WATER RESOURCES RESEARCH, VOL. 38, NO. 6, 1094, 10.

1029/2000WR000122, 2002

Seasonal variations in the evapotranspiration of a transitional

tropical forest of Mato Grosso, Brazil
George L. Vourlitis,1 Nicolau Priante Filho,2 Mauro M. S. Hayashi,3
José de S. Nogueira,2 Fernando T. Caseiro,3 and José Holanda Campelo Jr.3
Received 30 November 2000; revised 24 September 2001; accepted 25 September 2001; published 29 June 2002.

[1] The seasonal pattern of evapotranspiration (expressed as latent heat flux Qe) for a 28-
to 30-m-tall tropical transitional (ecotonal) forest was quantified over an annual cycle
using eddy covariance measurement and micrometeorological estimation techniques. The
study was conducted near the city of Sinop, in northern Mato Grosso, Brazil, which is
located within the ecotone of tropical rain forest and savanna (cerrado). Although the
majority of net radiation (Q*) was consumed by Qe (50–90%), seasonal variations in Qe
were large and positively correlated with precipitation. Total daily Qe for the dry season
(June–August) was on average 6.0 MJ m
2 d
1, while daily Qe for the transition
(October–November and April–May) and wet (December–March) season periods were
7.5 and 10.0 MJ m
2 d
1, respectively. The seasonal variation in midday (0900–1500 LT)
surface conductance (gs) was also positively correlated with precipitation. Analysis of the
‘‘decoupling factor’’ (
) indicated that the forest was strongly coupled to the atmosphere
(
= 0.1–0.3) over the dry season and transition periods, suggesting that Qe was under
relatively strong stomatal control. Although rainfall during the study period was above the
long-term (30-year) average, our results indicate that the seasonal dynamics of Qe for the
tropical transitional forest were more comparable to tropical savanna than to rain
forest. INDEX TERMS: 1818 Hydrology: Evapotranspiration; 1836 Hydrology: Hydrologic budget
(1655); 1851 Hydrology: Plant ecology; 1878 Hydrology: Water/energy interactions; 3374 Meteorology and
Atmospheric Dynamics: Tropical meteorology; KEYWORDS: energy balance, micrometeorology, Priestley-
Taylor alpha, Canopy conductance, decoupling factor

1. Introduction and validation of general circulation (GCM) and mesoscale

models used to assess the effects of deforestation on tropical
[2] Tropical forests and woodlands (savanna) exchange
ecosystem energy exchange.
vast amounts of water and energy with the atmosphere and
[3] Although ARME and ABRACOS generated substan-
are thought to be important in controlling local and regional
tial information on tropical rain forest and savanna micro-
climate [Avissar, 1991; Nobre et al., 1991; Grace, 1992].
meteorology and energy exchange, the tropical transitional
However, rapid deforestation of the Brazilian Amazon [Skole
(ecotonal) forest that separates these two regionally impor-
and Tucker, 1993; Moran et al., 1994] has the capacity to
tant ecosystems has largely been ignored. Aside from
destabilize the surface-atmosphere flux of water and energy
modeling studies [Grace, 1992], little is known about the
[Nobre et al., 1991; Wright et al., 1992; Bastable et al., 1993;
surface-atmosphere exchange of water and energy in the
Polcher and Laval, 1994; Xue et al., 1996]. Research con-
transition zone. However, transitional forests are spatially
ducted during the Amazon Region Micrometeorological
extensive in the Amazon Basin [Murca Pires, 1978; Ratter,
Experiment (ARME) and Anglo-Brazilian Amazonian Cli-
1992] and were subject to rapid rates of deforestation over the
mate Observation Study (ABRACOS) assessed the effects of
last 2 decades [Skole and Tucker, 1993; Moran et al., 1994].
deforestation on mass and energy exchange by quantifying
[4] To reduce uncertainty regarding tropical transitional
the micrometeorology [Shuttleworth et al., 1984a; Wright
forest energy and water cycling, measurements of mass
et al., 1992; Bastable et al., 1993; Culf et al., 1996], water
(H2O vapor) and energy were initiated in August 1999 as
balance [Shuttleworth et al., 1984b; Shuttleworth, 1988;
part of the NASA/INPE Large-Scale Biosphere-Atmosphere
Grace et al., 1995; Hodnett et al., 1995; Miranda et al.,
Experiment in Amazonia (LBA) [Cerri et al., 1995]. Here
1997], and physiology [Dolman et al., 1991; Sa et al., 1996]
we describe variations in evapotranspiration (expressed as
of rain forest, savanna, and cattle pasture. Data from these
latent heat flux, Qe) over the dry (June –September), wet
studies form the basis for the parameterization, calibration,
(December – March), and transition (October – November
and April – May) hydrologic periods and propose mecha-
1
Biological Sciences Department, California State University, San nisms for the temporal variations in the surface-atmosphere
Marcos, California, USA.
2
Departamento de Fı́sica, Universidade Federal de Mato Grosso, Cuiabá,
exchange of H2O vapor.
Mato Grosso, Brazil.
3
Departamento de Fı́sica, Núcleo de Tecnologia em Armazenagem, 2. Methods
Universidade Federal de Mato Grosso, Cuiabá, Mato Grosso, Brazil.
2.1. Site Description
Copyright 2002 by the American Geophysical Union. [5] The study was conducted near the city of Sinop, Mato
0043-1397/02/2000WR000122 Grosso, Brazil (11
24.750S, 55
19.500W), in an area of
30 - 1
30 - 2 VOURLITIS ET AL.: TROPICAL FOREST EVAPOTRANSPIRATION

