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

Abstract:

This study addresses transpiration in a tropical evergreen mountain forest in the Ecuadorian Andes from the leaf to the stand level, with emphasis on nocturnal plant-water relations. The stand level: Evapotranspiration (ET) measured over 12 months with the Eddy-Covariance (ECov) technique proved as the major share (79%) of water received from precipitation. Irre- spective of the humid climate, the vegetation transpired day and night. On average, 15.3% of the total daily ET were due to nocturnal transpiration. Short spells of drought increased daily ET, mainly by enhanced nighttime transpiration. Following leaf transpiration rather than air temperature and atmospheric water vapor deficit, ET showed its maximum already in the morning hours. The tree level: Due to the humid climate, the total water consumption of trees was generally low. Nevertheless, xylem sap flux measurements separated the investigated tree species into a group showing relatively high and another one with low sap flux rates. The leaf level: Transpiration rates of Tapirira guianensis, a member of the high- flux-rate group, were more than twice those of Ocotea aciphylla, a representative of the group showing low sap flux rates. Representatives of the Tapirira group operated at a rela- tively high leaf water potential but with a considerable diurnal amplitude, while the leaves of the Ocotea group showed low water potential and small diurnal fluctuations. Overall, the Tapirira group performed anisohydrically and the Ocotea group isohydrically. Grouping of the tree species by their water relations complied with the extents of the diurnal stem circum- ference fluctuations. Nighttime transpiration and hydrological type: In contrast to the isohy- drically performing trees of the Ocotea group, the anisohydric trees showed considerable water vapour pressure deficit (VPD)-dependent nocturnal transpiration. Therefore, we con- clude that nighttime ET at the forest level is mainly sourced by the tree species with aniso- hydric performance.

Abstract:

Nature Geoscience, doi:10.1038/s41561-022-00911-8

Abstract:

Tropical mountain ecosystems are threatened by climate and land-use changes. Their diversity and complexity make projec- tions how they respond to environmental changes challenging. A suitable way are trait-based approaches, by distinguishing between response traits that determine the resistance of species to environmental changes and efect traits that are relevant for species’ interactions, biotic processes, and ecosystem functions. The combination of those approaches with land surface models (LSM) linking the functional community composition to ecosystem functions provides new ways to project the response of ecosystems to environmental changes. With the interdisciplinary project RESPECT, we propose a research framework that uses a trait-based response-efect-framework (REF) to quantify relationships between abiotic conditions, the diversity of functional traits in communities, and associated biotic processes, informing a biodiversity-LSM. We apply the framework to a megadiverse tropical mountain forest. We use a plot design along an elevation and a land-use gradient to collect data on abiotic drivers, functional traits, and biotic processes. We integrate these data to build the biodiversity- LSM and illustrate how to test the model. REF results show that aboveground biomass production is not directly related to changing climatic conditions, but indirectly through associated changes in functional traits. Herbivory is directly related to changing abiotic conditions. The biodiversity-LSM informed by local functional trait and soil data improved the simulation of biomass production substantially. We conclude that local data, also derived from previous projects (platform Ecuador), are key elements of the research framework. We specify essential datasets to apply this framework to other mountain ecosystems.

Abstract:

Ratios of stable oxygen isotopes in tree rings (δ18O) are a valuable proxy for reconstructing past climates. Such reconstructions allow us to gain better knowledge of climate dynamics under different (eg warmer) environmental conditions, which also forms the basis for effective risk management. The latter aspect is particularly relevant for our study site on the western flanks of the Andes in Southern Ecuador, since the region is frequently affected by droughts and heavy precipitation events during the rainy season (January to April), leading to enormous social and economic losses. In particular, we focus on precipitation amounts and moisture source regions as they are known to influence the δ18O signature of tree rings. Moisture source regions are based on 240 h backward trajectories that were calculated with the trajectory model LAGRANTO for the rainy seasons 2008 to 2017. A moisture source diagnostic was applied to the air parcel pathways. The resulting moisture source regions were analyzed by calculating composites based on precipitation amounts, season, and calendar year. The precipitation amounts were derived from data of a local Automatic Weather Station (AWS). The analysis confirms that our study site receives its moisture both, from the Atlantic and the Pacific Oceans. Heavy precipitation events are linked to higher moisture contributions from the Pacific, and local SST anomalies along the coast of Ecuador are of higher importance than those off the coast toward the central Pacific. Moreover, we identified increasing moisture contributions from the Pacific over the course of the rainy season. This change and also rain amount effects are detectable in preliminary data of δ18O variations in tree rings of Bursera graveolens. These signatures can be a starting point for investigating atmospheric and hydroclimatic processes, which trigger δ18O variations in tree rings, more extensively in future studies.

