Tree Physiology 24, 1035–1044 © 2004 Heron Publishing—Victoria, Canada Water use by fast-growing Eucalyptus urophylla plantations in southern China JIM MORRIS,1,2 ZHANG NINGNAN,3 YANG ZENGJIANG,3 JOHN COLLOPY1 and XU DAPING 3 1 Forest Science Centre, Heidelberg, Victoria 3084, Australia 2 Corresponding author ([email protected]) 3 Research Institute for Tropical Forestry, Chinese Academy of Forestry, Guangzhou 510520, China Received October 6, 2003; accepted January 24, 2004; published July 1, 2004 Keywords: canopy conductance, Guangdong, heat pulse, sap flow, soil moisture, transpiration, vapor pressure deficit. Introduction Eucalyptus plantations in several countries have been the subject of criticism because of their high water use and other negative environmental impacts. Examination of the evidence for these claims has usually concluded that well-managed plantations are beneficial rather than detrimental to the environment (Poore and Fries 1985, White et al. 1995, Casson 1997). How- ever, studies in India (Calder et al. 1992) and in South Africa (Dye 1996) indicate that, when water resources are limited, the area, location and management of plantations must be carefully considered to avoid conflict with other water users. Detailed data on the water use and water balance of plantations are essential both to evaluate their environmental impacts and to design optimal land use strategies in catchments where wood production is an important economic component. Recently, research in several countries, including Australia (Honeysett et al. 1996, Myers et al. 1996, Morris and Collopy 1999), Brazil (Soares and Almeida 2001), Portugal (Osorio et al. 1998), South Africa (Scott and Smith 1997, LeMaitre et al. 2002), India (Kallarackal and Somen 1997) and Pakistan (Mahmood et al. 2001), has increased our understanding of the hydrology of eucalypt and other exotic tree plantations. Plantations of eucalypt and other exotic tree species have been extensively established in southern China during the past 50 years (Qi 2002). The total area of eucalypt plantations in China was around 460,000 ha in 1993 (Bai and Gan 1996), with a more recent estimate at 1.3 million ha (FAO 2001), increasing by 100,000 ha or more per year in response to government afforestation programs and cooperative joint ventures between provincial forestry authorities and foreign investors. In addition to their key role in timber and fiber production, eucalypt plantations in China have an important ecological role in the revegetation of degraded lands (Corlett 1999, Zhou et al. 2002). The Leizhou Peninsula of western Guangdong province (Figure 1) is an example of a region where eucalypt plantations have become a significant component of the regional economy. The forest estate of the region includes more than 200,000 ha of eucalypt plantations, from which more than 60% of the wood production is used as wood chips for export and for the domestic paper industry. It has been suggested that high water use by eucalypt plantations may contribute to the depletion of groundwater resources on the peninsula. Mean annual rainfall in the area is over 1300 mm, but the topography of the region and the deep permeable soils limit opportunities for storage of surface water, so farmers must often extract groundwater during the dry season to establish or maintain Downloaded from http://treephys.oxfordjournals.org/ by guest on August 29, 2014 Summary Tree growth, water use, climate and soil water conditions were monitored over 12 months in two 3– 4-yearold Eucalyptus urophylla S.T. Blake plantations on the Leizhou Peninsula of southern China. The Hetou plantation was established on a sandy soil of sedimentary origin with low water storage capacity, and the Jijia plantation was established on a clay soil formed on basalt. Sapwood area was ~50% higher at Jijia than at Hetou because of differences in plant spacing (1994 versus 1356 stems ha –1). Annual water use, assessed by heat pulse measurements, was 542 mm at Hetou and 559 mm at Jijia, with mean sap flux densities of 2772 and 1839 l m –2 day –1, respectively. Limitations to water use, imposed by climatic and soil factors, were quantified by analysis of daily canopy conductance in relation to daytime vapor pressure deficit (VPD) and soil water content. Similar annual water use at the two sites was a result of higher VPD and soil water availability at Hetou compensating for the greater sapwood area at Jijia. Potential annual water use in the absence of soil water limitation was estimated at 916 mm at Jijia and 815 mm at Hetou. Higher water availability during the dry season and early wet season at Hetou than at Jijia was the result of deep root systems. The results imply that water use by plantations on soils with high water availability and in areas of high VPD may be reduced by establishment at wider spacing. The environmental cost of water use by plantations must be weighed against their economic and environmental values to determine an appropriate mix of forestry, agriculture and other land uses in regions where water resources are limited. 