Journal of Hydrology 237 (2000) 40–57 www.elsevier.com/locate/jhydrol Gross rainfall and its partitioning into throughfall, stemflow and evaporation of intercepted water in four forest ecosystems in western Amazonia C. Tobón Marin a,b,*, W. Bouten a, J. Sevink a a Fysisch Geografisch Bodemkundig Laboratorium, University of Amsterdam, Nieuwe Achtergracht 166, 1018 WV Amsterdam, The Netherlands b The Tropenbos Foundation, Colombia Received 29 July 1999; revised 4 May 2000; accepted 4 July 2000 Abstract The partitioning of gross rainfall into throughfall, stemflow and evaporation of intercepted rainfall was studied in four forest ecosystems in the Middle Caquetá, Colombian Amazonia. Data on climate was collected automatically on an hourly basis during a five-year period. Weekly measurements of rainfall, throughfall and stemflow were carried out during a period of two years, while daily measurements, on an event basis, were carried out during two subsequent years. Throughfall, stemflow and evaporation in each forest were checked for correlations with gross rainfall characteristics, canopy gap fraction, tree crown area and bark texture. Canopy gap fraction differed between forests, ranging from 9% on the flood plain to 17% on the Tertiary sedimentary plain. Rainfall was rather evenly distributed over the year, with one dry period from December to February. 92% of the rain fell in single showers of less than 30 mm and most of the storms (56%) fell in less than one hour, during the afternoon or early night. Throughfall ranged from 82 to 87% of gross rainfall in the forests studied and varied with gross rainfall in all forests. It depended on the amounts and characteristics of rainfall, but differences in throughfall among forests, when comparing similar rainfall events, clearly indicated that throughfall also depends on forest structure. Stemflow contributed little to net precipitation (on average 1.1% of gross rainfall in all forests) and showed a power relation with gross rainfall. Correlations between stemflow per tree, projected crown area and bark texture were very poor as indicated by the low coefficients of determination. Evaporation during rainfall events exhibited a linear relation with rainfall duration and the ratio of evaporation over gross rainfall increased with forest cover (1-gap fraction) in the forests studied. The structure of the forests seemed to vary considerably and given its influence on rainfall partitioning it may explain both differences and similarities between results from this study and those from most other studies within Amazonia. 䉷 2000 Elsevier Science B.V. All rights reserved. Keywords: Amazonia; Tropical rain forest; Throughfall; Stemflow; Evaporation; Forest structure 1. Introduction The Amazonian rain forest seems to play an important role in the regulation of regional and global * Corresponding author. Tel.: ⫹31-205257442; fax: ⫹31205257431. E-mail address: [email protected] (C. Tobón Marin). climate (Salati and Vose, 1984). Using a modelling approach, Eltahir and Bras (1993) concluded that the atmosphere in the Amazon basin is an open system and that the net input of atmospheric moisture into the basin is about 32%, about 68% of the gross input leaving the basin. They also found that the recycling ratio of the Amazon basin is about 25–35%. This ratio differs from those found by others, which were based 0022-1694/00/$ - see front matter 䉷 2000 Elsevier Science B.V. All rights reserved. PII: S0022-1694(00 )00 301-2 C. Tobón Marin et al. / Journal of Hydrology 237 (2000) 40–57 on the erroneous hypothesis that the atmosphere over the Amazon basin is a closed system (Molion, 1975; Lettau et al., 1979; Salati et al., 1979). Eltahir and Bras (1993) furthermore concluded that deforestation of Amazonia would increase the surface temperature and decrease the heating of the upper troposphere, which would result in a reduction of precipitation. The foregoing illustrates that recent concern about tropical rain forest deforestation focuses on its impact on climate at regional and global scale (e.g. The Anglo Brazilian Amazonia Climate observation study ABRACOS, Gash et al., 1996; the Large Scale Biosphere–Atmosphere experiment in Amazonia, Nobre et al., 1996; Eltahir and Bras, 1993). Current research concentrates on global circulation models and on those parameters and fluxes that play a role in the global climate (e.g. Gash et al., 1996). Nevertheless, it is often argued that additional work is needed to explain and improve model predictions for impacts of deforestation (Bruijnzeel, 1990; Nobre et al., 1991). Evidently, local hydrological studies on undisturbed mature rain forests would provide base level information on initial conditions that might allow for an evaluation of the presumed influence of deforestation on regional and global climate. This paper concerns such local study. In the Middle Caquetá (Colombian Amazonia), the structure and species composition of the forest vary considerably between the different landscape units (Duivenvoorden and Lips, 1995; Londoño, 1993). This causes equally large variations in temporal and spatial patterns of water fluxes in these units, leading to local differences in water and nutrient stocks in the various forest compartments (Vitousek and Denslow, 1986; Tanner, 1985). Thus, the partitioning of rainfall into throughfall and stemflow leads not only to a more diffuse input of water into the forest floor, but also to local concentration around the base of tree stems, which is known to induce spatial variability in soil properties and soil moisture conditions (Waidi et al., 1992). In tropical forests, the abundance of epiphytes, climbers and aerial roots renders this partitioning much more complicated than in temperate forests (Longman and Jenı́k, 1990). It is well understood that in forested areas generally total evaporation is larger than in