JOURNAL OF GEOPHYSICAL RESEARCH, VOL. 115, G04021, doi:10.

1029/2009JG001179, 2010

Atmospheric versus vegetation controls of Amazonian tropical

rain forest evapotranspiration: Are the wet and seasonally dry
rain forests any different?
Marcos H. Costa,1 Márcia C. Biajoli,1 Luciana Sanches,2 Ana C. M. Malhado,1
Lucy R. Hutyra,3 Humberto R. da Rocha,4 Renata G. Aguiar,5
and Alessandro C. de Araújo6
Received 22 October 2009; revised 31 May 2010; accepted 4 June 2010; published 11 November 2010.

[1] This study analyzes evapotranspiration data for three wet and two seasonally dry rain
forest sites in Amazonia. The main environmental (net radiation, vapor pressure deficit,
and aerodynamic conductance) and vegetation (surface conductance) controls of
evapotranspiration are also assessed. Our research supports earlier studies that demonstrate
that evapotranspiration in the dry season is higher than that in the wet season and that
surface net radiation is the main controller of evapotranspiration in wet equatorial sites.
However, our analyses also indicate that there are different factors controlling the
seasonality of evapotranspiration in wet equatorial rain forest sites and southern seasonally
dry rain forests. While the seasonality of evapotranspiration in wet equatorial forests is
driven solely by environmental factors, in seasonally dry forests, it is also biotically
controlled with the surface conductance varying between seasons by a factor of
approximately 2. The identification of these different drivers of evapotranspiration is a
major step forward in our understanding of the water dynamics of tropical forests and has
significant implications for the future development of vegetation‐atmosphere models
and land use and conservation planning in the region.
Citation: Costa, M. H., M. C. Biajoli, L. Sanches, A. C. M. Malhado, L. R. Hutyra, H. R. da Rocha, R. G. Aguiar, and A. C. de Araújo
(2010), Atmospheric versus vegetation controls of Amazonian tropical rain forest evapotranspiration: Are the wet and seasonally dry
rain forests any different?, J. Geophys. Res., 115, G04021, doi:10.1029/2009JG001179.

1. Introduction precipitation cycle: more ET in the rainy season and less in

the dryer austral winter [Nobre et al., 1991; Costa and
[2] The seasonality of evapotranspiration (ET) of the
Foley, 1997; Hahmann and Dickinson, 1997; Werth and
Amazon rain forest and its controlling mechanisms has been
Avissar, 2002].
the subject of controversy for the last two decades. In an
[3] The strong seasonal cycle of Amazonian ET generated
early study, Shuttleworth [1988] used a Penman‐Monteith
by the models is a consequence of soil moisture stress. This is
model to estimate that evapotranspiration of a forest near
clearly illustrated in two studies by Costa and Foley [1997,
Manaus in central Amazon was almost constant throughout
2000], which used the same model but generated radically
the year, with small peaks in March and September, coin-
different seasonal cycles of ET by changing the root depth
ciding with periods of increased net radiation. These find-
parameter. In the earlier paper, Costa and Foley [1997] used
ings contrast with most subsequent modeling studies that
forest representations with a 2 m deep root system and sim-
have tended to generate ET annual cycles that follow the ulated a wet season peak in ET. In contrast, in their later study
Costa and Foley [2000] used 12 m deep roots and simulated a
1
Department of Agricultural Engineering, Federal University of Viçosa, dual‐equinox peak of ET, similar to that reported by
Viçosa, Brazil. Shuttleworth [1988]. In another study, Ichii et al. [2007]
2
Department of Environmental Engineering, Federal University of evaluated the sensitivity of simulated gross primary produc-
Mato Grosso, Cuiabá, Brazil. tion (GPP) to different root depths and concluded that only
3
Department of Geography and Environment, Boston University,
Boston, Massachusetts, USA. deeply rooted systems can successfully track flux‐based GPP
4
Department of Atmospheric Sciences, University of São Paulo, São seasonality. This divergence in model results is probably
Paulo, Brazil. attributable to the operation of two different controls on
5
Department of Environmental Engineering, Federal University of seasonal evapotranspiration. While the dual‐equinox peak
Rondônia, Ji‐Paraná, Brazil.
6
Amazonas State University, Manaus, Brazil.
implies that ET is energy controlled, the wet season peak
implies ET is influenced by water‐stressed vegetation.
Copyright 2010 by the American Geophysical Union. [4] Initial reports from tower‐based eddy covariance
0148‐0227/10/2009JG001179 studies for the Amazon also indicated a wet season evapo-

