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. Irrespective 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 relatively 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 circumference fluctuations. Nighttime transpiration and hydrological type: In contrast to the isohydrically performing trees of the Ocotea group, the anisohydric trees showed considerable water vapour pressure deficit (VPD)-dependent nocturnal transpiration. Therefore, we conclude that nighttime ET at the forest level is mainly sourced by the tree species with anisohydric performance.
Ecosystem water regulation couples energy and water balance, depends on the integrity of the ecosystem, and responds to changes in climate. Changes in tree-water relationships in the biodiversity hotspot of the tropical Andes in southern Ecuador might be potentially observed at the level of individual trees, thus providing an efficient ecosystem monitoring method with applications in forest management and conservation at the tree and landscape levels. In this study, we combine area-average measurements from a laser scintillometer above the forest with optical satellite data at high spatial resolution to obtain area-wide evapotranspiration data. The processing of field data includes the calculation of energy storage in forest biomass and the partitioning of evapotranspiration into transpiration and evaporation. Satellite-based estimates are calibrated by using tower flux measurements and meteorological data within periods of humid and less-humid atmosphere. The annual evapotranspiration was 1316mm, of which 1086mm per year corresponds to the forest transpiration at the study site. Average values of 4.7 and 4.1mm d−1 per tree crown are observed under humid and less-humid atmospheric conditions, respectively, when applying high-resolution area-wide evapotranspiration in individual crown analysis. Approximately 24% of the observed crowns show a positive monthly change in ET, and 51% of the crowns show a significant change in the daily ET, which can be considered sensitive individuals concerning water relationships. The limitations in the area-wide evapotranspiration at the crown level can be explained by considering the spectral responses of the crown individuals. The presented method can be robustly deployed in the ecological monitoring of mountain forests.
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 Spiro theca 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. (C) 2016 Elsevier Ltd. All rights reserved.
The relationship between canopy-level evapotranspiration (ETSci) and leaf-level transpiration (T-leaf) as well as photosynthesis (P-leaf) for a homogeneous tropical montane pasture was analyzed over five days using a combination of methods involving a laser scintillometer and a porometer. Weather conditions ranged from overcast to sunny during the period of study. The gas exchange of the leaves of the dominant pasture grass Setaria sphacelata (transpiration vs. photosynthetic CO2 net uptake) was measured with a porometer and physiologically interpreted on the background of microclimate variables (photosynthetic active radiation (PAR) as proxy for total light intensity, temperature, water vapor deficit of the air) and soil moisture data. Water use efficiency (WUE, photosynthetic CO2 net uptake vs water loss by leaf transpiration) of the pasture was used to analyze the grass' range of response to the environmental variables of the research area. PAR and water vapor deficit of the air (VPD) appeared to be the determinant factors for T-leaf and ETSci. WUE for the Setaria sphacelata pasture ranged from 1.9 to 5.8 mu mol CO2 mmol(-1) H2O day(-1) and is particularly low during periods of high VPD combined with enhanced insolation during cloudless periods. ET measurements collected by the scintillometer demonstrated a strong correlation with water flux calculated using the Penman-Monteith approach (TPM) (r(2) = 0.95). Also, T-leaf measured with the porometer showed reasonable coincidence with the ET observations (r(2) = 0.78). Values of ETSci ranged from 2.26 to 4.96 mm day(-1) and T-leaf ranged from 0.83 to 2.41 mm day(-1), but only ETSci showed good correspondence with the available energy (net radiation). The lower correlation between T-leaf and canopy-level ETSci compared to that between ETSci and T-PM was tested against contaminations from the adjacent fetch area of the scintillometer path, but no effects were found. Likewise, soil water limitations of T-leaf could be ruled out. Therefore, different correlations of ETSci and T-leaf with the incoming energy and VPD may be traced back to a direct effect of the VPD on ET in contrast to its indirect effect on T-leaf which is additionally regulated by physiological processes in the leaf stomata.
We investigated the seasonal gas exchange patterns of three different functional types of tropical afromontane trees, namely Podocarpus falcatus (Thunb.) Mirb. (evergreen gymnosperm), Prunus africana (Hook. f.) Kalkm. (broad-leaf evergreen), and Croton macrostachyus Hochst. ex Del. (broad-leaf deciduous) which grow side-by-side in the Munessa forest, southern Central Ethiopia. The hypothesis is that the trees can make different use of the environmental conditions which change seasonally and also from year to year. These changes can be understood as fluctuating niches, the utilization of which allows coexistence through balanced carbon gain. In this study, light and moisture were considered the two main fluctuating niches. Porometry was used to measure the daily and seasonal responses of the leaves to environmental variables under dry and wet season conditions in the course of the climatically differing years 2009 and 2010. Correlations of the patterns of these responses were analyzed by regression analysis. From daily integrated rates of irradiation, photosynthesis and transpiration, light and water use efficiencies of photosynthetic carbon gain and their mutual interdependences were determined. Except an extended dry season in 2009 when C. macrostachyus had shed its foliage, all three species continued photosynthetic net CO2 uptake concomitant with transpiration at all seasons, although at varying rates. Ecophysiological performance of P. falcatus leaves is mainly light-driven and responds relatively little to a change in moisture conditions. Its carbon and water relations are striving for stability rather than flexibility. As of a typically subdominant species of the forest, P. africana leaves can efficiently use low light intensities but suffer from photoinhibition at full light. Their performance, showing more dynamic response to the environment than P. falcatus, P. africana appears driven more by moisture than by light. Compared to the two evergreens C. macrostachyus exhibited the highest flexibility in its leaves’ physiological responses to environmental conditions, in particular to the light climate which is additionally potentiated by the fast turnover of its foliage. This species optimizes its carbon gain during the wet season and during the early dry period when cloud cover is minimal. Our findings thus reveal that elasticity of the response to fluctuating environmental conditions is an additional aspect in the assessment of the utilization of temporally fluctuating niches by adult tree individuals.