areas with shorter vegetation (e.g. grass) mainly due to the larger interception by the forest canopy (Bosch and Hewlett, 41 1982), which has been related to the large aerodynamic conductance of forest (Stewart, 1977). Additionally, there has been an increasing awareness that evaporation of intercepted rainfall has to be investigated separate from transpiration, especially in very humid areas (Hutjes et al., 1990; Shuttleworth and Calder, 1979). Furthermore, most studies on forest interception showed that this interception is closely related to gross rainfall amounts and characteristics. However, the influence of forest structure on interception is poorly known. Most rainfall interception and water balance studies in Amazonia were executed in Brazil (Ubarana, 1996; Leopoldo et al., 1995; Lesak, 1993; Lloyd and Marques, 1988; Shuttleworth, 1988) and only a few in other parts of Amazonia (Hölscher et al., 1997; Jetten, 1996; Wright et al., 1992; Jordan, 1978). Thus, in Colombian Amazonia, which represents one of the most humid areas within the basin, very little attention has been paid to the hydrology of forest ecosystems and to the effects of forest structure on water dynamics. This paper concerns a study designed to address this lack of knowledge by measuring rainfall and its partitioning after entering the canopy in four undisturbed rain forests in the Middle Caquetá, Colombian Amazonia. It focuses on the analysis of long-term hydrological measurements of rainfall, throughfall, stemflow, the resultant evaporation and the related structure of these forests. 2. The study area The study area is in Peña Roja (Nonuya Indian community) near Araracuara, Middle Caquetá, Colombia, (0⬚ 37 0 and 1⬚ 24 0 S, 72⬚ 23 0 and 70⬚ 43 0 W; Fig. 1). Climate is classified as equatorial superhumid Afi (Köppen, 1936). The research sites are permanent undisturbed forest plots, used by the Tropenbos Foundation for its research. They lie approximately 200–250 metres above sea level and form a sequence from the lower terrace of the River Caquetá to the Tertiary sedimentary plain. Based on data from the manual meteorological station at Araracuara (IDEAM), average annual rainfall in the area is about 3100, April being the wettest month and January the driest. 42 C. Tobón Marin et al. / Journal of Hydrology 237 (2000) 40–57 Fig. 1. Location of the research sites in the various land units in the Middle Caquetá, Colombian Amazonia. Colombian Amazonia comprises 403,000 km 2 and the major part of this area is covered by mature rain forests, classified as ombrophilous tropical forest (Duivenvoorden, 1995). The research plots are located in the four main land units in the area: the Tertiary sedimentary plain, the upland terraces of the River Caquetá (high and low terraces) and the flood plain. The vegetation is very rich in species C. Tobón Marin et al. / Journal of Hydrology 237 (2000) 40–57 (Duivenvoorden and Lips, 1995; Londoño, 1993) and is typical for undisturbed mature forests in the western part of the Amazon basin. The canopy reaches about 25–30 m above the forest floor with some emergent trees reaching up to 45 m in the rarely inundated flood plain. There are three to four canopy layers, but large tree crowns in the upper canopy form the bulk of the vegetation. Lower canopies contribute far less to the forest cover. Small palms reaching a height of 2–4 m constitute the lowest layer. The land units differ in the total standing biomass, species diversity and tree density (Duivenvoorden and Lips, 1995; Londoño, 1993). Other differences between plots pertain to the structure of the forest canopy (canopy cover) and the contribution of epiphytes, climbers and aerial roots. A more detailed description and vegetation classification of the research sites is given by Duivenvoorden and Lips (1995); and Londoño (1993). 3. Materials and methods The areas for the present study were selected as being representative for the natural vegetation in the main land units from this part of the Amazon basin. Three subplots were selected in the Tertiary sedimentary plain (SP) and two subplots in the high terrace (HT), the low terrace (LT) and the flood plain (FP), respectively, to measure gross rainfall above the forest canopy, throughfall and stemflow (Fig. 1). In 1992, approximately 3 km from the plots, in an open area of about 20 hectare (within an Indian community village), an automatic weather station (AWS) was installed to measure gross rainfall, temperature, air humidity, incoming radiation, wind speed, wind direction and Class A pan evaporation. Parameters were measured and recorded every 30 s using a datalogger (CR10 Campbell Scientific Instruments), which additionally recorded means or totals every 20 min. Rainfall in the open area was measured by a tipping bucket raingauge with a resolution of 0.2 mm providing information on the number and duration of showers and on total rainfall. Gross rainfall in each plot was measured in two ways: (1) automatic measurements with one tipping bucket installed in the top of an emergent tree crown after clearing all 43 branches; (2) manual measurements with two raingauges per subplot suspended from cords attached to two emergent trees in small gaps within the forest. Throughfall was measured in the same rainfall subplots using 20 collectors per subplot, randomly located in an area of 50 by 20 m (1000 m 2). Evaporation from the collectors was avoided by using an internal plastic tube running from the funnels to the bottom of the collectors. To allow for direct correlation, all funnels for gross rainfall above the forest canopy and for throughfall had an orifice of 298.6 cm 2. Throughfall and forest rainfall collectors were calibrated against standard raingauges in the open. Because of the large variability in throughfall due to the forest structure (Jetten, 1996; Ford and Deans, 1978) many readings are needed to