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G04021 COSTA ET AL.: CONTROLS OF AMAZONIAN EVAPOTRANSPIRATION
transpiration peak. Malhi et al. [2002] measured 1 year of
water flux using an eddy covariance system near Manaus
and concluded that wet season ET is about 20% higher than
dry season ET, the correlation between ET and net radiation
being 0.86 in the dry season and 0.94 in the wet season.
Vourlitis et al. [2002] analyzed 81 days of eddy covariance
ET data for the Sinop site (a transitional forest site in the
north of Mato Grosso state, Brazil), reporting wet season ET
about 70% higher than the dry season ET. However, their
wet season data were limited to only three days.
[5] Werth and Avissar [2004], in their review of Amazon
regional ET, concluded that (at that time) there were not
enough available ground observations to enrich the discus-
sion and evaluate which was the correct seasonal cycle:
strong seasonality due to water stress with a wet season peak
or mainly radiation controlled ET with a dual‐equinox peak.
However, as Costa et al. [2004] pointed out, the eddy
covariance data that were rapidly becoming available were
indicating that ET in Amazonia displays little seasonality,
with reported peaks during the dry season actually higher
than during the rainy season. They concluded that, at least
in the Caxiuanã sites (∼350 km west of Belém, Melgaço
municipality, Pará state), ET is largely controlled by the
atmospheric conditions and that control mediated through
surface conductance is probably secondary.
[6] Rocha et al. [2004] and Souza‐Filho et al. [2005]
reported eddy covariance ET results for the Santarém
region (Tapajós National Forest, km 83, Pará state) and the
Caxiuanã site. In both cases, they found higher values of ET
during the dry season, following higher net radiation and
vapor pressure deficit. However, Aguiar et al. [2006] ana-
lyzed ET from 1999 to 2002 and in 2004 in the Jaru Bio-
logical Reserve site (Ji‐Paraná, Rondônia state), reporting a
reduction of 19.6% in the dry season. Moreover, Hutyra et al.
[2007] analyzed 4 years of eddy covariance data at the
Santarém km 67 site, finding that rates of ET were inelastic
and did not appear to depend on dry season precipitation.
[7] Interestingly, in their study of lowland mixed forest in
Borneo, where there is no clear rainfall seasonality,
Kumagai et al. [2004] observed that discrepancies between
the equilibrium and actual evaporation rates, a measure of
water stress, were caused by unpredictable intra‐annual dry
spells, which reduced transpiration.
[8] More recently, Negrón‐Juárez et al. [2007] and Hasler
and Avissar [2007] presented the first analyses of eddy
covariance data from several rain forest sites in the Amazon.
Negrón‐Juárez et al. [2007] analyzed 10 sites, concluding that
average dry season ET is largely correlated with net surface
radiation. Hasler and Avissar [2007] analyzed data from six
sites, concluding that in the wet equatorial sites (2°S–3°S), ET
increases in the dry season and decreases during the wet season
and is in phase with the net radiation cycle. For the seasonally
dry tropical sites (9°S–11°S), no clear seasonality could be
identified although net radiation and ET are still strongly
correlated (r = 0.76–0.92) in the wet season and show
decreasing correlations in the dry season (r = 0.00–0.71).
[9] In summary, most previous studies [e.g., Costa et al.,
2004; Souza‐Filho et al., 2005; Negrón‐Juárez et al., 2007;
Hasler and Avissar, 2007], with the exception of Malhi et al.,
[2002], have established the dependence of seasonal ET on net
radiation, in particular for wet equatorial sites. This interpre-
