to show that basically any design may be optimised to represent the n-dimensional predictor space of a particular area, while selecting points is only permitted from a small accessible sub-area or from outside the area. Sampling efficiency may be evaluated based on the representation of the predictor space. However, not only each predictor’s probability function but also the interaction between predictors may have to be considered, to select a representative sample. Instead of sampling a previously un-sampled area with limited accessibility, the four sampling designs may also be used to subsample an existing dataset and, thereby, optimise a suboptimal dataset based on the predictor space of the area which shall be mapped by DSM.

Abstract:

From 10 selected tree species, Bursera graveolens and Maclura tinctoria exhibited distinct annual and crossdatable tree-rings. It was possible to synchronize individual tree-ring series and to establish two tree-ring chronologies of 203 and 87 years length, respectively. The characteristic ENSO frequency band is reflected in wavelet power spectra of both chronologies. Both species showa strong correlation between ringwidth and precipitation of thewet season (January–May). Strong El Niño events (1972, 1983 and 1998) lead to strong growth responses in the tree-ring chronologies, whereas ‘normal’ ENSO events do not trigger long-lasting growth responses. The first ring-width based wet-season precipitation reconstruction for the past 103 years was developed. Statistical and spatial correlation analysis verified the skills of the reconstructed precipitation which captures a great part of the Rainfall Index over the land area of Ecuador and the equatorial Pacific. Furthermore, teleconnections with central Pacific precipitation and SST patterns were found.

Abstract:

This paper presents a method to extrapolate wind-speed data and to calculate wind-speed and dynamic pressure maps for the complex topography of a mountain rainforest area in the tropical Andes of southeastern Ecuador. The spatial differentiation of dynamic wind pressure in this area is claimed to be a major determinant of the altitude of the tree-line ecotone and to affect the tree line’s physiognomy. The paper presents a hybrid method encompassing statistical data analysis using the Weibull distribution and a digital terrain analysis, taking topographical shelter effects into account. The method is used to derive mean and maximum wind-speed and dynamic pressure maps to reveal whether the tree-line ecotone is influenced by direct wind effects. On average, the tree-line ecotone on the eastern slopes shows a clear average depression of ~50 m. These slopes are affected by higher dynamic wind stress, so have a more disturbed canopy. These altered vegetation structures may be caused mainly by direct wind effects and to a smaller extent by indirect effects, such as high humidity.

Abstract:

In the megadiverse tropical mountain forest in the Andes of southern Ecuador, a global biodiversity hotspot, the use of fire to clear land for cattle ranching is leading to the invasion of an aggressive weed, the bracken fern, which is threatening diversity and the provisioning of ecosystem services. To find sustainable land use options adapted to the local situation, a profound knowledge of the long-term spatiotemporal patterns of land cover change and its drivers is necessary, but hitherto lacking. The complex topography and the high cloud frequency make the use of remote sensing in this area a challenge. To deal with these conditions, we pursued specific pre-processing steps before classifying five Landsat scenes from 1975 to 2001. Then, we quantified land cover changes and habitat fragmentation, and we investigated landscape changes in relation to key spatial elements (altitude, slope, and distance from roads). Good classification results were obtained with overall accuracies ranging from 94.5% to 98.5% and Kappa statistics between 0.75 and 0.98. Forest was strongly fragmented due to the rapid expansion of the arable frontier and the even more rapid invasion by bracken. Unexpectedly, more bracken-infested areas were converted to pastures than vice versa, a practice that could alleviate pressure on forests if promoted. Road proximity was the most important spatial element determining forest loss, while for bracken the altitudinal range conditioned the degree of invasion in deforested areas. The annual deforestation rate changed notably between periods: ~1.5% from 1975 to 1987, ~0.8% from 1987 to 2000, and finally a very high rate of ~7.5% between 2000 and 2001. We explained these inconstant rates through some specific interrelated local and national political and socioeconomic drivers, namely land use policies, credit and tenure incentives, demography, and in particular, a severe national economic and bank crisis.