Abstract:

Under drought conditions, even tropical rainforests might turn from carbon sinks to sources, and tree species composition might be altered by increased mortality. We monitored stem diameter variations of 40 tree individuals with stem diameters above 10 cm belonging to eleven different tree genera and three tree life forms with high-resolution dendrometers from July 2007 to November 2010 and additionally March 2015 to December 2017 in a tropical mountain rainforest in South Ecuador, a biodiversity hotspot with more than 300 different tree species belonging to different functional types. Although our study area receives around 2200 mm of annual rainfall, dry spells occur regularly during so-called “Veranillo del Niño” (VdN) periods in October-November. In climate change scenarios, droughts are expected with higher frequency and intensity as today. We selected dry intervals with a minimum of four consecutive days to examine how different tree species respond to drought stress, raising the question if some species are better adapted to a possible higher frequency and increasing duration of dry periods. We analyzed the averaged species-specific stem shrinkage rates and recovery times during and after dry periods. The two deciduous broadleaved species Cedrela montana and Handroanthus chrysanthus showed the biggest stem shrinkage of up to 2 mm after 10 consecutive dry days. A comparison of daily circumference changes over 600 consecutive days revealed different drought responses between the families concerning the percentage of days with stem shrinkage/increment, ranging from 27.5 to 72.5% for Graffenrieda emarginata to 45–55% for Podocarpus oleifolius under same climate conditions. Moreover, we found great difference of recovery times after longer-lasting (i.e., eight to ten days) VdN drought events between the two evergreen broadleaved species Vismia cavanillesiana and Tapirira guianensis. While Vismia replenished to pre-VdN stem circumference after only 5 days, Tapirira needed 52 days on average to restore its circumference. Hence, a higher frequency of droughts might increase inter-species competition and species-specific mortality and might finally alter the species composition of the ecosystem.

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:

Tropical forests and the trees as their principal components have been investigated in detail. However, due to its complexity, their interactions, adaptations and response to climate variations require much more research. In this study, dendrochronological techniques were applied to evaluate the potential of tree-rings from tropical tree species as climate records. Two ecosystems with very distinct climate scenarios were selected from a dry and humid forest in southern Ecuador. A comparative analysis between these two forest types was performed by applying three dendrochronological methods. First, Tree Ring Width (TRW) measurements from tree species with distinct ring boundaries were dated to develop ring-width chronologies. Second, stable carbon isotopes (?13C) were measured from whole-wood and alpha-cellulose of dated annual tree-rings. Finally, concentrations of more than 23 chemical elements were determined from individual dated tree-rings after dissolving the wooden material in HNO3. The results showed the high potential of tropical tree species as climate archives, Bursera graveolens and Maclura tinctoria for the dry forest and Cedrela montana for the humid forest. Radial growth variations in tree species from the dry forest revealed a strong and reliable precipitation signal. Then, for these tropical regions, the first ring-width based wet-season precipitation reconstruction over the past century was developed, and spatial correlations unraveled a strong connection to the climatic conditions of the central Pacific precipitation and temperature variability. Interseries correlations of the TRW from the trees of the humid forest revealed a weak common signal. Stable carbon isotopes evidenced higher climate sensitivity than TRW measurements in the humid forest. However, to infer a reliable climate reconstruction from stable carbon isotopes, more ?13C time series were needed. ?13C values from whole-wood and alpha-cellulose reflected local and regional signals of precipitation and humidity. Meanwhile, nutrient concentration in the wood was higher in the dry forest, but common patterns and trends of nutrients were more distinct in the humid forest. For both study sites, two groups of nutrients with opposite radial distribution were identified (Group 1: Ca, Sr, Ba, Ga; and Group 2: K, P, Rb). In conclusion, TRW of tree species from the dry forest have a high paleoclimate potential, especially to reconstruct precipitation amounts in arid zones of southern Ecuador. Stable carbon isotopes constitute a promising tool to perform climatic reconstructions in both ecosystems. Finally, the valuable historical information of nutrient concentration evidenced in tree-rings opens promising ways to study tree growth dynamics especially in the humid forest.