tation is supported by a recent review of 21 pan‐tropical eddy
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G04021
covariance sites that found that net radiation explained 87% of
the variance in monthly evapotranspiration across the sites
[Fisher et al., 2009]. Furthermore, the results of Vourlitis et al.
[2002], Aguiar et al. [2006], and Hasler and Avissar [2007]
suggest there is a difference in the dry season control of ET
between the wet equatorial forests and the seasonally dry
southern tropical forests. This finding implies it is important to
analyze wet equatorial sites and southern tropical sites sepa-
rately, because the dry season in Amazonia becomes increas-
ingly severe with increasing distance from the equator.
[10] In this study, we test the hypothesis that the factors
controlling ET are different in wet equatorial and seasonally
dry tropical forests. Specifically, we predict that the bio-
logical response of drought‐adapted plants in seasonally dry
forests will significantly influence ET during the dry season.
We evaluate this hypothesis through a comparison of the
abiotic and biotic drivers of ET for the wet and dry seasons
for five sites spread across the Amazon basin, including
three wet equatorial sites (around 2°S–3°S) and two sea-
sonally dry tropical sites (around 11°S).
2. Sites and Data
[11] Data used in this study were collected from five
Amazonian rain forest sites situated in different climatic
zones (equatorial and tropical) with different degrees of
seasonality and forest structure (Figure 1 and Table 1). The
three wet rain forest sites are Cuieiras Reserve (km 34,
hereafter referred to as the Manaus site), and two sites in the
Tapajós National Forest (km 67 and km 83, collectively
referred to as the Santarém sites), while the two seasonally
dry sites are located farther south in Jaru Biological Reserve
(Jaru site) and Fazenda Maracaí (Sinop site).
[12] The Manaus site is located about 60 km northwest of
Manaus city in central Amazonia and possesses a flux‐tower
managed by the Large Scale Biosphere‐Atmosphere Exper-
iment in Amazonia (LBA). The site is located in the km 34 of
the Cuieiras Biological Reserve, a protected area that belongs
to Instituto Nacional de Pesquisas da Amazônia (INPA). The
vegetation in this site is old‐growth closed‐canopy terra
firme (nonflooded) forest, about 30 m tall with emergent trees
reaching 45 m [Araújo et al., 2002]. This site is about 10 km
west of the site used by Malhi et al. [2002] for their study.
[13] There are two study sites (Santarém sites) at the Tapajós
National Forest (FLONA Tapajós), a primary forest reserve.
The reserve is on a broad, flat plateau along the Cuiabá‐San-
tarém highway. The sites located about 67 km and 83 km south
of Santarém. Vegetation is typically a tropical humid forest
with mostly evergreen and a few deciduous species. Average
canopy height is 40 m, with emergent specimens reaching 55 m
[Saleska et al., 2003; Goulden et al., 2004].
[14] The Jaru Biological Reserve site (Jaru site) is located
about 80 km north of Ji‐Paraná, Rondônia [Culf et al.,
1996]. The area contains seasonally dry tropical forest
with relatively closed canopy structure and emergent trees.
Understory vegetation of only a few meters height consists
mainly of palms [Rottenberger et al., 2004]. The mean
canopy height is 30 m, but the tallest emergent trees reach
44 m [McWilliam et al., 1996].
[15] The Sinop site is an intact transitional tropical forest
located about 53 km from Sinop, Mato Grosso state.
Although the mean canopy height is only 28 m, the tallest
G04021 COSTA ET AL.: CONTROLS OF AMAZONIAN EVAPOTRANSPIRATION G04021