study forest interception. However, when using average values, moving the collectors has a positive effect by reducing the standard error of estimations (Lloyd and Marques, 1988). Therefore, each month (after five measurements) collectors were randomly relocated within all subplots during the period of ‘single event measurements’, i.e. between 1996–1997. Stemflow was measured for 15 randomly selected trees in each subplot. Collars, constructed from 8 mm thick black polyethylene plastic, were sealed to the stems in an upward spiral pattern and the water diverted into bottle gauges on the forest floor. The opening of each collar extended only about 2–3 cm from the trunk to avoid drips from the branches or leaves being collected by the collars. The amount of water, which drops in a diffuse pattern around roughbarked trees rather than adhering to and flowing down the trunks, was considered throughfall. For practical reasons, only trees with diameter larger than 10 cm were selected for stemflow measurements. Where palm trees were present in the subplots, depending on their frequency of occurrence one or two of palms per subplot were randomly selected. In the SP and HT plots, two palms were selected, in the LT four palms in total and in the FP three palms. Stemflow measurements were rotated once by installing collars around stems of new trees within the same subplots. Horizontally or downwards inclined branches of trees may not direct intercepted rainfall to the centre of the tree to be drained as stemflow. Therefore, the flat area of the tree crown was mapped by means of 44 C. Tobón Marin et al. / Journal of Hydrology 237 (2000) 40–57 vertical projections from the edge of outstanding upward branches to the forest floor. At least six projection lines were drawn for each tree. The crown area was found by integrating measured areas of each triangle. Stemflow is expressed as millilitre of water coming in over the horizontal area of the tree crown and flowing down along the tree trunk. Thus, each tree was considered as a single catchment area. Estimates for stemflow at subplot level were obtained by multiplying the average stemflow per tree of a specific diameter by the total number of trees with approximately the same diameter within the subplot. Data on sampled trees with a diameter of about 10 cm were used to extrapolate the information to trees with smaller diameter. Average stemflow per subplot was related to gross rainfall measured in the same subplot to determine the percentage of rainfall coming to the forest floor as stemflow. Stemflow amounts from each rainfall event (ml) and from individual trees were correlated with the measured projected area of the central crown of the tree concerned and with some tree characteristics such as trunk surface area and bark texture. Consequently, the periphery of the trunks or surface area was determined by measuring the tree trunk circumference at breast height and the height of the tree trunk to the first set of divergent branches. Trunk area was determined by considering the tree trunk as being rectangular and applying a correction factor to account for the conical form of tree. Bark texture was classified according to the roughness of the bark in a range from smooth to very fibrous. Per plot, 40 hemispherical photographs were taken to estimate the gap fraction of the forest canopy in the subplots where throughfall collectors were installed. The photographic camera was installed horizontally at the same height as the throughfall collectors. The black and white photographs were digitised with a scanner and analysed with the Hemiphot program (Ter Stegee, 1994), which calculates the gap fraction of a forest and roughly estimates the LAI from the forest cover. Although the photographs were taken under covered sky with diffuse sun light, they were corrected for the reflected light from leaves, branches and trunks. It is assumed that the mean fraction of white pixels in the photographs gives the best estimate of the gap fraction. Most methods for the determination of the water storage capacity of the forest canopy (C), which have been presented thusfar (Klaassen et al., 1998), mainly differ in the way of accounting for the drainage before the forest is completely saturated and the gradual saturation of forest layers with continuously proceeding evaporation. In the present study, C is the intercept of the regression of the estimated evaporation versus gross rainfall. Therefore, only single, highintensity rainfall events of short duration (less than one hour) were used. Late afternoon and night rainfalls were preferred, representing rainfalls under conditions of a low moisture deficit. In total, 30–40 events were selected for each ecosystem. Free throughfall is considered as an important parameter, among others, in studies on nutrient cycling, since it may represent the fraction of throughfall which is not involved in washing out dry deposition, exudates and released nutrients from leaves, branches and trunks. Free throughfall in the forests may be taken as the gap fraction (ps). Therefore, it was estimated for each forest from the set of digitised and scanned black and white photographs taken in each subplot and from the regression coefficient of throughfall versus gross rainfall (pt), using data from small storms, which were insufficient to saturate the forest canopy (Gash and Morton, 1978). About 14 small storms were selected for each forest to determine the value of pt. Manual measurements of rainfall, throughfall and stemflow were carried out on weekly basis from December 1993 to February 1996, without moving collectors. During these years, some measurements were carried out on event basis during periods with limited or no rainfall (mostly in the dry season). Daily measurements were performed from February 1996 to August 1997. Readings were made early in the morning (around 