1036 MORRIS ET AL. Quaternary sediments. At both locations, the plantations are monocultures of E. urophylla planted in mid-1996. Tree spacing is 3 × 2.5 m at Hetou and 3.3 × 1.5 m at Jijia. An understory of native shrubs developed following the removal of all vegetation during planting preparations, but is patchy and relatively light at Hetou. Slopes at both sites are < 1%, near the top of a broad ridge defining the catchment boundary at Jijia (70 m above sea level) and on a low divide between streams in a broad valley at Hetou (25 m above sea level). Weather monitoring crops of sugar cane and rice. Water use by plantations has been shown to affect water table depth in other countries, but little research has been conducted on the hydrology of eucalypt plantations in China, and there have been no previous studies of plantation water balance on the Leizhou Peninsula. Zhou et al. (1995) estimated transpiration by a eucalypt plantation in western Guangdong at 614 mm year –1 based on an empirical function of rainfall and potential evaporation, and suggested that eucalypts are limited by water availability as a result of low water storage in the root zone. As part of an investigation of hydrological impacts of plantations on the Leizhou Peninsula, we studied the water use of plantations of Eucalyptus and other species at several locations. This paper reports daily and annual water use by stands of Eucalyptus urophylla S.T. Blake at two contrasting sites, and examines the influence of climate and soil water as controlling factors for plantation water use in the region. Methods Study areas Two sites were selected on forest farms near Hetou (21°05′ N, 109°54′ E) and Jijia (20°54′ N, 109°52′ E) in the Nandu River catchment of the Leizhou Peninsula (Figure 1). The topography of the peninsula is flat to undulating, with deeply weathered red soils derived from basaltic and sedimentary parent materials. The climate is tropical, with monthly mean temperatures of around 28 °C in July and 16 °C in January. Annual rainfall varies from 1300 mm in the south to 1800 mm in the north of the peninsula, with high year-to-year variability (standard deviation of 350 to 400 mm). Over 80% of the rain falls between April and September, up to half of this in typhoons that occur two to three times per year. The Jijia site is on a basalt-derived clay soil, whereas the Hetou site is about 40 km north on a sandy soil formed from λE = ε Rn + λρDq ga ε +1 + ( g a / g c ) (1) where ε is the dimensionless rate of change of saturated specific humidity with temperature (estimated as 2.2), Rn is net radiation, Dq is specific saturation deficit (calculated as 0.00622 × VPD), ga is aerodynamic conductance, gc is canopy conductance (assumed to be infinite for a wet canopy), λ is the latent heat of evaporation of water and ρ is air density. Daily net radiation was estimated from solar radiation as: Rn = 0.8R s – 90 W m –2 and aerodynamic conductance as 0.1 times wind speed in m s –1 (Landsberg 1986). Although both of these estimates are approximations of the true values of Rn and ga, neither variable is of critical importance to the calculations of this study and so the approximations were considered acceptable. Mean daytime values of VPD and potential E were calculated by averaging half-hourly data for each day when solar radiation was greater than zero. Soil conditions To monitor soil water, a set of four standing wave dielectric soil moisture probes with a waveguide length of 60 mm (MP406, Agri-Tech Instruments, Beijing, China) were buried at each site, at depths of 50, 150, 250 and 350 cm below the soil surface. Output from the probes was recorded at half-hourly intervals by the weather station data loggers. Calibration to volumetric water content (θ) was based on soil samples col- TREE PHYSIOLOGY VOLUME 24, 2004 Downloaded from http://treephys.oxfordjournals.org/ by guest on August 29, 2014 Figure 1. Leizhou Peninsula, western Guangdong province, showing the Nandu River system and location of the monitored plantations near Jijia and Hetou. A 0.1-mm tipping-bucket rain gauge and meteorological sensors (SC1 pyranometer, PTAT temperature sensor, Vaisala humidity sensor and cup anemometer from Tain Instruments, Box Hill, Australia) were mounted on a mast 7.5 m above ground level in a cleared area close to the plots monitored for water use within each of the plantations in September 1999. Half-hourly observations of solar radiation, rainfall, air temperature, relative humidity and wind speed were recorded by a Tain Micropower data logger. Data were downloaded at approximately monthly intervals. Daily values of solar radiation (Rs ), rainfall, maximum and minimum temperature and mean wind speed were calculated, and daily mean vapor pressure deficit (VPD; kPa) was derived from the half-hourly humidity data, with saturation vapor pressure estimated as an exponential function of temperature (Dilley 1968). Potential (wet canopy) daily evaporation (E; mm day –1) was calculated based on the Penman-Monteith combination equation, expressed by Landsberg and Gower (1997) as: EUCALYPTUS PLANTATION WATER USE IN CHINA Tree growth and sapwood area A 40 × 40 m monitoring plot was selected in each plantation in September 1999. Within each monitoring plot, diameters at breast height (DBH; cm, 1.3 m above ground level) were measured on all trees at intervals of about 6 months. For a small number of multi-stemmed trees, an equivalent diameter was calculated as the DBH of a single stem with the same basal area as the combined stems. A regression with diameter (H = 1.5584DBH + 0.0257DBH 2, R 2 = 0.88) was derived with heights (H; m) of a subset of 34 trees at each plantation, measured with a Vertex hypsometer (Forestor Instruments AB, Sweden). This estimated the heights of all trees and inferred mean dominant height (mean of the largest 200 trees ha –1), total volume (from single-tree volume approximated as DBH 2 × H/30,000 m3 ) and mean annual volume increment over the life of the stand. Comparison with the results of growth studies reported by Baker et al. (2003) suggests that the simple approximation