Figure 1. Orientation map. Sites are represented by their specified location in Table 1.

emergent trees can reach 42 m [Priante‐Filho et al., 2004].
It should be noted that the Fazenda Maracaí (our Sinop site)
has been erroneously labeled as Fazenda Continental in
previous literature.
[16] In the wet equatorial sites (Figure 2, thin lines),
although precipitation has some seasonality, it is rarely
below the annual mean ET line (3.4 mm d−1). In the Jaru
and Sinop sites (Figure 2, bold lines), the dry season is more
intense, with four to seven months of precipitation below the
3.4 mm d−1 threshold. Temperature at the sites is typical of
tropical regions, with monthly means of all sites varying
between 21°C and 27°C. Dry season temperatures are
slightly higher than the wet season ones.
[17] In all sites, latent heat flux (LE) and sensible heat flux
(H) were measured by eddy covariance systems, while an
automatic meteorological station measured the surface net
radiation (Rn), air temperature, humidity, and wind speed
among other variables. Details about the instrumentation for
each site may be found in the original publications [Araújo
et al., 2002; Saleska et al., 2003; Goulden et al., 2004;

Table 1. Main Characteristics of the Different Sites in the Study

Instrument Canopy
Site Name Location Vegetation Description Site Coordinates Height (m) Height (m) Period of Data Used
Manaus Cueiras Reserve (km 34), Primary tropical forest 60°13′W 2°36′S 54 30 16 Jun 1999–21 Sep 2000
Amazonas (evergreen)
Santarém km 67 Tapajós National Forest Primary tropical forest 54°58′W 2°51′S 65 40 19 Jun 2002–28 Aug 2003
(km 67), Pará (evergreen)
Santarém km 83 Tapajós National Forest Primary tropical forest 54°57′W 3°03′S 65 40 29 Jun 2000–28 Jun 2001
(km 83), Pará (evergreen)
Jaru Jaru Biological Reserve, Primary forest 61°56′W 10°46′S 60 30 08 Jan 2004–31 Dec 2005
Rondônia (semideciduous)
Sinop Fazenda Maracaí, Mato Primary transitional forest 55°19′W 11°25′S 42 28 01 Jan 2000–31 Dec 2003
Grosso (semideciduous)

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G04021 COSTA ET AL.: CONTROLS OF AMAZONIAN EVAPOTRANSPIRATION G04021

that, when used to evaluate land surface model (LSM)

Figure 2. Precipitation characterization of all sites. The
dashed line indicates the average annual mean evapotranspi-
ration for all sites. Periods when precipitation is below the
dashed line approximately represent periods of water deficit.
Priante‐Filho et al., 2004; Aguiar et al., 2006; Hutyra et al.,
2007].
[18] Leaf area index (LAI) is an important descriptive
variable that provides information about the biological
nature of a site and the response of the vegetation to sea-
sonal and annual changes in climatic conditions. Directly
measured LAI was only available for the Santarém sites
[Malhado et al., 2009], where LAI varied from 5.2 m2 m−2
in January to 4.9 m2 m−2 in May. At Sinop, indirect esti-
mates of LAI through the extinction of photosynthetic
photon flux density (PPFD) by the forest canopy indicates a
maximum LAI of 5.0 m2 m−2 in February and a minimum of
2.5 m2 m−2 in July [Vourlitis et al., 2004; Sanches et al.,
2005]. Moreover, MODIS LAI data for the sites are not
particularly reliable, as more than 95% of the data are either
missing or retrieved by the less reliable secondary algo-
rithm. Cohen et al. [2006] validated the MODIS LAI col-
lection 4 product against LAI‐2000 data for nine sites in the
Western Hemisphere, including the tropical rain forest at the
km 67 site in Santarém, finding a bias of 0.05 m2 m−2
(highest among nine land cover types analyzed), RMSE of
0.83 m2 m−2 (second highest), and a correlation of only 0.54
(second lowest). Despite the lack of local observations and
the limitations of remote sensing, there are indications that
the equatorial sites are evergreen, with LAI in the range of
5–6 m2 m−2, while the tropical sites (Jaru and Sinop) are
semideciduous.
performance, measurements of energy fluxes must satisfy
the energy budget closure prior to their use in LSM eva-
luations. Other authors, however, choose not to make any
corrections to the eddy covariance data.
[20] Here we use a filtering approach to ensure that all
data used are within specified bounds of energy closure.
For a specified value of d, we use only LE data when daily
values of LE + H are within the limits Rn(1 − d) and Rn(1 +
d). Otherwise, all LE data for that specific day are discarded.
[21] Figure 3a illustrates the data availability by site that
matches the criteria Rn(1 − d) < LE + H < Rn(1 + d) on a
daily basis. Data that passes a narrower energy imbalance
limit (smaller d) undoubtedly have better quality, although
the number of the data points that pass this threshold would
be correspondingly smaller. Conversely, there would be
more data points available for analysis when data are not
filtered for energy imbalance (d = infinite), but these data
would have no quality assurance. Thus, an operational
decision needs to be made between using more data points
with unverified energy closure or fewer data points of higher
quality.
[22] To make this decision, we plot yearly evapotranspi-
ration totals at all sites against the energy closure limit d for
all sites (Figure 3b). An analysis of this figure indicates that
when d is between 0.2 and 0.4, the calculated ET shows a
consistent pattern with little apparent variation (Figure 3b).
We therefore conclude that using d in the range between 0.2
and 0.4 does not significantly bias ET estimates. In contrast,
using d = infinite (no filter applied) causes the yearly ET
values to drop in three of the four sites, because many points

3. Methods
3.1. Filtering Technique
[19] The input data required to estimate the variables
described above are air temperature and humidity, hori-
zontal wind speed, sensible and latent heat flux, atmospheric
pressure, and friction velocity. However, eddy covariance
data are known to be associated with energy closure pro-
blems [Wilson et al., 2002; Fisher et al., 2009], and different
authors have come up with different solutions to deal with
this problem. For example, Twine et al. [2000] increased Figure 3. (a) Data availability under different levels of fil-
both H and LE by the same proportion to match Rn, keeping tering (d) for all four sites and (b) annual mean evapotrans-
the Bowen ratio constant. Maayar et al. [2008] recommend piration (ET) variation under different levels of d.