Abstract:

The seasonal climate drivers of the carbon cycle in tropical forests remain poorly known, although these forests account for more carbon assimilation and storage than any other terrestrial ecosystem. Based on a unique combination of seasonal pan-tropical data sets from 89 experimental sites (68 include aboveground wood productivity measurements and 35 litter productivity measurements), their associated canopy photosynthetic capacity (enhanced vegetation index, EVI) and climate, we ask how carbon assimilation and aboveground allocation are related to climate seasonality in tropical forests and how they interact in the seasonal carbon cycle. We found that canopy photosynthetic capacity seasonality responds positively to precipitation when rainfall is < 2000mm/yr (water-limited forests) and to radiation otherwise (light-limited forests). On the other hand, independent of climate limitations, wood productivity and litterfall are driven by seasonal variation in precipitation and evapotranspiration, respectively. Consequently, light-limited forests present an asynchronism between canopy photosynthetic capacity and wood productivity. First-order control by precipitation likely indicates a decrease in tropical forest productivity in a drier climate in water-limited forest, and in current light-limited forest with future rainfall < 2000mm/yr.

Abstract:

We present the first multi-year long time series of local climate data in the seasonally dry tropical forest in Southern Ecuador and related growth dynamics of Loxopterygium huasango, a deciduous tree species. Local climate was investigated by installing an automatically weather station in 2007 and the daily tree growth variability was measured with high-resolution point dendrometers. The climatic impact on growth behaviour was evaluated. Hydro-climatic variables, like precipitation and relative humidity, were the most important factors for controlling tree growth. Changes in rainwater input affected radial increment rates and daily amplitudes of stem diameter variations within the study period from 2009 to 2013. El Nin˜o Southern Oscillation (ENSO) related variations of tropical Pacific Ocean sea surface temperatures influenced the trees’ increment rates. Average radial increments showed high inter-annual (up to 7.89 mm) and inter-individual (up to 3.88 mm) variations. Daily amplitudes of stem diameter variations differed strongly between the two extreme years 2009 (wet) and 2011 (dry). Contrary to 2009, the La Nin˜a drought in 2011 caused a rapid reduction of the daily amplitudes, indicating a total cessation (‘growth collapse’) of stem increment under ENSOrelated drought conditions and demonstrating the high impact of climatic extreme events on carbon sequestration of the dry tropical forest ecosystem.

Abstract:

Regarding woody plant responses on higher atmospheric inputs of the macronutrients nitrogen (N) and phosphorous (P) on tropical forests in the future, an adaptive modification of wood anatomical traits on the cellular level of woody plants is expected. As part of an interdisciplinary nutrient manipulation experiment (NUMEX) carried out in Southern Ecuador, we present here the first descriptive and quantitative wood anatomical analysis of the tropical evergreen tree species Alchornea lojaensis (Euphorbiaceae). We sampled branch wood of nine individual trees belonging to treatments with N fertilization, N+P fertilization, and a control group, respectively. Quantitative evaluations of eleven different vessel parameters were conducted. The results showed that this endemic tree species will be able to adapt well to the future effects of climate change and higher nutrient deposition. This was firstly implied by an increase in vessel diameter and consequently a higher theo. area-specific hydraulic conductivity with higher nutrient availability. Secondly, the percentage of small vessels (0–20 ?m diameter) strongly increased with fertilization. Thirdly, the vessel arrangement (solitary vessels vs. multiple vessel groupings) changed toward a lower percentage of solitary vessel fraction (VS), and concurrently toward a higher total vessel grouping index (VG) and a higher mean group size of non-solitary vessels (VM) after N and N+P addition. We conclude that higher nutrient availability of N and N+P triggered higher foliage amount and water demand, leading to higher cavitation risk in larger vessels. This is counteracted by a stronger grouping of vessels with smaller risk of cavitation to ensure water supply during drier periods that are expected to occur in higher frequency in the near future.

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:

Systematic investigations of the upper forest line (UFL) primarily concentrate on mid and high latitudes of the Northern Hemisphere, whereas studies of Neotropical UFLs are still fragmentary. This article outlines the extraordinary high tree diversity at the UFL within the Andean Depression and unravels the links between the comparatively low position of the local UFL, high tree-species diversity, and climate. On the basis of Gentry?s rapid inventory methodology for the tropics, vegetation sampling was conducted at 12 UFL sites, and local climate (temperature, wind, precipitation, and soil moisture) was investigated at six sites. Monotypic forests dominated by Polylepis were only found at the higher located margins of the Andean Depression while the lower situated core areas were characterized by a species-rich forest, which lacked the elsewhere dominant tree-species Polylepis. In total, a remarkably high tree-species number of 255 tree species of 40 different plant families was found. Beta-diversity was also high with more than two complete species turnovers. A non-linear relationship between the floristic similarity of the investigated study sites and elevation was detected. Temperatures at the investigated study sites clearly exceeded 5.5°C, the postulated threshold value for the upper tree growth limit in the tropics. Instead, quasi-permanent trade winds, high precipitation amounts, and high soil water contents affect the local position of the UFL in a negative way. Interestingly, most of the above-mentioned factors are also contributing to the high species richness. The result is a combination of a clearly marked upper forest line depression combined with an extraordinary forest line complexity, which was an almost unknown paradox.