adopted for single-tree volume yields estimates within 5% of actual stem volume. Sapwood areas of 40 trees including those selected for water use measurements at each monitoring plot were calculated from measurements of DBH, bark thickness and sapwood width from increment cores taken at 2– 4 points around the stem. The sapwood–heartwood boundary was identified by color difference, light transmission or with the assistance of 0.1% aqueous methyl orange stain. Second-degree polynomial regressions of sapwood area against DBH were derived to estimate the sapwood area of all trees in each plot at the times of diameter measurement, and hence the total plot sapwood area. Leaf area Canopy leaf area index (LAI) was assessed at both sites in April and August 2000, with an Accupar ceptometer (Decagon Devices, Pullman, WA) held ~1 m above ground. The mean below-canopy photosynthetic photon flux (PPF) was calculated over 10 s at each point in a 5 × 5 m grid within each plot, and expressed as a fraction of PPF recorded simultaneously by a PR1 PPF sensor (Monitor Sensors, Caboolture, Australia) located in an open area outside the plantation. We calculated LAI from functions provided with the ceptometer, assuming a leaf distribution parameter of 1 (approximately spherical canopy elements) and calculating zenith angle from latitude, longitude and time. The LAI values at all grid points were averaged to derive a single plot LAI for each site. Measurements at Jijia, but not at Hetou, unavoidably included a small but unknown contribution from understory shrubs and small trees up to 5 m in height. Tree water use Water use by E. urophylla plantations at Hetou and Jijia was monitored from September 1999 to September 2000, for a total of 583 and 544 tree-days, respectively, by the heat pulse method described by Edwards and Warwick (1984) and modified by Olbrich (1991). A set of 18 trees were selected for water use measurement at each site, matching the diameter distributions of the monitoring plots. From these 18 trees, two were randomly selected per site at intervals of 3 to 4 weeks for implantation with teflon thermistor probes and stainless steel heaters for heat pulse monitoring with HeatPulser instruments (Edwards Industries, Otaki, New Zealand) and CR10X data loggers (Campbell Scientific, Logan, UT). Measurements of apparent heat pulse velocity were recorded at 30-min intervals with four pairs of thermistor probes per tree and a 1-s heat pulse duration. The data were corrected for the effects of wound diameter, probe separation and volume fractions of water and woody matrix in the sapwood to derive estimates of sap flux density (SFD; cm 3, cm –2 h –1). A whole-tree SFD estimate was derived from each set of four point estimates by calculating a second-degree polynomial regression against implantation depth below the cambium, then integrating this function around the stem. In some trees where probe failures reduced the number of point measurements to less than four, a weighted mean calculation similar to that of Hatton (1990) was adopted. Instantaneous tree water use was derived by multiplying SFD by sapwood area, estimated from under bark diameter and sapwood widths measured at two to four points around the stem. Total daily water use was obtained by integration of the half-hourly values over each day, and dividing by the sapwood area to give a mean daily SFD in units of dm 3 m –2 day –1. Scaling of individual tree observations to stand water use was based on sapwood area, as described by Khanzada et al. (1998). The daily mean SFD values recorded for individual trees were assumed to be unbiased estimates of the mean SFD of the whole stand. This assumption is supported by the observation that differences in SFD between trees measured simultaneously or sequentially were usually small compared with day-to-day SFD variation within a tree. Daily plot water use was therefore estimated as the product of plot sapwood area (interpolated for each day from the six monthly measurements, assuming continuous year-round growth) and mean daily SFD of the monitored trees, and annual water use was calculated as 365 times the mean of the daily values. Canopy conductance For evaporation from rough forest canopies, the radiation term of Equation 1 is often small compared with the vapor deficit TREE PHYSIOLOGY ONLINE at http://heronpublishing.com Downloaded from http://treephys.oxfordjournals.org/ by guest on August 29, 2014 lected at the time of installation and 6 months later from auger holes within 1 m of the probes. The volume of stored soil water to 4 m depth was calculated by taking the water content at each probe as an estimate of mean water content in a 1 m stratum. Water release characteristics and particle size distribution of the soils were assessed from soil samples collected at both sites in September 1999 (end of the wet season) and April 2000 (end of the dry season). Matric potential (Ψm ) of samples from 30-cm depth intervals to 390 cm was determined by the filter paper method (Greacen et al. 1989), and gravimetric water content of the same samples was measured. Bulk density was determined at 20-cm intervals to 80 cm as the mean of three samples collected with 200-cm3 fixed-volume sampling cylinders. Plant available water was determined as the difference between field water content and water content at wilting point (estimated as –1500 kPa), based on a power curve fitted to the water release data for each site (Ψm = –0.05θ –10.95 at Jijia, Ψm = – 0.05θ –5.63 at Hetou). 