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Table 2. Annual Means, Seasonal Variability, and Percentage and environmental factors, we calculate hourly and then
Increase in the Dry Season Value Compared to the Wet Season monthly averages of surface and aerodynamic conductance.
Value of the Evapotranspiration in Four Amazon Rain Forest Sites, These are analyzed together with other variables measured
According to the Energy Closure Limit d a in each site: net radiation (Rn), evapotranspiration (ET) and
ET (mm d−1)
vapor pressure deficit (VPD). Each variable was estimated
half‐hourly or hourly, depending upon data availability.
No
Site Season d = 0.1 d = 0.2 d = 0.3 d = 0.4 d = 0.5 Filter From the 48 half hour period estimates, a typical day was
derived for each month. The average of the all hourly
Manaus Year 3.73 3.58 3.58 3.52 3.50 2.93 averages of the typical day constituted the monthly mean,
Wetb 3.62 3.42 3.41 3.38 3.37 2.66
Dry c
3.84 3.74 3.75 3.67 3.64 3.20
and in this way, yearly variation could be constructed.
Increment 6% 9% 10% 9% 8% 20% [26] Following Brutsaert [1982, p.112], aerodynamic
Santarém sites Year 3.73 3.55 3.49 3.44 3.40 3.30 conductance ga (m s−1) is calculated using the following
Wet 3.78 3.45 3.40 3.35 3.32 3.20 equation:
Dry 3.67 3.65 3.59 3.52 3.48 3.40
Increment −3% 6% 5% 5% 5% 6% u2
Jaru Year 3.56 3.54 3.57 3.53 3.51 3.52 ga ¼ * : ð1Þ
Wet 3.78 3.77 3.86 3.82 3.81 3.87 u
Dry 3.35 3.32 3.27 3.24 3.22 3.18
Increment −11% −12% −15% −15% −15% −18%
This formulation, that depends on the friction velocity u*
Sinop Year 3.06 3.09 3.11 3.12 3.10 2.87 and above canopy mean horizontal wind, avoids all the
Wet 3.04 3.05 3.04 3.06 3.07 3.25 potential errors in computing the roughness length, displace-
Dry 3.08 3.13 3.18 3.18 3.12 2.49 ment height, and stability functions. Surface conductance is
Increment 1% 3% 5% 4% 2% −23% calculated using the inverted form of the Penman‐Monteith
a
Italicized values indicate estimates that lie outside the range (ET − equation:
1.2sET, ET + 1.2sET).   1
a cp VPD 1 DH
b
Wet season is between November and April.
c
Dry season is between May and October. gs ¼ ðrs Þ1 ¼  1 ; ð2Þ
LE ga LE