Figure 1. Map of Brazil and false-color image of the study area located 50 km N of the city of Sinop, Mato
Grosso, Brazil (11
24.750:55
19.50W). The location of the eddy covariance tower and study site is visible
on the false-color image as filled white circle. Mature, intact forest is visible as dark red surfaces, secondary
forest is visible as light red surfaces, and pasture is visible as pink, white and light blue surfaces. Note the
logging roads and "patios" (light blue trails and clearings, respectively) associated with selective logging.
The shaded area on the map displays the approximate boundaries of the Brazilian Legal Amazon. The false-
color image is produced from IKONOS imagery courtesty of the University of New Hampshire. Image
courtesy of the University of New Hampshire’s EOS-WEBSTER Earth Science Information Partner (ESIP)
(http://www.eos-webster.sr.uhn.edu).

extensive mature, intact forest, selectively logged forest,
and pasture (Figure 1). The measurements were con-
ducted in intact, mature forest with a relatively continu-
ous, 28- to 30-m-tall canopy (Figure 2). The terrain is
generally flat within approximately 5 km of the tower,
and the fetch (the upwind distance sampled by the eddy
covariance sensor array) is dominated by a relatively
continuous canopy of intact mature forest for at least
1.5 km in all directions.
[6] Soils of the study site are largely sandy, nutrient-poor
(dystrophic) ultisols [Ratter, 1992], which are common to
the Amazon Basin [Moraes et al., 1995]. These soils have
high porosity and drain rapidly following rainfall events
(i.e., within 4– 7 days) (P. Girard et al., unpublished data,
2001). The vegetation consists of evergreen trees that are
characteristic of transitional forest in Mato Grosso [Ratter,
1992], such as Qualea sp., Vochysia sp., Ocotea sp., and
Mezilaurus itauba [Vourlitis et al., 2001], and diversity is
 VOURLITIS ET AL.: TROPICAL FOREST EVAPOTRANSPIRATION
Figure 2. Schematic of the eddy covariance/micrometeor-
ology measurement system used to measure the latent heat
flux (Qe) of the transitional tropical forest near Sinop, Mato
Grosso. Shown are the locations of the vertical wet-bult (Tw)
and dry-bulb (Td) temperature measurements, net radio-
meter (Q*), tipping-bucket rainfall gauge (PPT), and ground
heat flux transducers (Qg). The eddy covariance array,
consisting of the open-path infrared gas analyzer for
measuring the mean and fluctuating quantities of water
vapor (H2OEC), and the three-dimensional sonic anem-
ometer for measuring the mean and fluctuating quantities of
virtual temperature (Tv) and wind speed (u, v, and w), was
mounted at a height of 42 m above ground level. Canopy
height is 28– 30 m.
high so there are no obvious dominant tree species.
Overstory trees are 28– 30 m, and although the canopy
is relatively closed and continuous, understory trees (1 –
5 m tall) are widespread especially in gaps formed by fallen
and/or dead overstory trees. Leaf area index (LAI), esti-
mated from measurements of the extinction of photosyn-
thetic photon flux density by the forest canopy [Goudriaan,
1988], reaches a maximum of 4.5– 5 during the wet season
(January) and a minimum of 4 – 4.5 during the dry season in
July (G. Vourlitis et al., The role of seasonal variations in
30 - 3
meteorology on the net CO2 exchange of a Brazilian
transitional tropical forest, submitted to Ecological Appli-
cations, 2001, hereinafter referred to as submitted manu-
script, 2001). The decline in LAI during the transition from
the wet to the dry season corresponds with an increase in
leaf and stem litter production (G. Vourlitis et al., submitted
manuscript, 2001). The transitional forest LAI contrasts
with a maximum LAI for tropical rain forest of 5– 6 [Malhi
et al., 1999] and 1 for savanna [Miranda et al., 1997].
[7] The 30-year mean annual temperature in the Sinop
area is 24
C with little seasonal variation, and rainfall is
approximately 2000 mm yr
1, with a 4-month dry season
between June and September (Table 1). The seasonal
climatology for the transitional forest is similar to rain forest
and savanna; however, transitional forest typically receives
about 200 mm less rainfall per year than rain forest and 500
mm more rainfall than savanna (Table 1). Average air
temperature is similar for transitional and rain forests;
however, savanna is typically 2
– 3
C cooler than transi-
tional forest (Table 1).
2.2. Eddy Covariance Instrumentation and Data
Treatment
[8] Latent heat flux (Qe) and sensible heat flux (Qh) were
quantified using tower-based eddy covariance [Baldocchi
et al., 1988; Verma, 1990; Vourlitis et al., 2001]. This
micrometeorological technique directly quantifies the sur-
face-atmosphere exchange of mass and energy by measur-
ing the turbulent transport of H 2 O vapor and heat
[Baldocchi et al., 1988; Verma, 1990]. Eddy covariance
sensors were mounted at a height of 42 m above ground
level or 12– 14 m above the forest canopy (Figure 2). The
eddy covariance system utilized a three-dimensional sonic
anemometer-thermometer (SWS-211/3K, Applied Technol-
ogies, Inc., Boulder, Colorado) and an open-path infrared
gas analyzer (NOAA-ATDD, Oak Ridge, Tennessee) to
measure the mean and fluctuating quantities of wind speed
and temperature and H2O vapor, respectively. Both sensors
sampled and output data at 10 Hz and were physically
oriented into the direction of the mean wind at the upwind
side of the tower to minimize the potential for flow
distortion from the tower. The H2O vapor channel of the
gas analyzer was calibrated every 2 – 4 weeks using a
portable dew-point generator (LI-610, LI-COR, Inc., Lin-
coln, Neb-raska).
[9] Raw H2O vapor fluctuations were output as mean
voltages and converted to densities by multiplying by the
requisite calibration constant [Leuning and Moncrieff,
1990], and H2O vapor and sensible heat fluxes were
computed following a coordinate rotation of the wind
vectors [McMillen, 1986, 1988]. Fast response (10-Hz)
fluxes were calculated and stored on a laptop computer as
30-min averages using a 200-s running mean and digital
recursive filtering technique. Water vapor flux was corrected
for the simultaneous fluctuations in heat [Webb et al., 1980].
2.3. Micrometeorological Measurements and Data
Treatment
[10] Net radiation (Q*) was measured above the canopy
(40 m above ground level; Figure 2) using a ventilated net
radiometer (Q*7.1, REBS, Inc., Seattle, Washington). Soil
heat flux was measured using heat flux transducers (n = 2)
buried approximately 2 cm into the surface litter layer
30 - 4 VOURLITIS ET AL.: TROPICAL FOREST EVAPOTRANSPIRATION