Abstract:

In undisturbed tropical montane rainforests massive organic layers accommodate the majority of roots and only a small fraction of roots penetrate the mineral soil. We investigated the contribution of vegetation to slope stability in such environments by modifying a standard model for slope stability to include an organic layer with distinct mechanical properties. The importance of individual model parameters was evaluated using detailed measurements of soil and vegetation properties to reproduce the observed depth of 11 shallow landslides in the Andes of southern Ecuador. By distinguishing mineral soil, organic layer and aboveground biomass, it is shown that in this environment vegetation provides a destabilizing effect mainly due to its contribution to the mass of the organic layer (up to 973 t ha
1 under wet conditions). Sensitivity analysis shows that the destabilizing effect of the mass of soil and vegetation can only be effective on slopes steeper than 37.9. This situation applies to 36% of the study area. Thus, on the steep slopes of this megadiverse ecosystem, the mass of the growing forest promotes landsliding, which in turn promotes a new cycle of succession. This feedback mechanism is worth consideration in further investigations of the impact of landslides on plant diversity in similar environments.

Abstract:

An arbuscular mycorrhizal fungus (AMF) was isolated from a south Ecuadorian reforestation site in the vicinity of a pristine mountain rainforest ecosystem. Morphologically, it corresponded with the description of an AMF first described from Cuba as Gigaspora savannicola, before it was placed in Scutellospora. No living culture of this species has been available previously to allow detailed morphological review or DNA barcoding. After comparison of the morphology of the holotype and authenticated specimens with that of the Ecuadorian isolate, it was concluded that they are conspecific. A detailed redescription of the species is provided, including morphological characters not included in the original description, and an epitype is designated. To provide an extended DNA barcode for this AMF species, intraspecific variants of the near full-length small subunit rRNA gene (SSU) and a rDNA region comprising part of the SSU, the internal transcribed spacer (ITS) region including the 5.8S rRNA gene, and part of the large subunit rRNA gene (LSU) were sequenced. Phylotaxonomic analyses confirmed the classification in the genus Scutellospora sensu lato in a phylogenetic clade that, based on questionable evidence, was considered to circumscribe a distinct genus, Fuscutata. Recently, this genus was merged with Dentiscutata. Consequently, we move S. savannicola (=Fuscutata savannicola?=?Gigaspora savannicola) to the genus Dentiscutata (Gigasporaceae).

Abstract:

To reconstruct the environmental history including vegetation, fire and climate dynamics, from the Cajanuma valley area (3285 m elevation) in the Podocarpus National Park, southern Ecuador, we address the following major research question: (1) How did the mountain vegetation developed during the late Glacial and Holocene? (2) Did fire played an important control on the vegetation change and was it natural or of anthropogenic origin?. Palaeoenvironmental changes were investigated using multiple proxies such as pollen, spores, charcoal analyses and radiocarbon dating. Pollen data indicated that during the late Glacial and transition to the early Holocene (ca. 16 000–10 500 cal yr BP) herb páramo was the main vegetation type around the study area, while subpáramo and mountain rainforest were scarcely represented. The early and mid-Holocene (ca. 10 500 to 5600 cal yr BP) is marked by high abundance of páramo during the early Holocene followed by a slight expansion of mountain forest during the mid-Holocene. During the mid- to late Holocene (ca. 5600–1200 cal yr BP) there is a significant presence of páramo and subpáramo while Lower Mountain Forest decreased substantially, although, Upper Mountain Forest remained relatively stable during this period. The late Holocene, from ca. 1200 cal yr BP to present, was characterized by páramo; however, mountain forest and subpáramo presented significantly abundance compared to the previous periods. Fires became frequent since the late Holocene. The marked increased local and regional fire intensity during the wetter late Holocene strongly suggests that were of anthropogenic origin. During the late Glacial and early Holocene, the upper forest line was located at low elevations; but shifted slightly upslope to higher elevations during the mid-Holocene.

Abstract:

Late Pleistocene and Holocene vegetation, climate, and fire dynamics, as well as human impact, were studied in the El Cristal region, which is part of the Andean Depression in southeastern Ecuador. The sediment core from a small basin at 2056 m a.s.l. was analyzed by pollen and charcoal analysis and dated by six radiocarbon dates. The record indicates that during the late Pleistocene (ca. 19 750–12 500 cal yr BP) the Upper Mountain Rainforest (UMF), with some smaller areas of páramo and subpáramo were the main vegetation type in the study area. Additionally, there is evidence for Polylepis woodland in the region during this period, which does not exist in the area today. During the transition from the late Pleistocene to the early and mid-Holocene between ca. 12 500 to 3600 cal yr BP, there is a significant presence of mountain rainforest and stable proportions of subpáramo and páramo taxa. The early Holocene from ca. 11 500 to 7800 cal yr BP is characterized by UMF vegetation at the study site. Polylepis became more frequent and reached its maximum. During the mid-Holocene between 7800 to 3600 cal yr BP, the composition of the UMF changed. Polylepis decreased after ca. 4000 cal yr BP, probably due to higher fire frequency. During the mid- to late Holocene, human landscape dis- turbance is evident, especially between ca. 2000 and 1380 cal yr BP, with Asteraceae, Muehlenbeckia/Rumex, Osmunda and Huperzia all increasing in abundance. The late Holocene, from ca. 1380 cal yr BP, was characterized by an increased abundance of páramo taxa and a decrease in mountain rainforest taxa. The expansion of grassland during the late Holocene is interpreted as a result of an increase in human activity, which limited forest vegetation to isolated patches. However, between ca. 1200 and 900 cal yr BP the UMF expanded, probably due to forest recovery after the high frequency of fire. Between ca. 90

Abstract:

Late Holocene human impact and environmental changes were reconstructed from a sediment core of the Laguna Daniel Alvarez (2200 m asl) located on the outskirts of the city of Loja, southeastern Ecuador. Palaeoenvironmental changes were investigated by pollen, spore, algae and charcoal analysis in com- bination with X-Ray Fluorescence (XRF) scanning and element analysis of d13C, Total Organic Carbon (TOC), Total Nitrogen (TN) and radiocarbon dating. This multi-proxy study provides in detail the set- tlement history in the inner-Andean dry valley in southern Ecuador over the last ca. 1400 years. Between 630 and 1470 AD, Zea mays was intensively cultivated around the studied lake by the native Palta culture in the Loja region. After ca. 1470 AD, Z. mays cultivation collapsed, accompanied by an increase in fallow vegetation, such as Mimosa and Poaceae, probably as a result of the Inca invasion and occupation from 1463 to 1531 AD in southern Ecuador. After ca. 1570 AD, Amaranthaceae/Chenopo- diaceae markedly increased, re?ecting the beginning of the Spanish Conquest. In 1531 AD, Loja became Spanish and, during the ?rst ca. 100 years of the Spanish regime, Loja developed into the forti?ed capital of the province. In the 17th century, crop growing strongly declined due to the diminished indigenous population that probably suffered from new diseases introduced by the Spanish invaders. Pinus and Eucalyptus as well as Plantago lanceolata were introduced in the Loja region about 220 years ago.

Abstract:

Inverse parameter estimation of process-based models is a long-standing problem in many scientific disciplines. A key question for inverse parameter estimation is how to define the metric that quantifies how well model predictions fit to the data. This metric can be expressed by general cost or objective functions, but statistical inversion methods require a particular metric, the probability of observing the data given the model parameters, known as the likelihood. For technical and computational reasons, likelihoods for process-based stochastic models are usually based on general assumptions about variability in the observed data, and not on the stochasticity generated by the model. Only in recent years have new methods become available that allow the generation of likelihoods directly from stochastic simulations. Previous applications of these approximate Bayesian methods have concentrated on relatively simple models. Here, we report on the application of a simulation-based likelihood approximation for FORMIND, a parameter-rich individualbased model of tropical forest dynamics. We show that approximate Bayesian inference, based on a parametric likelihood approximation placed in a conventional Markov chain Monte Carlo (MCMC) sampler, performs well in retrieving known parameter values from virtual inventory data generated by the forest model. We analyze the results of the parameter estimation, examine its sensitivity to the choice and aggregation of model outputs and observed data (summary statistics), and demonstrate the application of this method by fitting the FORMIND model to field data from an Ecuadorian tropical forest. Finally, we discuss how this approach differs from approximate Bayesian computation (ABC), another method commonly used to generate simulation-based likelihood approximations. Our results demonstrate that simulation-based inference, which offers considerable conceptual advantages over more traditional methods for inverse parameter estimation, can be successfully applied to process-based models of high complexity. The methodology is particularly suitable for heterogeneous and complex data structures and can easily be adjusted to other model types, including most stochastic population and individual-based models. Our study therefore provides a blueprint for a fairly general approach to parameter estimation of stochastic process-based models.