08:00 local time) or after the storm. Personal observation in the plots showed that the upper canopy dried out within 6 h after rainfall, if this event occurred during the daytime or part of it. Therefore, the criteria to separate the single events was that only those events where no rainfall occurred in the previous 6 h were considered. In those cases where two or more events occurred during the night time, since evaporation during the night is expected to be low, they were considered as a single event. Most analysed single events in this study are events, preceded by a dry period of more than 10 h. Evaporation (E), which is taken here as the total C. Tobón Marin et al. / Journal of Hydrology 237 (2000) 40–57 amount of water intercepted and evaporated from the forest canopy, is calculated on event basis from the difference between gross rainfall (Pg) and net rainfall (throughfall Th plus stemflow Sf) of the single rainfall events. Cumulative evaporation during the rainfall events (Ew) is calculated from the difference between gross rainfall and net precipitation plus the water stored in the forest canopy after rainfall ceased (S), deduced from the estimated evaporation of single events minus the storage capacity (C) Ew Pg ⫺ Th ⫹ Sf ⫹ S 1 To avoid negative values in Eq. 1, although they may reflect those events where the forest canopy does not reach saturation, the value for S was inferred from the single rainfall events where E was larger than C. Statistical analyses were made for the entire collected data. However, for the assessment of the effect of rainfall sizes on net rainfall and resulting evaporation, measurements of single rainfall events are required. Therefore, throughfall percentages relative to gross rainfall and evaporation were calculated from these single events. Moreover, to determine the effect of forest structure on throughfall percentage and evaporation, we used only data from single rainfall events collected during the period of weekly measurements when collectors were not relocated. 4. Results 4.1. Forest structure Average values of measured variables of forest structure in each forest ecosystem studied are presented in Table 1. The largest mapped crown 2 45 2 area was 62.2 m and the smallest 2.4 m , both in the high-terrace plot. For the trunk surface area, the correction factor of 0.5 was applied to trees with diameter larger than 0.1 m. The apparent inconsistency in the flood plain data, of a large tree crown area in combination with the lowest tree trunk surface area, is mainly due to the abundance of small trees with a diameter of about 0.1 m. The data on the estimated gap fraction of the canopy and the free throughfall coefficient show that considerable differences exist between forests, the largest values being observed in the sedimentary plain. Moreover, the leaf area index appeared to decrease from the sedimentary plain to the flood plain of the River Caquetá. The estimated LAI values are within the range of values found for similar forests in Brazilian Amazonia (Roberts et al., 1996; Klinge et al., 1975). The decrease in gap fraction and increase in LAI towards the flood plain is in line with the results from Duivenvoorden and Lips (1995), who observed that litter fall in the flood plain of the River Caquetá is highest when compared to the other ecosystems in the area. 4.2. Rainfall characteristics Gross rainfall above the forest canopy did not vary considerably within plots. On average, the differences in the amounts of gross rainfall between subplots in the SP were 5.5% (^5.6) with n 3; whereas in the HT these were 3.7% (^3.5), in the LT gauges 3.7% (^3.0) and in the FP 3.1% (^2.7), all with n 2: Rainfall distribution differs between plots when examining separate storms, although annual totals are rather similar. During the measurement period (1992–1997), the mean annual rainfall at the AWS was about 3400 mm y ⫺1 and the average period with rainfall was 616 h y ⫺1. In total, 1584 rainfall events Table 1 Forest structure characteristics of four forest ecosystems in the Middle Caquetá, Colombian Amazonia Parameter Sedimentary plain High terrace Low terrace Flood plain Crown area (m 2) Stem trunk area (m 2) Gap fraction (%) Free throughfall coefficient, pt LAI Forest storage capacity (mm) 9.5 (^7.3) 7.2 (^4.6) 16.8 (^2.4) 0.59 4.4 (^0.7) 1.16 17.8 (^14.0) 8.2 (^5.2) 15.4 (^3.4) 0.52 4.9 (^0.8) 1.28 9.8 (^7.2) 8.4 (^5.4) 11.7 (^1.5) 0.49 5.6 (^0.6) 1.32 12.1 (^8.3) 5.4 (^4.6) 8.2 (^1.5) 0.27 6.6 (^0.4) 1.55 46 C. Tobón Marin et al. / Journal of Hydrology 237 (2000) 40–57 Fig. 2. Rainfall characteristics at the research site in the Middle Caquetá, Colombian Amazonia.(August 1992 to August 1997) C. Tobón Marin et al. / Journal of Hydrology 237 (2000) 40–57 Fig. 2. (continued) 47 48 C. Tobón Marin et al. / Journal of Hydrology 237 (2000) 40–57 Table 2 Throughfall percentages of daily gross rainfall for 5 storm classes, in four forests in the Middle Caquetá, Colombian Amazonia. (SP) sedimentary plain, (HT) high terrace, (LT) low terrace, (FP) flood plain, (std) standard deviation of the means and (n) number of events Rainfall ranges Throughfall % SP HT LT FP (mm) % std n % std n % std n % std n ⬍5 5–20 20–40 40–80 ⬎ 80 Total 58.7 81.4 88.9 90.6 92.8 87.2 11.4 6.3 2.8 2.1 – 2.4 41 78 39 19 1 178 56.2 80.5 87.9 90.0 92.2 86.7 12.1 5.6 2.6 1.9 0.8 2.4 34 68 36 19 3 160 52.3 79.8 87.7 88.8 92.4 85.8 9.6 6.8 3.4 2.5 1.0 1.2 32 71 41 17 2 163 47.4 74.5 83.0 84.6 88.5 81.9 13.2 6.9 2.5 3.1 1.2 1.0 27 57 32 19 5 140 were recorded at the AWS in Peña Roja, with storms ranging from 0.2 to 161.6 mm and lasting between 20 min and 13 h. During the total period, 37% of the incident rain fell in single showers of less than 2 mm and 92.3% of these showers contributed with less than 30 mm. Rainfall intensity, calculated for the total period with some rainfall, averaged 5.46 mm h ⫺1 with a maximum of 78.16 mm h ⫺1 (Fig. 2a). Most showers (63%) fell during the afternoon and at night (Fig. 2b) and 56% of these storms fell in less than 1 h (Fig. 2c). Monthly rainfall distribution during the fiveyear period shows that there was a slightly drier period from December to February (Fig. 2d). Comparing our data on five years rainfall with data from earlier years in the Middle Caquetá (Duivenvoorden and Lips, 1995), rainfall characteristics appear to be similar to the long-term average. 