Table 1. Monthly Summary of the Average Daily Air Temperature and Total Precipitation During the Study Period and for Tropical
Transitional Forest (TF), Rain Forest (RF), and Savanna (S)a
Total Monthly Precipitation, mm Average Daily Air Temperature, °C

99 – 00 RF TF S 99 – 00 RF TF S

Aug. 0 38 10 12 23.9 26.1 23.2 21.2

Sept. 71 120 71 52 25.3 26.3 24.6 22.5
Oct. 195 190 189 172 25.4 26.2 25.6 22.1
Nov. 235 230 267 238 24.4 26.1 24.9 21.7
Dec. 139 322 365 249 24.6 25.5 24.5 21.5
Jan. 704 312 320 241 24.6 25.5 24.5 21.6
Feb. 394 306 347 215 24.6 25.5 24.3 21.8
March 153 266 266 189 25.0 25.5 24.7 22.0
April 204 251 139 124 24.9 25.6 24.8 21.4
May 0 132 45 39 24.5 25.4 23.9 20.2
June 0 49 15 9 24.5 24.9 22.7 19.1
July 0 27 3 12 22.8 25.0 21.9 19.1
Annual 2095 2243 2037 1552 24.6 25.6 24.1 21.2
a
Data for the study period are for 15 August 1999 to 31 July 2000 (99 – 00), while data for TF are 30-year averages from Vera, Mato Grosso located
approximately 60 km southeast of Sinop (N. Priante-Filho et al., unpublished data, 2001). Data for S are 27-year averages from Brasilia, Districto Federal
[Miranda et al., 1997], and data for RF are 30-year averages from Porto Velho, Rondonia (N. Priante-Filho et al., unpublished data, 2001).

(HFT-3.1, REBS, Inc., Seattle, Washington). Air temper-
ature and vapor pressure were measured at the top of the
tower (40 m above ground level; Figure 2) using a relative
humidity sensor (HMP-35, Vaisala, Inc., Helsinki, Finland).
After approximately 1 March 2000, the vertical vapor
pressure profile was also measured using wet- and dry-bulb
psychrometers mounted at heights of 1, 4, 12, 20, 28, and
40 m above ground level (Figure 2). Vapor pressure deficit
of the atmosphere (VPD) at the top of the tower was
calculated as the difference between saturation vapor pres-
sure (es) and actual vapor pressure (ea) from temperature
and humidity data collected by the relative humidity sensor
and/or the wet- and dry-bulb psychrometer. Precipitation
was measured at the top of the tower using a tipping-bucket
rainfall gauge (2501, Sierra-Misco, Inc., Berkeley, Califor-
nia). Micrometeorological data were averaged over 30-min
intervals from observations made every 60 s and stored
using a solid-state data logger (CR-10, Campbell Scientific,
Inc., Ogden, Utah).
2.4. Statistical Analysis and Derived Quantities
[11] Diurnal Q* and Qe were summarized as diurnal
averages calculated over seasonal intervals. Diurnal aver-
ages were calculated by averaging each 30-min energy flux
density measurement for a particular time (e.g., 0900 –0930
LT) over the dry (June – September), transition (October –
November and April – May), or wet (December – March)
season periods. This averaging process was conducted to
allow the utilization of all data collected and to provide an
indication of how the diurnal trend in energy flux density
varied, on average, during the different hydrological peri-
ods. Daily total values of Qe, Q*, and precipitation were
summed over a daily (24-hour) period.
[12] Total daily Qe and the evaporative fraction (Qe/Q*)
were bootstrapped over monthly, intraseasonal (2-month),
and seasonal intervals to estimate the random variance
(95% confidence interval) about the mean [Efron and
Tibshirani, 1993]. The bootstrap calculated a confidence
interval by (1) constructing 4000 bootstrapped sample data
series by randomly sampling (with replacement) the
observed average daily Qe and Qe/Q* time series, (2)
calculating an average Qe or Qe/Q* from each constructed
sample data series, and (3) calculating the grand mean
±95% CI from the distribution of averages calculated from
the bootstrapped sample data series [Efron and Tibshirani,
1993].
[13] Least squares linear regression was used to assess the
degree of energy balance closure and the performance of the
eddy covariance flux system [McMillen, 1986, 1988]. For
each day, the sum of the instantaneous (i.e., 30-min average)
sensible and latent heat flux (Qh + Qe) measured from the
eddy covariance system (n = 48) was regressed against the
difference between the instantaneous net radiation and
ground heat flux (Q*
Qg) measured from the micro-
meteorological sensors (n = 48) [Vourlitis et al., 2001]. This
yielded a slope and intercept quantifying the degree of
energy balance closure for each day of the measurement
campaign. A slope of 1 and an intercept of 0 J m
2 s
1
indicates complete closure of the energy balance and
excellent performance of the measurement system
[McMillen, 1986, 1988]. Results indicate that the slope of
the regression was on average (±1 standard deviation) 0.92
± 0.03 and the intercept was 11.03 ± 14.67 J m
2 s
1 (n =
72 individual regressions) [Vourlitis et al., 2001], which are
well within the range reported for other forest measurement
systems [e.g., Goulden et al., 1996]. These data indicate
satisfactory performance of the eddy covariance measure-
ment system.
[14] Power system failure limited the amount of data
collected from the eddy covariance system. However, the
micrometeorological system used a separate power system
and was operational more frequently (75% of the total
possible time) than the eddy covariance system (26% of
the total possible time). Therefore data collected from the
more consistent micrometeorological measurements, along
with the direct measurements of Qe from the eddy cova-
riance system, were used to estimate Qe for the measure-
ment period. Instantaneous (30-min average) rates of Qe
were calculated from micrometeorological data using the
Priestley and Taylor [1972] expression,