1037 1038 MORRIS ET AL. term, and aerodynamic conductance is large compared with canopy conductance (Jarvis 1981). Under these conditions, canopy conductance (gc; m s –1) may be estimated from observations of daily water use and daytime VPD as: gc = γ λT ρ C pV (2) Figure 2. Monthly rainfall recorded at Jijia and Hetou between October 1, 1999 and September 30, 2000. study sites was unaffected by this brief chilling period, although frost-sensitive trees were killed in plantations 100 km north. Measurements of wind speed were affected by shelter from the surrounding plantations as a result of the sub-canopy height position of the anemometers, but maximum half-hour means reached 2.5 m s –1 at Jijia and 1.8 m s –1 at Hetou. The dry season was windier than the wet season at both sites. Daily Results Weather conditions Rainfall in the 12 months from October 1, 1999 to September 30, 2000 was 1539 mm at Jijia and 1531 mm at Hetou. Long-term meteorological records for the Leizhou Peninsula indicate that both 1998 and 1999 were years of below average rainfall (73.5 and 80.8% of the 21-year mean at Zhanjiang), whereas in 2000, the rainfall returned to the 21-year mean. There was a clear differentiation in monthly rainfall distribution between the wet and dry seasons (Figure 2). The wettest days were June 26, 2000 at Hetou (163 mm) and June 27, 2000 at Jijia (128 mm), and the highest rainfall intensity recorded was 48.6 mm in 30 min. Daily total solar radiation at Jijia is shown in Figure 3A. Radiation at Hetou followed a similar pattern to that at Jijia, but was slightly less, averaging 94% of the Jijia daily values. Overcast skies are common at all times of the year in this region of the Leizhou Peninsula. Mean daytime VPD (Figure 3B) rarely exceeded 2.5 kPa, and was less than 1.5 kPa throughout the dry season from December to March. The VPD was 17% higher on average at Hetou than at Jijia, as a result of temperature and relative humidity differences. Throughout the year, monthly means of daily maximum temperature were 1 to 2 °C higher at Hetou than at Jijia, reaching a maximum of 36.8 °C at Hetou and 34.6 °C at Jijia in June. Daily minimum temperatures rarely fell below 7 °C, except during the week of December 21–28 when temperatures of zero or below were recorded at both sites. Vegetation at the Figure 3. Climate and soil conditions at Jijia and Hetou, October 1999–September 2000. (A) Daily solar radiation; (B) daily mean vapor pressure deficit during daylight hours; and (C) available soil water (0 to 4 m depth), estimated as the difference between observed field water content and water content at –1500 kPa. TREE PHYSIOLOGY VOLUME 24, 2004 Downloaded from http://treephys.oxfordjournals.org/ by guest on August 29, 2014 where T is mean rate (mm s –1) of plot water use for the day, and V (kPa) is mean daily daytime vapor pressure deficit. This formula was applied to estimate gc for the plantations at Hetou and Jijia on each day when the vapor deficit term in the numerator of Equation 1 exceeded the radiation term by a factor of five or more. The psychrometric constant (γ), latent heat of evaporation of water (λ), density of air (ρ) and specific heat of air at constant pressure (Cp ) were estimated as 66 Pa K –1, 2450 kJ kg –1, 1.2 kg m –3, and 1010 J kg –1 K –1, respectively. To calculate T, the estimated daily stand water use was divided by day length, calculated from latitude and date following Sellers (1965). To test the assumption that aerodynamic conductance exceeded canopy conductance sufficiently to make Equation 2 valid, the calculated daily values of gc were sorted into five classes of mean daily wind speed, and plotted against V. There was no evidence of a difference in the relationship of gc and V between days of high and low wind speed, confirming that aerodynamic conductance was high enough to permit use of Equation 2 on a daily time scale. EUCALYPTUS PLANTATION WATER USE IN CHINA 1039 potential evaporation calculated from the meteorological data was between 0–15 mm day –1 during the dry season and 5– 25 mm day –1 during the wet season, with high short-term variation. Daily potential evaporation was on average 16% higher at Hetou than at Jijia during both dry and wet seasons. Table 1. Soil properties at Hetou and Jijia. Bulk density was averaged over 20 – 80 cm depth; water content at –1500 kPa (WC) was estimated from a water release curve derived from matric potential and volumetric water content measurements on 104 soil samples from 0 – 3.9 m depth. Soil water content Location Bulk density Water content at saturation (%) Estimated WC at –1500 kPa (%) Hetou Jijia 1.64 1.01 38 62 16 38 Tree growth Tree density at Hetou was approximately 70% of that at Jijia. Plot measurements in September 1999, March–May 2000 and August 2000 indicated that both stands grew rapidly during the study, with an increase in basal area of 30% at Hetou and 19% at Jijia for the study year despite declining stand density as a result of mortality of suppressed trees (Table 2). Mean tree diameter and height were slightly greater at Hetou, but the higher stocking at Jijia led to greater sapwood area and volume growth. Leaf area index was relatively low at both sites, reflecting the open canopy structure; LAI increased during the wet season, although at Jijia the measured increase was partly a result of understory development during the period. Polynomial regressions derived to relate sapwood area (SA; cm2 ) to overbark diameter (DBH cm) of a sub-set of 40 trees ranging in DBH from 4 to 16 cm were: SA = 0.359DBH2 + Figure 4. Variation in soil particle size distribution with depth at Hetou and Jijia. 