4.3. Throughfall The variability of throughfall within a subplot was large, with the smallest variation in the FP forest, although differences in average values between subplots were small. The average coefficient of variation (CV) of individual gauges in each plot was 0.285 (^0.10) in the SP, 0.306 (^0.07) in the HT, 0.279 (^0.09) in the LT and 0.225 (^0.08) in the FP forest. The CV of the mean throughfall in each subplot was 0.062 (^0.058) in the SP, 0.043 (^0.05) in the HT, 0.046 (^0.04) in the LT and 0.047 (^0.04) in the FP forest. As a general trend, for small rainfall events the value of the standard deviation (std) of throughfall (expressed as a percen- tage of mean throughfall) over gross rainfall varied more than for major events. Furthermore, for some individual throughfall gauges values exceeded gross rainfall (e.g. 29% of the individual gauge values in the SP were larger than gross rainfall, whereas in the HT this was 30%, in the LT 27% and in the FP forest 21%), but the average (of 60 and 40 gauges) was always lower than gross rainfall. Throughfall was calculated as a percentage of gross rainfall for five different rainfall sizes and from the totals of the measured daily gross rainfall and throughfall during the study. Throughfall ranged from zero, with events below 2 mm, to 95% in storms larger than 100 mm, but mean throughfall varied from 50 to 93% depending on gross rainfall amounts and the type of forest (Table 2). The calculated value of total throughfall relative to total gross rainfall ranged from 82 to 87% in the four forests. Although empirical regression equations provide only site-specific information, they may indicate a trend especially if explained variance is high. Therefore, Table 3 Regression parameters of throughfall versus gross rainfall in four different forest ecosystems in Colombian Amazonia. (se) standard error of regression coefficient (Note: The equation for linear form is T a ⫹ bPg ; where T is throughfall amount and Pg is gross rainfall (mm)) Landscape unit a b se R2 n Sedimentary plain High terrace Low terrace Flood plain ⫺1.02 ⫺1.02 ⫺1.07 ⫺1.48 0.926 0.918 0.906 0.887 0.003 0.003 0.004 0.003 0.99 0.99 0.99 0.99 102 97 97 84 C. Tobón Marin et al. / Journal of Hydrology 237 (2000) 40–57 49 Fig. 3. Average throughfall and its standard deviation (std) against gross rainfall in a forest ecosystem (Sedimentary plain) in the Middle Caquetá, Colombian Amazonia. regressions of throughfall versus gross rainfall were computed from single storms for each forest (Table 3). Average throughfall per plot was highly correlated with gross rainfall in all forests (Fig. 3). ANOVA analysis showed that the ratio of mean throughfall over gross rainfall in the FP forest was significantly different from the other forests (at 95% level). 4.4. Stemflow Large differences were observed in the amount of stemflow of individual trees and among subplots. In general, however, the contribution of stemflow to net rainfall was very low. The average CV in each plot was 0.295 (^0.12) in the SP, 0.207 (^0.12) in the HT, 0.323 (^0.20) in the LT and 0.303 (0.26) in the FP Fig. 4. Average stemflow and its standard deviation (std) against gross rainfall in a forest ecosystem (Sedimentary plain) in the Middle Caquetá, Colombian Amazonia. 50 C. Tobón Marin et al. / Journal of Hydrology 237 (2000) 40–57 Table 4 Summary statistics for regressions of daily stemflow against gross rainfall, in four forest ecosystems in the Colombian Amazonia (Note: The power form is Ps c Pdg where c and d are the regression coefficients for stemflow (mm)) Landscape unit Regression coefficient c d se R2 n Sedimentary plain High terrace Low terrace Flood plain 0.0015 0.0020 0.0029 0.0031 0.049 0.038 0.035 0.050 0.92 0.94 0.95 0.91 86 92 87 73 1.53 1.467 1.423 1.325 forest. The percentage of stemflow in all plots varied from 0.2 to 3.2% of gross rainfall. The total average percentage of stemflow relative to gross rainfall was 0.85% (^0.46) in the SP, 0.94 (^0.51) in the HT, 1.45 (^0.88) in the LT and 1.12 (^0.56) in the FP forest. Differences are mainly due to the higher contribution of tree palms to the total stemflow per plot. For palms, high-capacity collectors (more than 35 l) were required to measure the incoming water. In subplots with abundant palms, these palms produced about 43% of total stemflow. Upon rainfall, in all forests stemflow increased very gradually until a threshold of about 25 mm gross rainfall is reached (Fig. 4). However, values tend to scatter with increasing rainfall. The relationship between measured stemflow and gross rainfall could be described with a power function (Table 4). Some rainfall events smaller than 3 mm did not produce stemflow in most plots, which explains the lower number of events (n) reported for stemflow regressions. For rainfall events with an intensity ⬎5 mm h ⫺1, stemflow showed no clear relationship with tree trunk area or bark texture R2 0:3: Nevertheless, there seems to be an inverse relationship between crown area and the amount of collected stemflow for each tree. We also observed that lower parts of tree trunks with fibrous bark texture were slowly wetted during long storm events, which points to high water storage. 