Abstract:

We analyzed the stocks of Ca, Mg, P, and S in tree rings of Cedrela montana Moritz ex Turcz in order to evaluate the dynamics of element input to a lower montane rain forest in South Ecuador. Samples were taken by a Pressler drill from the living tree (n=5). Afterwards tree rings were separated and digested with concentrated HNO3 under pressure (Heinrichs et al.,1986). Concentrations of Ca and Mg were determined by atomic absorption spectroscopy (AAS), concentrations of P and S by inductively coupled plasma optical emission spectrometry (ICP-OES), respectively. Mean element concentrations in tree rings between 1957 and 2005 were 2.2 g kg-1 for Ca, 1.1 g kg-1 for P and S and 0.4 g kg-1 for Mg. Radial distribution of element concentrations was heterogeneous with exception of the tree rings close to the phloem, where concentrations were generally higher. The pattern of annual radial increment showed significant growth peaks related to strong La Niña events. Also concentrations of P and S peaked during the La Niña events of 1989, 1996, and 2000. For Ca and Mg a similar effect was observed with a time shift of one year. Highest Ca peaks during the monitored period were associated with the 1974 La Niña event, which was also the strongest event on the record. We conclude that tree rings are a suitable tool for recording element inputs of Ca, Mg, P, and S at lower montane rain forests of South Ecuador. Furthermore we found element inputs and growth signals linked to the ENSO cycle.

Abstract:

Stem diameter increments of the broadleaved deciduous tree species Tabebuia chrysantha were measured with high-resolution dendrometers in a tropical lower montane forest and in a dry forest in southern Ecuador, the latter showing a distinct dry season. Those analyses were complemented by wood anatomical studies on regularly collected microcores to determine the season of active cambial growth and the time of formation of annual growth boundaries. The length of the cambial active period varied between 3 and 7 months at the tropical lower montane forest and 2 and 4 months in the dry forest, respectively. During dry days, amplitudes of daily stem diameter variations correlated with vapour pressure deficit. During October and November, inter-annual climate variations may lead to dry and sunny conditions in the tropical lower montane forest, causing water deficit and stem diameter shrinkage in T. chrysantha. The results of the climate– growth analysis show a positive relationship between tree growth and rainfall as well as vapour pressure deficit in certain periods of the year, indicating that rainfall plays a major role for tree growth.

Abstract:

Cedrela montana is a deciduous broad-leaved tree species growing in the humid mountain rainforests of southern Ecuador. High-resolution dendrometer data indicate a regular seasonal growth rhythm with cambial activity during January to April. Amplitudes of daily radial stem diameter variations are correlated with the amount of the maximum daily vapour pressure deficit. During humid periods, daily stem diameter variations are considerably smaller than during drier periods. This indicates that cambial activity is limited by available moisture even in such a very humid mountain climate. Wood anatomical studies on microcores show the formation of a marginal parenchyma band at the beginning of the growth period. This parenchyma band can be used to delineate annual growth rings. We were able to establish the first ring-width chronology from Cedrela montana which covers the time until 1840. However, the chronology is presently statistically robust back to 1910 only. Correlation functions calculated with NCEP/NCAR data indicate a significantly positive relationship of tree growth with temperatures during the growth period during January to April. However, only 8% of the growth variance is explained by this climatic factor. In the future, this relationship may be useful to reconstruct past temperature conditions of the study area.

Abstract:

Abstract. Available proxy records witnessing palaeoclimate of the tropical Andes are comparably scarce. Major implications of palaeoclimate development in the humid and arid parts of the Andes are briefly summarized. The long-term behaviour of ENSO has general significance for the climatic history of the Andes due to its impact on regional circulation patterns and precipitation regimes, therefore ENSO history derived from non-Andean palaeo-records is highlighted. Methodological constraints of the chronological precision and the palaeoclimatic interpretation of records derived from different natural archives, such as glacier sediments and ice cores, lake sediments and palaeo-wetlands, pollen profiles and tree rings are addressed and complementary results concerning former climatic conditions are discussed in terms of possible implications of former atmospheric circulation patterns and main climatic forcing factors. During the last years, increasing tree-ring information is getting available from the tropical Andes, providing high-resolution climate-sensitive records covering the past centuries for the study of climate variability.