 
Qe ¼ a s=ðs þ gÞ Q*
Qg ; ð1Þ
 VOURLITIS ET AL.: TROPICAL FOREST EVAPOTRANSPIRATION 30 - 5

the eddy covariance array, and Q*, Qg, and vapor pressure
measured from the micrometeorological instruments. Aero-
dynamic conductance (ga) was calculated as [u/(u*)2]
1
corrected for atmospheric stability [Grace et al., 1995],
where u is wind speed measured from the triaxial sonic
anemometer and u* is frictional velocity calculated from
eddy covariance measurements of momentum flux [Baldoc-
chi et al., 1991]. The average midday (0900 – 1500 LT)
‘‘decoupling factor’’ (
) [Jarvis and McNaughton, 1986]
was calculated to assess the relative importance of physio-
logical and meteorological limitations to canopy water
vapor exchange.

3. Results
3.1. Correspondence Between the Long-Term
Average and Observed Meteorology
[17] Temperature and precipitation during the study
period (15 August 1999 to 31 July 2000) were similar
to the long-term (30-year) average; however, approxi-
mately 25% of the micrometeorological data were missing
Figure 3. Estimates of the calibrated Priestley-Taylor due to system failure. Total annual rainfall during the
alpha (a) calculated from equation (2) (circles) using least study period was 2095 mm compared with the long-term
squares linear regression between Qe measured from eddy average of 2037 mm (Table 1), but with missing precip-
covariance (dependent variable) and (s/s + g)(Q*
Qg) itation data, it is likely that the study period was substan-
derived from micrometeorological data (independent vari- tially wetter than the long-term average. However, the
able). The curve depicts interpolated values of a estimated seasonal distribution of rainfall during the study was
using a polynomial function that was fit to the data using identical to the long-term average. For example, most of
nonlinear, least squares regression. the rainfall recorded during the study period (66%)
occurred during the wet season (December –March), while
wet season rainfall historically accounts for 65% of the
where a is the Priestley-Taylor coefficient (usually taken as total annual rainfall (Table 1). Similarly, rainfall recorded
1.26), s is the slope of the saturation vapor pressure versus during the dry season (June – September) and transition
temperature curve at the appropriate temperature, g is the periods (October – November and April – May) accounted
psychrometric constant, and Q* and Qg are the net radiation for 4 and 30%, respectively, of annual rainfall, which is
and ground heat flux measured from the micrometeorolo- similar to the long-term seasonal distribution (Table 1). Air
gical sensors. The advantage of the Priestley-Taylor temperature was on average 24.6
C during the study
relationship is that few data (Q*, Qg, and temperature) are period, compared with a long-term average temperature
required to estimate the Qe of saturated surfaces [Priestley of 24.1
C (Table 1).
and Taylor, 1972]. However, the theoretical basis of a is
unclear, and a can vary substantially depending on canopy 3.2. Temporal Dynamics of the Calibrated
roughness and surface water content [Shuttleworth et al., Priestley-Taylor A
1984a; Xu and Singh, 2000]. [18] Estimates of a calculated from the eddy covariance
[15] Given a known Qe, such as that measured from the measurements of Qe (equation (2)) ranged between 0.57
eddy covariance system, a can be estimated by rearranging
equation (1) [Priestley and Taylor, 1972], where
nh i o Table 2. Estimates of the Priestley-Taylor Alpha (a) Calculated as
a ¼ Qe s=ðs þ gÞ Q*
Qg : ð2Þ the Slope of the Least Squares Linear Regression With Latent Heat
Flux Measured From the Eddy Covariance System (Qe) as the
With (2), monthly values of a were estimated for the _
Dependent Variable and [(s/s + g)](Q*
Qg) as the Independent
annual measurement period using least squares linear Variablea
regression with instantaneous (30-min average) Qe mea- nb a (±1 SD) r2
sured from the eddy covariance system as the dependent
variable and instantaneous [s/(s + g)](Q*
Qg) measured Aug. 1999 700 0.57 ± 0.01 0.86
from the micrometeorological data as the independent Sept. 1999 561 0.58 ± 0.01 0.77
Oct. 1999 1174 0.65 ± 0.01 0.83
variable. The calibrated values of a were interpolated Nov. 1999 653 0.74 ± 0.01 0.88
between August 1999 and July 2000 using a polynomial Feb. 2000 96 1.07 ± 0.06 0.77
function to provide an estimate of a for the measurement April 2000 142 0.84 ± 0.03 0.82
period (Figure 3). July 2000 394 0.59 ± 0.01 0.86
[16] Average midday (0900 – 1500 LT) surface conduc- a
See equation (2).
tance (gs) was estimated by inversion of the Penman-Monteith b
Here n is the number of 30-min observations used in the linear
equation [Baldocchi et al., 1991], with Qe measured from regression.
30 - 6 VOURLITIS ET AL.: TROPICAL FOREST EVAPOTRANSPIRATION
Figure 4. (a) Latent heat flux (Qe) measured from eddy
covariance (Qe(EC); independent variable) versus Qe
estimated by the calibrated Priestley-Taylor expression
(equations (1) and (2) (Qe(PT); dependent variable). (b)
Sensible heat flux (Qh) measured from eddy covariance
(Qh(EC); independent variable) versus Qh estimated by the
calibrated Priestley-Taylor expression (Qh(PT); dependent
variable). The solid line depicts the best fit line estimated
from least squares, linear regression, and the dashed line
depicts the 1:1 relationship. The equations correspond to the
slope, intercept, and coefficient of determination (r2) of the
regression. Data are 30-min. averages (n = 3729 observa-
tions).
during the dry season (August) and 1.07 during the wet
season in February (Table 2). The maximum value of a
(1.07) was well below the value of 1.26 for saturated
surfaces suggested by Priestley and Taylor [1972] but is
similar to the value estimated by Viswanadham et al. [1991]
(1.03) for Amazonian rain forest. Although data were
limited, the seasonal variation in estimated monthly a was
highly correlated with total monthly rainfall (a = 0.54 +
0.0012  rainfall; r2 = 0.84; p < 0.01; n = 7). The close
correspondence between a and precipitation is expected
given that a relates evapotranspiration to surface features
such as the amount of available water [Priestley and Taylor,
1972]. These data suggest that seasonal (and possibly
interannual) variations in a can be estimated from relatively
simple data such as total monthly rainfall.
3.3. Correspondence Between Eddy Covariance and
Priestley-Taylor Qe
[19] Instantaneous (30-min average) estimates of latent
heat flux (Qe) calculated from the calibrated Priestley-
Taylor (PT) expression (equations (1) and (2)) were
similar to those measured directly from eddy covariance
(EC) (Figure 4a). Although there was substantial varia-
bility about the 1:1 line and few actual measured data,
the slope of the regression was 0.91 and the intercept was