4.5. Evaporation Evaporation of intercepted water by the forest Fig. 5. Evaporation against gross rainfall in the sedimentary plain forest ecosystem, Middle Caquetá, Colombian Amazonia. C. Tobón Marin et al. / Journal of Hydrology 237 (2000) 40–57 51 Table 5 Statistics of evaporation from wet forest canopy in single rainfall events in four forest ecosystems in the Middle Caquetá, Colombian Amazonia. Equations are of the form Ew e ⫹ ft ; (e in mm and f in mm h ⫺1) (Note: e and f are the regression coefficients for the linear function of evaporation loss during rainfall events) Forest SP HT LT FP Total rainfall (mm) 3273.8 3293.0 3158.4 3120.9 Total throughfall (mm) 2853.7 2854.8 2711.7 2555.0 Total stemf. (mm) 32.4 36.2 38.6 30.5 time (h) 557.0 464.2 487.8 472.3 Evap. during rainfall (Ew) mm 190.1 207.4 201.1 320.0 canopy is calculated by subtracting the measured daily throughfall and stemflow from gross rainfall. Furthermore, it is related to gross rainfall characteristics and forest system parameters. Following net throughfall trends, the percentage of evaporation relative to gross rainfall varied from 6 to 100% in all forests, depending on rainfall size. Mean evaporation, expressed as percentage of total gross rainfall, also differed between the forests: 11.84 (^2.4) in the SP, 12.24 (^1.2) in the HT, 12.92 (^1.1) in the LT and 17.15 (^0.96) in the FP forest. For small storms (less than 2 mm), evaporation values were very close to those of gross rainfall. For heavy showers, however, the relative value of evaporation became smaller (Fig. 5). An assessment of the cumulative evaporation during rainfall (Ew) was calculated with Eq. 1 for the single rainfall events in each forest. Additionally, Ew was related to rainfall duration. The average evaporation rate during the rainfall varied from 0.34 to 0.68 mm h ⫺1 among the forests (Table 5) and Ew exhibited a linear relation with rainfall duration for all forests (Fig. 6). Negative values correspond to rainfall events of short duration and low intensity, indicating that the forest canopy did not reach saturated conditions during such events. Though climatic conditions are similar in the forests studied, there is a clear difference in amounts of evaporation when comparing similar rainfall events. This implies that amounts of evaporation from the wet forest canopy did not only depend upon gross rainfall and climate conditions. Differences in evaporation between these close-by forests may be related to differences in their structure. For Evap. rate from wet canopy mm h ⫺1 0.342 0.447 0.412 0.677 Ew versus rainfall duration. Regression coeffficient e f ⫺0.424 ⫺0.263 ⫺0.351 ⫺0.366 0.46 0.52 0.52 0.78 R2 n 178 0.86 0.75 0.82 0.88 178 160 163 140 that reason, the ratio of evaporation over gross rainfall was plotted against the mean forest cover (1-gap fraction) established for each forest. Fig. 7 indicates that there is an increase of evaporation from the wet forest canopy with increasing canopy cover. This figure also shows an inflexion in the curve, i.e. a steeping from the low terrace to the flood plain, which indicates that also other forest structural parameters (e.g. leaf surface characteristics) affected canopy interception. 5. Discussion Storage capacity values of the forests studied on the whole resemble the values found by Ubarana (1996) in the reserves Vale do Rio Doce and Jaru Duke in Brazil, which were based on linear regressions of throughfall against gross rainfall. However, Ubarana concluded that this method results in an overestimation of evaporative losses. Since our estimates of the canopy storage capacity were based on specific events for which it was assumed that evaporation was negligible, it might be that we somewhat overestimated the storage capacity by neglecting evaporation during the selected events. The storage capacity of the flood plain forest is higher than the values commonly reported in Amazonian rain forest studies. This can be explained by the fact that most of the latter studies were executed in so called “terra firme” forests (non-flooded ecosystem). As we studied a broader range of ecosystems, differences between parameters values reported here and those reported in other studies therefore should be interpreted in terms of 52 C. Tobón Marin et al. / Journal of Hydrology 237 (2000) 40–57 differences in forest structure between the ecosystems studied. The free throughfall coefficients of our forests, derived from the data on selected small storms (pt), conform to the values found by Jetten (1996). However, these values which range from 0.27 to 0.59 clearly differ from those derived from the scanned photographs (ps) and their application resulted in an overestimation of net rainfall rates in all forests. According to Ubarana (1996) this may be explained by the waxy nature of the tree leaf surfaces causing the drops to splash off, which thus contribute to throughfall before the canopy is saturated. Additionally, our values were based on the results for small showers producing throughfall and this may have influenced the estimation of the free throughfall because of the low frequency of such events (for the research sites only 7–10 events were registered during the total period). Photographs taken from the forest canopy under non-direct sunlight and with covered sky can easily be analysed, the white pixels reflecting the non-covered part. This estimate may provide better results, especially if large numbers of photographs have been taken and analysed. The range in our values for throughfall and stemflow, expressed as percentage of gross rainfall, is similar to the range in values reported in earlier studies on rainfall partitioning in Amazonian rain forests (Table 6). This most probably also explains why such variability exist in the latter values, i.e. it is probably largely due to differences in rainfall characteristics and forest structure between the forests studied. That coefficients of determination for our regressions are significantly higher than most values presented in the literature can be explained by the