0.537DBH (R 2 = 0.96) at Hetou and SA = 0.331DBH2 + 1.298DBH (R 2 = 0.92) at Jijia. These site-specific relationships were used to calculate the stand sapwood areas in Table 2 and for scaling up single-tree water use measurements to a stand basis, although the regression lines did not differ significantly within the DBH ranges at Jijia and Hetou. Tree water use Single-tree water use averaged 13.9 dm 3 day –1 at Hetou with a maximum of 49 dm 3 day –1 compared with 8.6 dm 3 day –1 and a maximum of 27 dm 3 day –1 at Jijia. Two small suppressed trees at Hetou (DBH = 6.2 and 7.3 cm) were excluded from the calculation of stand water use because their sap fluxes were anomalously low compared with other trees measured during the same period, suggesting that their canopy exposure and access to soil water were atypical of the stand as a whole and that they are more appropriately considered as part of the understory. Trees of this size or smaller contributed about 10% of the sapwood area at both sites, but not all are necessarily suppressed. Daily sap flux density (SFD) and daily stand water use recorded at the two sites are shown in Figure 5. Estimates of annual stand water use based on the daily mean values at Hetou and Jijia (Table 3) were 542 and 559 mm, respectively, equivalent to 35.4 and 36.3% of rainfall for the period, respectively. Wet season (April–September) water use was 59.6% of the annual total at Hetou, but only 53.9% at Jijia, where water use was greater than at Hetou in the dry season (1.39 versus 1.25 mm day –1, averaged over 137 and 172 days, respectively) and less than at Hetou in the wet season (1.62 versus 1.85 mm day –1, averaged over 112 and 101 days, respectively). Water-use efficiency, defined as total stem volume growth per unit of water transpired, was 4.8 × 10 –3 ± 0.2 × 10 –3 m3 m –3 at Hetou and 4.2 × 10 –3 ± 0.2 × 10 –3 m3 m –3 at Jijia (P > 0.95). Despite lower planting density and hence lower stand volume at Hetou, volume growth at Hetou in 1999–2000 exceeded that of the higher density Jijia plantation (stand volume increased by 23.8 and 21.6 m3 ha –1, respectively, between September 1999 and August 2000), leading to a higher water-use efficiency at Hetou during the study period. Despite the similar annual and shorter-term water use estimates for Hetou and Jijia in Figure 5B, SFD differed between plantations, with an annual mean daily value about 51% higher at Hetou than at Jijia (Table 3). Sapwood area averaged 53% more at Jijia than at Hetou during the study year. Because water use is the product of SFD and sapwood area, the similarity of these ratios suggests that SFD varies between stands to TREE PHYSIOLOGY ONLINE at http://heronpublishing.com Downloaded from http://treephys.oxfordjournals.org/ by guest on August 29, 2014 Soil texture varied with depth at both sites (Figure 4). The coarser textured soil at Hetou had markedly higher bulk density and lower water-holding capacity than the clay-dominated soil at Jijia (Table 1). Half-hourly data from the buried soil moisture sensors showed extraction of soil water to at least 350 cm depth from September 1999 until the end of the dry season in early May 2000. Rain events during that period usually had no measurable effect on soil water content below 50 cm at Hetou or below 150 cm at Jijia. Replenishment of water at 350-cm depth occurred when about 200 mm of rain fell at both sites over four days in late June. Infiltration through the profile was notably more rapid at Hetou than at Jijia, consistent with the greater storage capacity of the clay soil at Jijia. Maximum available soil water in the upper 4 m of the profile was estimated to reach 450 mm at Jijia and 200 mm at Hetou, and was depleted at both sites to around 150 mm by the end of the dry season (Figure 3C). 1040 MORRIS ET AL. Table 2. Growth characteristics of 1600 m2 plots of 3–4-year-old E. urophylla plantations assessed at Hetou and Jijia in 1999–2000. Hetou Number of trees Stocking (stems ha –1) Mean diameter (cm) ± SE Basal area (m2 ha –1) Mean dominant height (m) ± SE (n = 32) Mean annual volume increment (m3 ha –1 year –1) Sapwood area (m2 ha –1) Leaf area index ± SE (n = 64) Jijia September 1999 March 2000 August 2000 September 1999 May 2000 August 2000 217 1356 8.5 ± 0.2 9.0 16.7 ± 0.2 213 1331 9.5 ± 0.3 11.0 18.0 ± 0.2 208 1300 9.9 ± 0.3 11.7 18.5 ± 0.2 319 1994 8.5 ± 0.2 12.9 16.3 ± 0.2 311 1944 9.3 ± 0.2 15.1 17.5 ± 0.2 303 1894 9.5 ± 0.2 15.3 17.8 ± 0.2 16.3 17.9 18.3 22.6 24.7 22.6 4.74 5.69 1.14 ± 0.03 6.03 1.53 ± 0.12 7.65 8.71 1.70 ± 0.08 8.77 2.98 ± 0.11 Canopy conductance Evaporation from a forest canopy is driven by the same factors that drive evaporation from any wet surface (cf. Equation 1). Thus, low net radiation and VPD may limit plantation water use, and the response to these factors will be tempered by aerodynamic and canopy conductance. We found a close relationship between observed monthly water use and potential evaporation from a wet surface calculated from Equation 1 (Figure 6). The ratio of the vapor deficit term to the radiation term in Equation 1 was greater than five on 76% of days at Jijia and 92% of days at Hetou, with mean values of 8.4 and 14.3, respectively. Although these values are subject to errors in the approximation of net radiation from solar radiation, and of aerodynamic conductance from wind speed at less than canopy height, they confirm the common observation (Jarvis 1981) that evaporation from tall rough canopies is usually dominated by atmospheric vapor deficit. Under such conditions, Equation 2 provides a valid estimate of canopy conductance. Values of gc calculated for all days when the vapor