1.7 J m
2 s
1 (Figure 4a), indicating that on average,
Qe(PT) underestimated Qe(EC) by <10% at the highest
levels of Qe(EC). The relatively close correspondence
between Qe(PT) and Qe(EC) was expected considering
that a was calibrated from Qe(EC) (equation (2)), and
much of the variability about the regression was presum-
ably due to the fact that the measured values of Qe from
eddy covariance were sensitive to variations in evapora-
tive demand (VPD) and wind speed and direction, while
the estimated values of Qe from Preistley-Taylor were
insensitive to these environmental fluctuations [Priestley
and Taylor, 1972; Shuttleworth et al., 1984a; Xu and
Singh, 2000]. Similar results were obtained in the regres-
sion between Qh(PT) and Qh(EC), as the slope of the
regression was 0.95 and the intercept was 19.9 J m
2 s
1
(Figure 4b).
3.4. Temporal Trends in Qe and the Evaporative
Fraction (Qe/Q*)
[20] There were large differences in the magnitude of
diurnal Qe observed over the measurement period. For
example, peak daytime values of Qe averaged over the
dry season were 270 J m
2 s
1 (Figure 5a). Diurnal
temperature fluctuations and daytime VPD were highest
during the dry season, as the diurnal temperature range was
14
C and the maximum midday VPD was 3 kPa (Figure
5d). Rates of midday Qe increased to 320 J m
2 s
1 during
the transition period (Figure 5b). Diurnal temperature and
VPD fluctuations were somewhat smaller during the tran-
sition period, as the average diurnal range in air temperature
was 8
C and the maximum midday VPD was 1.75 kPa
(Figure 5e). Maximum midday rates of Qe averaged over
the wet season were 410 J m
2 s
1 (Figure 5c and Table 3).
These high rates of wet season Qe coincided with the lowest
average daytime Q* (Figure 5c), ambient temperature, and
VPD observed over the study period (Figure 5f).
[21] Rates of total daily Qe were 4 –6 MJ m
2 d
1 during
the dry season, and 50% of the total daily Q* was
dissipated by Qe (Table 3). Rates of daily Qe at the end
of the dry season (August – September) varied substantially
(2 – 8 MJ m
2 d
1; Figure 6a) presumably because of
fluctuations in average daily Q* (Figure 6b). August –
September corresponded to the peak of the biomass burning
season in Mato Grosso, which can reduce incident radiation
by 20– 45% and cause substantial variations in daily Q* [Eck
et al., 1998]. The atmospheric evaporative demand (meas-
ured as the average daily VPD) was at a seasonal maximum
at the end of the dry season (Figure 6c).
[22] The first precipitation event was recorded during
the second week of September, but this and subsequent
 VOURLITIS ET AL.: TROPICAL FOREST EVAPOTRANSPIRATION 30 - 7

Figure 5. (a– c) Average diurnal trend in net radiation (Q*, solid curves), latent heat flux measured by
eddy covariance (Qe(EC), solid circles), and latent heat flux estimated from the calibrated Priestley-
Taylor expression (equations (1) and (2)) (Qe(PT); open circles). (d – f) Average diurnal trend in air
temperature (solid curves) and vapor pressure deficit (VPD, open diamonds) measured at the top of the
tower (40 m above ground level). Data represent means ±1 standard deviation calculated over the dry
season (June – September, Figures 5a and 5d), transition (October – November and April – May, Figures 5b
and 5e), and the wet season hydrologic periods (December – March, Figures 4c and 4f).

events were relatively small (<20 mm d
1) and infrequent
until late October (Figure 6d). As rainfall increased in
late October (Figure 6d), the VPD declined as well
(Figure 6c), and depending on Q*, total daily Q e
increased to on average 6 – 8 MJ m
2 d
1 (Figures 6a
and 6b). When averaged over the October – November
transition period, total daily Qe was 6.8 ± 0.5 MJ m
2 d
1
and approximately 65% of daily Q* was dissipated by Qe
(Table 3).
[23] Total daily Qe was on average 10 MJ m
2 d
1
during the wet season, and nearly 90% of the total daily
Q* was dissipated by Qe (Table 3). Although Qe increased
rapidly with rainfall, day-to-day variations in the estimated
rate of Qe were substantial (Figure 6a). Total daily Qe was
estimated to be 10– 12 MJ m
2 d
1 in December– February
(Figure 6a), when average daily VPD was at an annual
minimum (Figure 6c) and rainfall was at an annual max-
imum (Figure 6d).