size of our data set, which is much larger than in earlier studies. Our results also indicate that the partitioning of rainfall depends, among others, on the size of the rainfall event. Moreover, it is clear from the relation between throughfall and storm size (Table 2) that the high CV of throughfall is the result of the large variability in rainfall classes. To define the total error (t.e.) of the mean throughfall as a percentage of gross rainfall, we applied the proposed formula for random relocation of n gauges Fig. 6. Evaporation from the wet forest canopy (Ew) in relation to rainfall duration, in a forest ecosystem (sedimentary plain) in the Middle Caquetá, Colombian Amazonia. C. Tobón Marin et al. / Journal of Hydrology 237 (2000) 40–57 53 Fig. 7. Fraction of evaporation from gross rainfall as a function of forest cover fraction (FC 1-gap fraction) in the four forest ecosystems. Middle Caquetá, Colombian Amazonia. by Lloyd and Marques (1988), although larger diameter funnels were used in the current study p t:e: s:e: 1 ⫹ N=nm 2 where s.e. is the standard error resulting from the random relocations of collectors, expressed as the best estimate of the standard error of mean throughfall in each collector, under the assumption that the specific canopy structure is properly described by N (number of grid points) and m (the relocation of collectors). Based on the formula of Lloyd and Marques (1988), the arrangement of 60 funnels with 23 relocations in the SP forest results in a total error in measured throughfall of 3.5% of gross rainfall, whereas in the other ecosystems with 40 gauges and 23 relocations the error is 3.8% due to variation in canopy structure. These figures are lower than those found by other authors, which can be explained by the continuous relocation of our collectors. Nevertheless, our values for t.e. are larger than those found by Lloyd and Marques (1988). Whether throughfall percentage depends on storm size remains to be established, as clearly stated by Lloyd and Marques (1988). Accordingly, we investigated the effect of storm size on the variability of throughfall percentage by using only those single events that were measured during the 20-months period in which weekly data of throughfall and gross rainfall were collected. During that period, the same methodologies were used with the exception that gauges were not relocated. Fig. 8, given as an example, shows that the variation in the ratio of throughfall from a single funnel over the average of 20 funnels tends to decrease as storm size increases. This is a trend observed for most non-moving collectors but also for the relocated collectors, as stated earlier in this paper, which suggests that storm size also affect throughfall variability in our ecosystems. We did not fully investigate the relation between storm size and throughfall (expressed as percentage of gross rainfall). Nevertheless, we conclude from our results that when the method of relocation of collectors is used to estimate this throughfall, it is essential 54 C. Tobón Marin et al. / Journal of Hydrology 237 (2000) 40–57 Table 6 Partitioning of gross rainfall (percentages) in Amazonian rain forests Location Forest type Throughfall % Stemflow % Evaporative loss % Reference Venezuela Brazil Brazil Brazil Colombia Catinga Rain forest Rain forest Rain forest Rain forest 91 80.2 87–91 86–87 82–87 0.8–14 – 1.8 0.8–1.4 0.9–1.5 – 19.8 8.9(^3.6) 11.6–12.9(^5.9) 12–17 Herrera (1979) Franken et al. (1992) Lloyd and Marques (1988) Ubarana (1996) This study that this relocation is preceded by sampling of a wide range of storms sizes with a fixed set of collectors, in order to assess the combined effect of site forest structure and rainfall characteristics on throughfall. Lastly, although litter fall in the ecosystems studied exhibits some temporal dynamics (Duivenvoorden and Lips, 1995), no relation was observed between throughfall percentage and litter fall. Such relation was reported for a Bornean rain forest by Burghouts et al. (1988). The variability of stemflow in mature tropical rain forest has been attributed to the high species diversity (Hutjes et al., 1990; Hertwitz, 1985) and this variability certainly is larger in tropical forests than in temperate forest plantations (Lloyd and Marques, 1988). In the present study, this parameter was estimated for different tree species with different diameter. Stemflow values from this study ranged from 0.9 to 1.5% of gross rainfall, which is within the range of values presented in other studies on similar forest types (Table 6). Although the contribution of stemflow to net rainfall was very low, it probably causes an important input of solutes to the forest floor, concentrated around the base of trees. Results suggest that little water was stored in excess of the storage capacity of the stem elements, as indicated by the very small stemflow quantities collected once rainfall has ceased or during small storms. This can be explained by the presence of some tree trunks with hydrophobic bark (personal observations 1992–1997) and of bark with fibrous texture. Upon rainfall, tree species with these characteristics exhibited significant stemflow, even without being completely wet. However, once rainfall stopped, there was a sharp decline in stemflow. We found static models to be capable of describing rainfall partitioning for the forests studied. The applicability of these models is most probably restricted to the area and conditions during the period of research. Though the observed relationships may contribute little to the explanation of the hydrological processes at canopy level, the models nevertheless provide clear indications for the extent to which this partitioning is controlled by the parameters used. While linear functions produce better fits for correlation between throughfall and gross rainfall, power functions produce better fits