Foliage dynamics of three functional tree types representing major components of the tropical montane evergreen forest in southern part of Central Ethiopia were compared. The species were Podocarpus falcatus (evergreen gymnosperm), Prunus africana (evergreen broadleaf), and Croton macrostachyus (facultative deciduous). The hypothesis examined is that in such tropical trees, endogenous control of foliage dynamics by the leaf life-spans (LLS) is largely dominant over external signals. Crown foliage turnover, leafiness of twigs, LLS, photosynthetic performance, respiration rate, specific leaf area, and relative growth rates of the stems were investigated. Foliage dynamics and leafiness of the twigs were monitored over 2 years while leaf traits were followed over 3 months. The degree of inter and intra-individual synchronization of foliage phenophases was examined to get an estimate of the contributions of endogenous and external signals to the dynamics of the foliages. Autoregression analysis indicated significant influence of the moisture regime on leaf sprouting of Croton and Podocarpus. During pronounced dry periods, new leaves were not developed. Analysis of phenological data using circular statistics revealed that in spite of strong inter-individual synchronization of leaf flush and fall (Podocarpus and Croton), the dynamics of individual parts of the crowns were less synchronized. LLS was independent of climate factors and it had substantial contribution to the control of foliage turnover. Moreover, examination of ecophysiological traits of developing leaves of the studied functional types showed differing patterns with LLS corroborating the ecophysiological characteristics. Although overlaid by fungal infestation, both the foliage and ecophysiological properties of Prunus resemble that of Podocarpus but the former exhibited a shorter LLS and slightly higher metabolic rates. Nevertheless, all species reacted positively to high moisture with respect to stem growth. In spite of largely differing weather conditions of the 2 years, direct competitive advantage of one of the species over the others could not be detected.
Plantations of exotic trees on areas where tropical forest has been clear-felled have been reported to facilitate regrowth of indigenous tree species. This so-called nurse-or shelter tree effect was investigated in a montane semihumid site in southern Ethiopia where plantations of Pinus patula and Eucalyptus saligna grow in close vicinity to the natural Podocarpus falcatus mixed forest. Physiological performance of P. falcatus saplings growing in the exotic and the natural forests was investigated over the year. Compared with the natural forest, photosynthetic carbon gain and growth of the saplings were significantly enhanced under Pinus patula while likewise retarded under Eucalyptus saligna. Diverging effects of the differently dense shelter canopies on the saplings could be traced to differences in the sub-canopy microclimates and particularly to the intensities and temporal distribution of photosynthetic active radiation (PAR). Moisture also played an important role for photosynthetic carbon gain: while the morning patterns of CO2 uptake were significantly correlated with the patterns of sunflecks, stomatal conductance was the determinant factor in the afternoon. Analysis of the photosynthetic efficiency of sunflecks revealed a lower quantum yield than the basic diffuse PAR intensity. Compared with a theoretically even distribution of the daily PAR, CO2 uptake under the real light climate was 70% under Pinus and in the natural forest, and 59% under Eucalyptus. Relating growth rates of Podocarpus saplings to photosynthesis the microclimate under Pinus was 2.5 times as effective as that in the natural forest and five times more effective than under Eucalyptus.
Munessa Forest is a mountain forest in south-eastern Ethiopia experiencing seasonal rainfall variation. We investigated seasonal cambial activity and dormancy from increment rates of four different tree species belonging to varying life forms, namely, evergreen native conifer (Podocarpus falcatus), evergreen introduced conifer (Pinus patula), evergreen broadleaved tree (Prunus africana) and deciduous broadleaved tree (Celtis africana). Measurements of stem radius fluctuations were registered with the help of high-resolution electronic dendrometers. Daily amplitudes of stem diameter variations and daily and monthly net growth rates were determined and related to climatic variables measured at local climate stations. Thin sections of wood collected with a microcorer every 3–6 weeks allowed a visual control of newly formed wood cells during consecutive time intervals. Lack of water availability during the long dry season induced cambial dormancy of 5–7 months depending on life forms. After the onset of the short rainy season, stem swelling started quite synchronously with a variation of only single days in all studied species. Evergreen tree species were able to initiate wood formation during the short rainy season, whereas growth in the deciduous broadleaved species started in the long rainy season. The acquired data provide a basis for delineating the species-specific growth boundaries and the duration of the cambial growing season.