Table 3. Total Daily Qe and the Fraction of Total Daily Q* Dissipated by Qe(Qe/Q*) Averaged Over Monthly, Seasonal, and Annual
Intervalsa
Eddy Covariance Priestley-Taylor

2
1
n, days Qe, MJ m d Qe/Q* n, days Qe, MJ m
2 d
1 Qe/Q*

Aug. 1999 17 5.58 ± 1.68 0.56 ± 0.16 17 4.20 ± 0.67 0.42 ± 0.02
Sept. 1999 30 6.42 ± 1.66 0.80 ± 0.53 30 3.65 ± 1.72 0.46 ± 0.02
Oct. 1999 31 7.22 ± 1.50 0.67 ± 0.20 31 6.67 ± 1.94 0.54 ± 0.03
Nov. 1999 17 6.91 ± 2.61 0.76 ± 0.39 17 5.99 ± 2.18 0.61 ± 0.02
Dec. 1999 0 ND ND 20 9.93 ± 3.62 0.77 ± 0.02
Jan. 2000 0 ND ND 31 10.36 ± 3.74 0.81 ± 0.02
Feb. 2000 3 10.45 ± 1.34 0.97 ± 0.18 16 8.34 ± 4.18 0.80 ± 0.02
March 2000 0 ND ND 16 10.14 ± 3.12 0.71 ± 0.02
April 2000 2 9.27 ± 0.28 0.63 ± 0.07 30 8.77 ± 1.69 0.64 ± 0.03
May 2000 0 ND ND 31 8.08 ± 0.90 0.54 ± 0.03
June 2000 0 ND ND 30 6.36 ± 0.89 0.46 ± 0.02
July 2000 7 6.73 ± 0.67 0.43 ± 0.08 14 5.95 ± 0.99 0.42 ± 0.01
Dry 34 6.22 ± 1.02 0.57 ± 0.12 91 5.00 ± 0.70 0.45 ± 0.01
Transition 44 7.20 ± 1.04 0.70 ± 0.16 110 7.52 ± 0.76 0.58 ± 0.02
Wet 3 10.45 ± 2.42 0.97 ± 0.33 83 9.83 ± 1.55 0.78 ± 0.02
Annual 81 6.91 ± 0.80 0.66 ± 0.11 284 7.39 ± 0.89 0.60 ± 0.02
a
Data are means ±95% confidence intervals calculated by bootstrapping (see text). ND means no data.
30 - 8 VOURLITIS ET AL.: TROPICAL FOREST EVAPOTRANSPIRATION

Figure 6. (a) Total daily latent heat flux (Qe) measured fromeddy covariance (shaded circles, shaded
curve) and estimated from the calibrated Priestley-Taylor expression (equations (1) and (2)) (solid curve),
(b) total daily net radiation (Q*), (c) average daily air temperature (solid curve) and vapor pressure deficit
(VPD, dashed curve) measured at the top of the tower (40 m above ground level), and (d) total daily
precipitation for the measurement period.

[24] Total daily Qe declined during the April – May tran-
sition period and was <8 MJ m
2 d
1 by July (Figure 6a
and Table 3). During this period, Q* was on average
14 MJ m
2 d
1 (Figure 6b) and VPD increased from
<0.5 kPa in April – May to 1 kPa by July 31 (Figure 6c).
April – May precipitation was <50 mm d
1, and after 27
April, no precipitation was recorded for the remainder of
the study period (Figure 6d). However, estimated rates of
Qe during the April – May transition period were signifi-
cantly higher than those observed during the October –
November transition period (Table 3). Although the decline
in precipitation during the transition period was presumably
the proximate cause for the decline in Qe over the April –
May transition period, the relatively high rates of Qe
suggest that wet season rainfall significantly enhanced Qe
into the dry season.
3.5. Surface Conductance and Atmospheric Coupling
[25] The seasonal variations in Qe imply concomitant
variations in stomatal (gst) and surface conductance (gs). To
partially assess this hypothesis, the seasonal trend in midday
(0900 – 1500 LT) gs was estimated by inversion of the Pen-
man-Monteith equation. When midday gs was averaged over
monthly intervals, there was a consistent increase in gs from
the dry to the wet season, as late dry season (August –
September) rates of gs were between 2 and 3 mm s
1 and
 VOURLITIS ET AL.: TROPICAL FOREST EVAPOTRANSPIRATION 30 - 9

Table 4. Average (±1 Standard Deviation) Daytime (0900 – 1500 LT) Wind Speed (u), Frictional Velocity (u*), Aerodynamic
Conductance (ga), Surface Conductance (gs), and the ‘‘Omega Factor’’ (
) Calculated Over Monthly Intervalsa
n, days u, m s
1 u*, m s
1 ga(EC), mm s
1 gs(EC), mm s
1 gs(PT), mm s
1
(EC)
(PT)

Aug. 1999 17 2.38 ± 0.75 0.53 ± 0.18 127.3 ± 41.1 2.0 ± 0.5 2.1 ± 0.5 0.10 ± 0.05 0.11 ± 0.05
Sept. 1999 12 1.51 ± 0.49 0.34 ± 0.15 88.2 ± 48.0 2.7 ± 1.4 3.0 ± 2.0 0.21 ± 0.12 0.22 ± 0.09
Oct. 1999 25 1.65 ± 0.43 0.35 ± 0.09 95.6 ± 33.9 3.6 ± 3.4 3.7 ± 2.2 0.23 ± 0.11 0.22 ± 0.07
Nov. 1999 12 1.81 ± 0.49 0.35 ± 0.09 94.6 ± 36.0 7.9 ± 4.2 5.5 ± 2.6 0.29 ± 0.11 0.27 ± 0.09
Feb. 2000 3 1.06 ± 0.53 0.39 ± 0.13 156.4 ± 19.2 11.1 ± 1.8 7.6 ± 1.5 0.51 ± 0.03 0.47 ± 0.01
April 2000 3 2.50 ± 0.24 0.30 ± 0.06 42.8 ± 14.9 9.2 ± 0.3 9.2 ± 1.5 0.24 ± 0.07 0.25 ± 0.11
July 2000 7 2.59 ± 0.77 0.43 ± 0.23 85.0 ± 49.7 7.1 ± 3.7 8.8 ± 4.9 0.29 ± 0.07 0.35 ± 0.10
a
Surface conductance was calculated from inversion of the Penman-Monteith equation (see text).