deficit to radiation term ratio was greater than five are plotted against mean daytime VPD in Figure 7. The value of five, chosen as a cutoff point for data filtering, is entirely arbitrary and is clearly affected by the approximations adopted for net radiation and aerodynamic conductance. However, a comparison of gc data calculated for all days with filtered data sets based on cutoff values of three to 10 for the ratio of vapor deficit to radiation terms indicated that a factor of five was effective in removing days when VPD was low enough or radiation high enough to possibly invalidate Equation 2, while retaining a Table 3. Sap flux density and water use by E. urophylla plantations monitored at Hetou and Jijia between September 1999 and September 2000. Figure 5. Sap flux density (A) and daily water use (B) of E. urophylla at Jijia and Hetou, September 1999 –September 2000. Hetou Jijia Days monitored 291 266 Mean sap flux density (dm3 m –2 day –1) ± SE 2772 ± 66 1839 ± 50 Mean daily water use (mm) ± SE 1.49 ± 0.04 1.53 ± 0.04 Annual water use 542 559 TREE PHYSIOLOGY VOLUME 24, 2004 Downloaded from http://treephys.oxfordjournals.org/ by guest on August 29, 2014 compensate for differences in stand sapwood area, such that variation in stand water use is minimized within a region of generally similar environmental conditions. This would further imply that soil and climate factors may wholly determine stand water use, independently of tree density and stand sapwood area, over the range of the plantations studied. However, the similarity of water use at Jijia and Hetou despite their different sapwood areas might be coincidental, resulting from differing responses to environmental conditions at the two sites. It is therefore pertinent to examine the extent to which canopy conductance and hence water use at the Leizhou Peninsula sites are limited by evaporative demand and soil water availability. EUCALYPTUS PLANTATION WATER USE IN CHINA data set containing a broad range of daily climate conditions for analysis. The daily gc data in Figure 7 show the expected decline in canopy conductance with increasing VPD, resulting from the stomatal response to atmospheric humidity (Monteith 1995). The envelope curves in Figure 7, fitted to define the approximate upper limit to observed gc at a given VPD, are described by the functions gc = 0.87(VPD + 0.12) –0.86 at Jijia and gc = 0.75(VPD + 0.12) –0.86 at Hetou. These curves were derived by assuming functions of the form gc = a (b + VPD)c with a maximum conductance of 5 cm s –1 at VPD = 0 kPa, inserting the values of gc and VPD for two points at the upper limit of the observed data and solving the resulting simultaneous equations for a, b and c. Schäfer et al. (2000) proposed an alternative method for fitting envelope curves to similar data sets. The relatively small (16%) difference in VPD response for the two sites is magnified by the higher VPD at Hetou. The tendency for higher VPD at Hetou to lead to higher water use is there- Figure 7. Daily canopy conductance of E. urophylla at Hetou and Jijia plotted against mean daytime vapor pressure deficit (VPD) on days when the ratio of the vapor pressure deficit to radiation terms in Equation 1 was greater than five. Envelope curves are fitted to define the approximate upper limit of observations at each site. fore partly compensated for by reduced stomatal conductance. Over the year, the combination of higher sapwood area, higher stomatal conductance and possibly other factors, led to a mean gc at Jijia that was 1.34 times that at Hetou. If it is assumed that the envelope curves of Figure 7 represent the maximum possible gc at each site under conditions where transpiration is limited only by VPD, then we can calculate the corresponding daily potential water use of the plantations by inverting Equation 2. Figure 8 shows the resulting curves and mean rates of water use observed on days with vapor deficit to radiation term ratios greater than five. Weekly mean values of potential water use are compared with potential evaporation in Figure 9A. Potential water use was usually 10 to 30% of potential evaporation during the wet season, rising to 20 to 50% in the dry season at both sites, and was slightly higher at Jijia than at Hetou all year round. The observed daily rates of water use were 60 to 100% of potential rates during the wet season (Figure 9B), falling during December and January to less than 40%. Although limited data are available for the late dry season, it appears that the recovery of water use to greater than 60% of potential rates occurred during March at Hetou but not until late May at Jijia. Potential water use for the year (calculated as 365 times the daily mean) was 815 mm at Hetou and 916 mm at Jijia. The reduction in estimated actual annual water use below its potential value was therefore 273 mm (33%) at Hetou and 357 mm (39%) at Jijia. The observed difference in soil water availability (Figure 3C) is the most obvious cause of the variations with time and between sites displayed in Figure 9B. To test this hypothesis, the residuals of daily values of gc below the envelope curves of Figure 7 were expressed as a percentage of the potential (envelope) value and compared with soil water content and matric potential at 50-cm depth by regression. Regressions with total available water to 400 cm were also calculated. Highly significant relationships (P < 0.0001) were found for both sites, indicating that low values of gc compared with the envelope value were associated with periods of low soil water content. The nature of these relationships