where LE + H underestimates Rn are being used in the
calculation of ET. In addition, using a very strict energy
closure criteria (d = 0.1) significantly reduces the number of
data points used (Figure 3a), causing variations in the cal-
culated ET. Based on this analysis, we decided to use the
central point of the d interval that does not bias the results
(d = 0.3) as the filter threshold. In other words, we only
used data when daily LE + H was within 30% of the daily
Rn, otherwise the entire eddy covariance and Rn data for
that day were discarded.
[23] We also analyzed the outliers, i.e., ET estimates that
were outside of the range (ET − 1.2sET, ET + 1.2sET),
where sET is the standard deviation of ET in a table row, and
1.2 was chosen so that 20% of the points would be outliers
(Table 2). Most of the outliers (shaded cells) reside in datasets
that were not checked for energy closure (d = infinite) or
where the energy closure filter limit is too rigorous (d = 0.1),
drastically reducing the number of data points available.
[24] We believe that by filtering the data with this energy
closure criteria, we maximize the number of data points with
acceptable quality, without significantly biasing the results.
We should emphasize that if the data are not filtered, a
procedure adopted by many authors, much lower estimates
of ET are generated, as anticipated from the typical under-
estimation of eddy covariance energy fluxes.
3.2. Computation of Biotic and Abiotic Components
[25] Evapotranspiration is influenced by four main vari-
ables: net radiation available at the surface (Rn), the vapor
pressure deficit between the evaporating surface and the
atmosphere (VPD), the conductances of the water vapor
flow known as aerodynamic conductance (ga), and surface/
stomatal conductance (gs). Rn, VPD and ga are the abiotic
environmental controls on ET, while gs is the biological
control. To characterize the evapotranspiration process and
how this process is controlled in different periods by biotic
where gs is the surface conductance (m s−1), LE is the latent
heat flux (W m−2), ra is the air density (kg m−3), cp is the
specific heat of air at constant pressure (J kg−1 °C−1), VPD
is in hPa, g is the psychometric constant (hPa °C−1), H is the
sensible heat flux (W m−2), and D is the slope of the satu-
ration vapor pressure curve (hPa °C−1).
4. Results and Discussion
[27] The monthly results for ET, Rn, VPD, ga, and gs for
the five sites are shown in Figure 4, while Table 3 sum-
marizes the variables above for the wet (Nov–Apr) and dry
(May–Oct) seasons. Data from Santarém sites (km 67 and
km 83) are highly concordant, so for the purpose of pre-
sentation, they are averaged and presented together. Inter-
estingly, the data for these sites do not overlap as much if
the eddy covariance data are not filtered. The Sinop site has
operated intermittently in the period of study. Although our
average methodology was designed to not bias the results
under heavy data missing, in some months it was not pos-
sible to define a typical day, and therefore considered the
entire month to be missing. In Table 3, sites are arranged
from the wettest dry season to the driest dry season. In
addition, Table 3 shows the results of a statistical test:
seasonal means within each column followed by the same
letter are significantly different from each other at the 0.05
significance level, according to the t test.
[28] In three out of the four sites (except Jaru), dry season
ET is higher than wet season ET. In these cases ET increases
by about 5%–10% in the dry season, while in Jaru it de-
creases by 15% (Table 3). Increases in ET are not significant
at the 5% level, while the decrease in Jaru is significant at
the 5% level, according to the t test. The results for the wet
equatorial sites support the findings of Rocha et al. [2004],
Souza‐Filho et al. [2005], Hasler and Avissar [2007], and
Rocha et al. [2009]. Moreover, the Jaru data, which depart