wet season rates of gs were on average 9 mm s
1 (Table 4).
As with Qe, the transition period following the wet season
(April – May) had substantially higher rates of gs than the
transition period before the wet season (October – Novem-
ber), suggesting that wet season rainfall stimulated gs long
after the cessation of rainfall in late April.
[26] The close correspondence between Qe and gs sug-
gests that limitations to gst played an important role in the
seasonal variations in Qe. At the scale of the canopy,
however, gs may not be directly synonymous with gst
because gs is also sensitive to nonphysiological processes
such as the net radiation balance, aerodynamic conductance
inside the canopy, and surface evaporation [Baldocchi et al.,
1991]. Furthermore, rates of Qe may be relatively unre-
sponsive to variations in gst if low wind speed and/or
canopy properties cause aerodynamic conductance (ga) to
be low relative to gs [Jarvis and McNaughton, 1986]. Under
these conditions the canopy atmosphere will be relatively
uncoupled from the overlying atmosphere, so changes in gst
will have a smaller influence on Qe [Jarvis and McNaugh-
ton, 1986]. Given this, the decoupling factor (
) [Jarvis and
McNaughton, 1986] was calculated to assess the relative
importance of physiological (i.e., stomatal) versus physical
(VPD, Q*) limitations to Qe.
ranged between 0.1 and 0.2
during the late dry season (August – September) and
increased only slightly during the transition period of
October and November (Table 4), suggesting close coupling
between the canopy and the overlying atmosphere. Because
average daytime ga was at least an order of magnitude
greater than gs (Table 4), the low values of daytime