differed between sites. At Jijia, canopy conductance was most strongly correlated with total available soil water (Figure 10A, r 2 = 0.50 for Figure 8. Observed mean daily water use and calculated potential water use at Hetou and Jijia corresponding to the canopy conductance observations and envelope functions in Figure 7. TREE PHYSIOLOGY ONLINE at http://heronpublishing.com Downloaded from http://treephys.oxfordjournals.org/ by guest on August 29, 2014 Figure 6. Monthly values of mean daily water use by E. urophylla at Hetou and Jijia, and mean daily potential evaporation from a wet canopy. Monthly water use at Jijia in April and at Hetou in July were estimated by regression from daily observations at the other site, because insufficient observations were recorded in these months to calculate a reliable monthly mean. 1041 1042 MORRIS ET AL. Figure 9. (A) Weekly means of potential daily water use by E. urophylla plantations at Hetou and Jijia as a percentage of daily potential evaporation. (B) Weekly means of observed daily water use as a percentage of potential water use. 242 days from September to early May), whereas at Hetou, canopy conductance was most strongly correlated with matric potential at 50 cm, and the correlation was significantly increased by a time lag of 2 days between the matric potential and conductance observations (r 2 = 0.22 for 242 days from September to early May). Yin et al. (2003) also reported that correlations between SFD and soil water content were strongest in the lower profile at Jijia but in the upper profile at Hetou. Correlations calculated over the full period of measurement (382 days) that included the wet season of May–September 2000 were stronger at Hetou (Figure 10B, r 2 = 0.29) but weaker at Jijia (r 2 = 0.21). These results suggest that soil water was limiting at Hetou during both wet and dry seasons, but only during the dry season at Jijia. The 2-day lag between canopy conductance and matric potential at 50 cm at Hetou suggests that the major part of the root system was deep in the profile; a comparison of soil moisture sensor responses to individual rain events shows that infiltrating water reached 150 cm one day after reaching 50 cm, and 250 cm after a further 2 days, suggesting a major root presence at around 200 cm depth. At Jijia, good correlations with both total available soil water and soil water content at 50 cm indicate that the upper limit of the zone of root water uptake was shallower than at Hetou, consistent with the greater soil water storage capacity of the clay soil. Discussion Automated data collection at the study sites produced a detailed data set for relating plantation water use to climate and soil conditions in remote locations on the Leizhou Peninsula. Although measurements of net radiation and above-canopy wind speed would have improved the precision of daily potential evaporation estimates, the finding that vapor deficit exceeds radiation by a factor of five or more on most days would be expected to remain true, so that the relationship between plantation water use and climate is adequately described by the relationship with VPD in Equation 2. The heat pulse equipment and techniques adopted for water use measurement were a reliable and convenient means of determining SFD in the range from about 1.5 to over 80 ml cm –2 h –1. The 1127 treedays of water use observations recorded over 12 months at the two sites are expected to provide a satisfactory estimate of mean daily water use and hence seasonal and annual water use. Annual water use at the sites was similar, despite a 53% greater sapwood area at Jijia than at Hetou. The interpretation that SFD varies to compensate for differences in sapwood area, such that variation in stand water use is minimized conflicts with the hypothesis of constant leaf water efficiency advanced by Hatton et al. (1998) and confirmed in terms of SFD by Khanzada et al. (1998), Benyon et al. (1999) and other studies. Furthermore, if soil and climate determine annual evapotranspiration largely independently of vegetation, then the denser understory and higher soil evaporation at Jijia (Lane et al. 2003) would be expected to reduce overstory water use relative to that at Hetou, rather than only reducing SFD. We therefore conclude that the similarity in water use between stands on an annual scale is coincidental, arising from soil and climate differences at the two sites that have compensated for the difference in sapwood area of the plantations studied, and that the observed data do not support the hypothesis of SFD varying to maintain constant stand water use across the region. The greater sapwood area at Jijia would be expected to increase water use by increasing the conductance of the stand as a conduit for water from soil to atmosphere. The difference in conductance, and hence water use, would be less than 50%, TREE PHYSIOLOGY VOLUME 24, 2004 Downloaded from http://treephys.oxfordjournals.org/ by guest on August 29, 2014 Figure 10. (A) Residual canopy conductance (gc; %) at Jijia in relation to total available soil water to 4 m depth (September 1999–May 2000). (B) Residual gc (%) at Hetou in relation to soil matric potential at 50 cm depth, with a 2- day lag time (September 1999–September 2000). EUCALYPTUS PLANTATION WATER USE IN CHINA tations studied. This environmental cost must be weighed against the acknowledged non-wood benefits of the plantations to agriculture including wind shelter, control of erosion and soil loss, and contribution to atmospheric humidity (Zhou et al. 2002). Our results provide some of the information needed for making