for such correlation with stemflow, in terms of the significance levels and standard deviation of residuals. The linear regression equations of throughfall versus gross rainfall fit most points and have a high coefficient of determination in all ecosystems. Nevertheless, their application to very small storms (lower than 2 mm) results in negative values for throughfall and they slightly underestimate throughfall for very high-rainfall events, which illustrates the limitation of regressions, which fit a curve to a set of data. Throughfall and gross rainfall were highly correlated in all forests, which is probably due to the similarity of our forests with regard to relevant system parameters. However, the correlation between interception values and gross rainfall is less prominent, the coefficient of determination of the regression being distinctly lower (R 2 between 0.66 and 0.83). In other words, throughfall percentages can be predicted with a high accuracy based on data on rainfall amounts and characteristics, whereas for the prediction of interception other parameters, such as forest structure, must be included. Although our values are within the range of interception values reported in other studies from the Amazon basin (Table 6), the values found for the FP and LT forests are rather high compared to those reported in earlier Amazonian studies (Lloyd and Marques, 1988). This is in line with the higher canopy storage capacity of our forests as compared to those described in these other studies (e.g. Lloyd and C. Tobón Marin et al. / Journal of Hydrology 237 (2000) 40–57 55 Fig. 8. The ratio of single gauge throughfall over average throughfall from 20 gauges, as a function of gross rainfall size. Marques, 1988). Thus, even though climatic conditions for our forests were similar, forest interception varied, due to differences in amounts and characteristics of the rainfall and in forest structure. The relative proportion of evaporation from our forests was also higher than the values reported in these earlier studies. This can be attributed to the higher gross rainfall in our forests, at least when compared with the mean annual value of 2500 mm reported from central Amazonia (Leopoldo et al., 1987). In many studies on rainfall interception, it is concluded that leaf surfaces determine the interception storage capacity of woody plants (Hertwitz, 1985; Gash, 1979; Singh, 1977; Rutter et al., 1975). Studies in the research plots, using destructive methods and derived regression equations for leaf biomass estimation (Overman et al., 1990; Alvarez, 1993), showed that leaf biomass is higher in the flood plain forest (9.5 tonnes/ha) than in the other forests from which a higher leaf surface area can be inferred. Therefore, the relatively high interception by the flood plain forest may be explained by its higher leaf biomass. Cumulative evaporation during rainfall events (Ew) from our forests strongly depended on rainfall dura- tion. Although not evaluated in this study, it might also depend on specific climatic conditions (e.g. wind speed). Additionally, a distinct relationship seemed to exist with the forest cover fraction (Fig. 7). Although it should be realised that the number of forests studied is small and the relationship is rather uncertain, it may serve for the estimation of evaporation by a forest for which measurements are not available. Provided that climatic conditions are similar, such estimations mainly rely upon an adequate estimation of the gap fraction or LAI. 6. Conclusions Of the gross rainfall of about 3400 mm y ⫺1, most fell in small showers during the afternoon and at night. The overall average rainfall intensity was about 5 mm h ⫺1. These rainfall characteristics largely explain the partitioning of rainfall into throughfall, stemflow and ensuing evaporation in the forests studied. Water fluxes in the forest canopy of four forest ecosystems in western Amazonia have been quantified 56 C. Tobón Marin et al. / Journal of Hydrology 237 (2000) 40–57 as a percentage of gross rainfall. Amounts of net precipitation reaching the forest floor and evaporation from the wet forest canopy varied for the forests studied: the SP forest had the highest percentage of throughfall relative to gross rainfall and the FP forest the lowest. The observed differences in throughfall, stemflow and evaporation can partly be attributed to differences in forest structure (gap fraction). Their range is similar to the overall range in these parameters as published in earlier studies from the Amazon basin, implying that the latter variability may very well be connected with differences in forest structure. Results from the forests studied provide some insight into the rates of evaporation from a wet forest canopy and strengthen the understanding of the contribution of forests to atmospheric moisture. The mean evaporation rate from a wet forest canopy during rainfall events in the Middle Caquetá, (Colombian Amazonia) was estimated at 0.47 mm h ⫺1 and it increased with increasing forest cover. Moreover, this study of throughfall, stemflow and evaporation in a range of forests demonstrates the relevance of forest structure for the evaporation of rainfall intercepted by the forest canopy and for the net precipitation reaching the forest floor. The results show that within the scope of this research forest structure can be adequately characterised by the gap fraction and LAI. These structural characteristics together with the rainfall amount and rainfall duration are the main parameters determining rainfall partitioning in the western Amazonian rain forests. Acknowledgements We are grateful to Dr John Gash from the UK Institute of Hydrology and to Dr Sampurno Bruijnzeel from the Free University, Amsterdam for their suggestions and corrections of earlier drafts of this paper. 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