indicate that physiological limitations to Qe were relatively
more important than physical limitations during the dry and
transition periods [Jarvis and McNaughton, 1986]. During
the wet season, however, midday values of
were on
average 0.5, and
remained relatively high (i.e., 0.3) into
July (Table 4). The increase in
suggests that the relative
importance of physical limitations to Qe increased during
the wet season.
4. Discussion
4.1. Comparisons With Other Studies
[27] Maximum rates of midday Qe during the dry season
are reportedly 400 J m
2 s
1 for rain forest [Shuttleworth,
1988; Roberts et al., 1993] and 150 J m
2 s
1 for savanna
(cerrado) [Miranda et al., 1997; Meinzer et al., 1999].
Maximum rates of midday Qe during the wet season are
reported to be 300 – 400 J m
2 s
1 for rain forest [Shuttle-
worth, 1988; Roberts et al., 1993] and 400 J m
2 s
1 for
cerrado [Miranda et al., 1997; Meinzer et al., 1999]. The
maximum midday rate of Qe estimated for transitional forest
during the dry season (260 J m
2 s
1; Figure 5) was
intermediate to that reported for tropical savanna and rain
forest, while wet season rates for all three tropical ecosys-
tems were nearly identical. Similarly, surface conductance
(gs) during the dry season reportedly varies between 1.5 and
2.5 mm s
1 for cerrado [Miranda et al., 1997; Meinzer et al.,
1999] and 2 and 5 mm s
1 for rain forest [Meinzer et al.,
1995, 1997; Andrade et al., 1998]. Although wet season
rates of gs for tropical rain forest are lacking, average
daytime gs for cerrado was reported to be 7 mm s
1
[Miranda et al., 1997], which is on the low side of what
was estimated for transitional forest (e.g., 7.5– 11 mm s
1;
Table 4). As with Qe, dry season gs estimated for the
transitional forest (2 – 3 mm s
1; Table 4) was intermediate
to that reported for tropical savanna and rain forest. Thus,
even though the study period was wetter than the long-term
average (see Table 1), the seasonal pattern of midday Qe and
gs for transitional tropical forest appeared to be more similar
to savanna than to rain forest.
4.2. Physiological and Environmental Regulation of
Transitional Forest Qe
[28] Results from a variety of studies indicate that rain
forest and cerrado experience declines in soil water content,
leaf water potential, and leaf area index (LAI) during the dry
season [Hodnett et al., 1995; McWilliam et al., 1996;
Roberts et al., 1996; Sa et al., 1996; Meinzer et al.,
1999]. Although emergent trees may have access to deep
water reserves [Nepstad et al., 1994; Hodnett et al., 1996],
declines in soil water content can limit evaporation and
water availability to shallow rooted plants [Hodnett et al.,
1995; Sa et al., 1996; Meinzer et al., 1999]. The decline in
ecosystem water content and LAI can lead to enhanced
surface heating, an increase in sensible heat flux and
evaporative demand [Grace, 1992; Bastable et al., 1993],
and declines in gst and gs [Roberts et al., 1993; Sa et al.,
1996; Meinzer et al., 1993; Miranda et al., 1997]. Each of
these processes can individually and/or collectively limit
rates of Qe.
[29] High rainfall during the wet season can promote leaf
production, increase plant water potential, and lower surface-
atmosphere vapor pressure deficit [Roberts et al., 1993;
Hodnett et al., 1995; McWilliam et al., 1996; Miranda et
al., 1997]. These seasonal changes in microclimate and
physiology are reportedly the cause of the large increase in
wet season Qe observed for cerrado [Miranda et al., 1997].
In rain forest, however, frequent could cover during the wet
30 - 10 VOURLITIS ET AL.: TROPICAL FOREST EVAPOTRANSPIRATION
season effectively limits Q*, and as a result, Qe [Shuttle-
worth, 1988; Bastable et al., 1993; Roberts et al., 1993]. Thus
in rain forest, dry season limitations to Qe and gs are balanced
by wet season limitations to Qe caused by declines in Q*,
which dampen the annual variations in Qe and gs [Shuttle-
worth, 1988; Bastable et al., 1993; Roberts et al., 1993].
[30] Our data suggest that the tropical transitional forest
studied here may be more similar to the cerrado, where the
seasonal variation in gs (and hence gst) caused by seasonal
fluctuations in precipitation is an important limitation to Qe.
The tropical transitional forest was closely coupled to the
atmosphere, implying that variations in gst were important
in limiting seasonal rates of Qe. The dry season decline in gs
was presumably in response to a decline in surface soil
water content and in plant water potential. Measurements of
light-saturated leaf conductance and water potential of
understory trees at the study site confirm that leaf gas
exchange and plant water potential can be 2 – 3 times lower
during the dry season than during the wet season [Vourlitis
et al., 2001]. Furthermore, canopy LAI can decline by
almost 1 m2 m
2 between the wet and the dry season (G.
Vourlitis et al., submitted manuscript, 2001), which will
limit the transpirational surface area and reduce rates of
evapotranspiration. The August – September decline in Q*
from widespread biomass burning [Eck et al., 1998] may
exacerbate the dry season decline in Qe [Shuttleworth, 1988;
Bastable et al., 1993].
[31] Similarly, the increase in Qe during the wet season
presumably reflects an increase in surface and soil water
content. Average midday gs increased as rainfall increased,
and analysis of
suggests that Qe became relatively less
dependent on gst and more dependent on physical factors,
such as the evaporation of intercepted rainfall, which can
account for 10– 13% of rainfall [Ubarana, 1996]. Given
these trends, our data suggest that seasonal variation in
rainfall, and associated variations in surface water avail-
ability and VPD, caused substantial variation in annual Qe.
4.3. Uncertainties
[32] The failure of the eddy covariance power system
substantially limited the amount of data collected during the
wet season. Although estimates of Qe(PT) during these
hydrologically important periods were similar to those
measured directly from eddy covariance (Figures 4 – 6 and
Table 3) and reported from previous studies of tropical
savanna [Miranda et al., 1997; Meinzer et al., 1999] and
rain forest [Shuttleworth et al., 1984a; Shuttleworth, 1988;
Hodnett et al., 1995], the lack of direct measurements adds
uncertainty to Qe(PT).
[33] It is also unknown whether estimates of a, Qe, and gs
calculated for the study period are applicable to other years.
Our data suggest that seasonal patterns of rainfall are
important in controlling seasonal patterns of a, Qe, and gs,
so assuming that the seasonal distribution of rainfall is
similar from year to year (see Table 1), the seasonal patterns
of, and controls on, Qe are presumably applicable from year
to year. In contrast, the observed rainfall was higher than the
long-term average (Table 1), so the magnitude of a, Qe, and
gs reported here may not be applicable to drier years.
Although the measured and estimated values of Qe are
limited to the surface moisture conditions of the study period,
the close correspondence between a and rainfall suggests
that a can be estimated from total monthly rainfall data. This
result is not surprising given the fact that a is sensitive to
surface moisture availability [Priestley and Taylor, 1972]. If
a can be estimated from rainfall records, then interannual
variations in Qe estimated from this calibrated Priestley-
Taylor approach should be sensitive to interannual variations
in rainfall. Unfortunately, this approach could not be tested
during the current study because of missing precipitation data
and the short duration of the field campaign.
[34] Another limitation of the current study was the lack
of data regarding the seasonal variations in soil water
content and plant water potential. Limited measurements
of the predawn and midday (1300 LT) water potential of
understory trees were conducted in July 2000. During this
time, mean ±1 standard error predawn water potential was

0.3 ± 0.05 MPa (n = 2 plants), suggesting adequate soil
water and/or access to deep-water reserves. However, mid-
day (1300 LT) water potential was
2.7 ± 0.17 MPa (n = 2
plants), which suggests the possibility of midday plant water
stress of understory tree species early in the dry season. The
low midday plant water potential is consistent with our
interpretation of dry season limitation to gst, and hence, Qe.
[35] Acknowledgments. Funding was provided by the National Geo-
graphic Society, Committee for Research and Exploration, and the National
Science Foundation, Division of International Programs. Additional support
was provided by California State University, San Marcos (CSUSM); the
Universidade Federal de Mato Grosso (UFMT); Conselho Nacional de
Desenvolvimento Cientı́fico e Tecnológico (CNPq); Mato Grosso State
Research Support Foundation (FAPEMAT); Northern Mato Grosso Wood-
cutters Trade Union (SINDUSMAD); the Coordenação de Aperfeiçoa-
mento de Pessoal de Nı́vel Superior (CAPES); Fundaçao de Promoçao
Social do Estado de Mato Grosso (PROSOL); Telemat Brasil Telecom;
Corpo de Bombeiros do Estado de Mato Grosso; NASA-LBA; and the
Brazilian Institute for Space Research (INPE). Equipment support from the
Global Change Research Group of San Diego State University is gratefully
appreciated. The authors also thank the subject and associate editors and
two anonymous reviewers for helpful comments and suggestions.
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J. H. Campelo Jr., F. T. Caseiro, and M. M. S. Hayashi, Núcleo de
Tecnologia em Armazenagem, Universidade Federal de Mato Grosso,
78060-900 Cuiabá, Mato Grosso, Brazil.
N. Priante Filho and J. de S. Nogueira, Departamento de Fı́sica,
Universidade Federal de Mato Grosso, 78060-900 Cuiabá, Mato Grosso,
Brazil.
G. L. Vourlitis, Biological Sciences Department, California State
University, San Marcos, CA 92096, USA. (georgev@mailhost1.csusm.edu)