rational decisions about the management of the tree growing areas of the Leizhou Peninsula and nearby provinces of southern China. Acknowledgments The study was funded by the Australian Centre for International Agricultural Research (ACIAR) as part of project FST 97/77, Eucalypts and Water. We acknowledge the support of the Forest Science Centre, Research Institute of Tropical Forestry (RITF), China Eucalypt Research Centre (CERC) and South China Institute of Botany (SCIB) in making staff and other resources available for the project. We are grateful to the management and staff of Leizhou Forest Bureau, the Tangjia, Jijia and Hetou forest farms, site watchmen and their families, and other members of the Chiniwei and Tiantou workgroups who made it possible to carry out the monitoring reported here over a protracted period in remote locations. Finally, we thank the staff and students of RITF, CERC and SCIB, especially Zhou Guangyi, Wu Zhongmin, Zhou Tao, Hua Lizhong, Chen Shaoxiong, Zhang Kejian, Zhou Guoyi, Li Zhi’an, Shen Weijun, Chu Guowei and Huang Zhihong, for assistance with data collection, establishment and maintenance of the field sites. References Bai, J.Y. and S.M. Gan. 1996. Eucalyptus plantations in China. In Reports Submitted to the Regional Expert Consultation on Eucalyptus, October 1993. Eds. M. Kashio and K. White. RAPA Publication 1996/44, FAO Regional Office for Asia and the Pacific, Bangkok, Thailand, pp 23–32. Baker, T., J. Morris, M. Duncan, N.N. Zhang, Z.J. Yang, Z.H. Huang and G.W. Chu. 2003. Tree and stand growth and biomass relationships for Eucalyptus urophylla and E. 12ABL on the Leizhou Peninsula, Guangdong province, China. In Eucalypts in Asia. Ed. J. Turnbull. ACIAR Proc. No. 111, ACIAR, Canberra, pp 174–182. Benyon, R.G., N.E. Marcar, D.F. Crawford and A.T. Nicholson. 1999. Growth and water use of Eucalyptus camaldulensis and E. occidentalis on a saline discharge site near Wellington, NSW, Australia. Agric. Water Manage. 39:229–244. Calder, I.R., R.L. Hall and P.G. Adlard. 1992. Growth and water use of forest plantations. John Wiley and Sons, Chichester, 381 p. Casson, A. 1997. The controversy surrounding eucalypts in social forestry programs of Asia. 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TREE PHYSIOLOGY ONLINE at http://heronpublishing.com Downloaded from http://treephys.oxfordjournals.org/ by guest on August 29, 2014 because soil, root and canopy components also influence total conductance. Higher VPD at Hetou than at Jijia, particularly during the wet season, would offset this difference by increasing water use, even though canopy conductance would have been lower. In the range of VPDs observed at our sites (usually averaging < 2 kPa during daylight hours), there was no evidence of a reduction in water use as a result of stomatal response to low humidity as observed in more arid conditions (Monteith 1995). The net result of these opposing factors is that the potential water use was estimated as 916 and 815 mm year –1 for Jijia and Hetou, respectively. The effect of limited soil water availability on water use was greater at Jijia than at Hetou, because of a shallower root system and hence greater dependence on water in the upper soil profile. The reduction in actual water use below its potential value amounted to 33% at Hetou and 39% at Jijia, bringing annual water use at both sites to about 550 mm. Statistical analysis of deviations from maximum conductance in relation to soil water availability provided evidence of a strong relationship at both sites. Direct measurements of root density variation with depth in the profile would be required to confirm the interpretation that a deeper root zone at Hetou alleviated the reduction in water use caused by limited soil water availability compared with at Jijia. Water balance studies reported by Lane et al. (2003) found evidence of dry season water uptake from below 4 m depth at Hetou but not at Jijia. Recovery of water use after the dry season in 2000 was later at Jijia than at Hetou, implying a need to replenish the soil water content to some depth before a measurable increase in water use was achieved; lower storage capacity of the sandy soil at Hetou than at Jijia allowed this to occur several weeks earlier, despite the greater root system depth of trees at Hetou. The analysis of potential water use at the study sites allows some qualitative predictions of water use of plantations under other conditions. If the Hetou plantation had the same soil conditions as Jijia, we may infer that its annual water use would be less than 550 mm because of reduced water availability in the dry season, and because the positive influence of higher VPD would still only partly offset the lower sapwood area. On the other hand, if the Hetou plantation was established at the same density as Jijia, its water use would be greater than 550 mm because of the combined effects of higher VPD and higher root zone soil water availability during the dry season than Jijia. For plantation managers seeking to minimize water use, our results indicate that one solution is to plant trees at lower density on sandy soils and in areas with warmer, less humid climates. The possibility that eucalypt plantation water use may impact excessively on groundwater resources on the Leizhou Peninsula remains unresolved. Plantations in the region are usually established on higher ground, with agriculture in the valleys and on lower slopes; hence in most locations there may be limited opportunity for direct uptake of shallow groundwater. 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