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G04021 COSTA ET AL.: CONTROLS OF AMAZONIAN EVAPOTRANSPIRATION
Figure 4. Seasonality of evapotranspiration: (a) main con-
trolling variables, (b) net radiation, (c) vapor pressure deficit,
(d) aerodynamic conductance, and (e) surface conductance.
from the high dry season ET pattern, are similar to those
reported by Aguiar et al. [2006] and Rocha et al. [2009].
[29] The long‐term data from the Sinop site have been
analyzed by Hasler and Avissar [2007] and Rocha et al.
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G04021
[2009]. Here, discrepant results are found: Hasler and Avissar
[2007] did not find any seasonality, whereas Rocha et al.
[2009] found ET in the dry season 29% less than in the rainy
season, while the present study found dry season ET 5% higher
than during the wet season. There are two possible explana-
tions for these differences. One possibility is the researchers
analyzed different periods of data with different climate con-
ditions. Hasler and Avissar [2007] used data from 1999 to
2003, and Rocha et al. [2009] used data from 1999 and 2001,
while the present study used data from 2000 to 2003. Such an
explanation, though possible, seems unlikely, and we believe a
more robust explanation lies in the filtering technique we
applied. If data were not filtered, we would find dry season ET
to be 23% lower than wet season ET (Table 2), a result that is
similar to Rocha et al. [2009]. However, after applying the d =
0.3 filter, ET actually increases during the dry season. Again,
the increase in the Sinop ET during the dry season is not sig-
nificant at the 5% level.
[30] Vourlitis et al. [2008] have suggested that the Sinop
site seasonality may be affected by deep water reserves,
given the lack of available water in the soil surface during
the dry season. They speculate that given the relatively
shallow depth (3.0–3.6 m) of the water table in this region,
the trees are likely to have access to this stable water source
during the dry season.
[31] At the wet equatorial sites, there is a general trend of
increasing ET toward the end of the dry season (Figure 4a,
thin lines). Although small, the net radiation in these sites
(Figure 4b, thin lines) is higher during the dry season than
during the rainy season, due to less cloudy conditions during
the dry season (Figure 4b and Table 3). By contrast, in Jaru
and Sinop (bold lines), Rn is relatively constant through the
year. Only in Santarém, the increase in Rn during the dry
season is significant at the 5% level of significance (Table 3).
[32] As predicted, vapor pressure deficit is larger in the dry
season than in the wet season in all sites (Table 3 and Figure 4c).
The seasonal amplitude is smaller in Santarém (1.0 hPa) and
larger in Sinop (5.8 hPa) possibly because the Sinop site has a
significantly drier year‐round atmosphere than the other sites.
Dry season VPD is significantly higher in Jaru and Sinop (at
the 5% level of significance), but not in Manaus and Santarém.
[33] Aerodynamic conductance, dependent on the horizontal
wind speed, does not show a significant seasonal variability,
with seasonal changes smaller than 8% (Table 3 and Figure 4d).
Changes are not significant at the 5% level in any of the sites.
[34] The three controlling variables so far discussed (Rn,
VPD and ga) correspond to the three environmental controls
of ET. VPD increases in the dry season in all cases, Rn
increases in the dry season in the wet equatorial sites, while
ga is relatively constant throughout the year. The remaining
controlling variable, surface conductance (gs), is the only
biotic control on ET, and our analysis indicates important
differences in this variable between the wet equatorial and
the seasonally dry tropical sites. While the two equatorial
wet sites have relatively minor (and not significant) drops in
gs during the dry season (Table 3), the two southern tropical
sites show much lower values with a dramatic decrease in gs
(up to 58%) in the dry season (Table 3). This decrease is due
to the biological response of the stomata to decreased water
availability and indicates that in these sites the control of
evapotranspiration in the dry season is primarily biological.
The Jaru gs drop in the dry season is significant at the 5%
G04021 COSTA ET AL.: CONTROLS OF AMAZONIAN EVAPOTRANSPIRATION G04021

Table 3. Annual Means, Seasonal Variability, and Percentage Increase in the Dry Season Value Compared to the Wet Season Value of
the Atmospheric Variables in Four Amazon Rain Forest Sites, Filtered Using d = 0.3a
Site Season P (mm d−1) T (°C) ET (mm d−1) Rn (W m−2) VPD (hPa) ga (m s−1) gs (m s−1)
Manaus Year 8.4 25.6 3.58 135.0 6.0 0.062 0.020
Wetb 10.9 25.3 3.4 129.9 4.9 0.062 0.022
Dryc 5.9 25.9 3.7 140.0 7.1 0.062 0.018
Increment 10% 8% 45% 0% −22%
Santarém sitesd Year 5.7 25.2 3.49 128.8 4.1 0.083 0.015
Wet 7.3 24.8 3.40 117.9e 3.6 0.081 0.015
Dry 4.1 25.6 3.59 139.6e 4.6 0.084 0.014
Increment 5% 18% 27% 3% −6%
Jaru Year 4.5 22.9 3.57 135.8 6.8 0.046 0.013
Wet 7.5 20.9 3.86e 136.1 4.1e 0.044 0.017e
Dry 1.4 24.9 3.27e 135.4 9.4e 0.047 0.008e
Increment −15% −1% 129% 6% −53%
Sinopf Year 5.0 25.5 3.11 130.2 10.1 0.069 0.010
Wet 9.5 25.5 3.04 130.7 7.2e 0.066 0.013
Dry 0.6 25.5 3.18 129.6 13.0e 0.071 0.007
Increment 5% −1% 80% 8% −46%
a
P, precipitation; T, temperature; ET, evapotranspiration; Rn, net radiation; VPD, vapor pressure deficit; ga, aerodynamic conductance; gs, surface
conductance.
b
Wet season is between November and April.
c
Dry season is between May and October.
d
Data for Santarém are the average of km 67 and km 83 sites.
e
Seasonal means that are significantly different from each other at the 0.05 significance level, according to the t test.
f
At Sinop, ga and gs were not tested because of the reduced number of observations (2 in the wet season, 4 in the dry season); for reference, standard
deviation of ga and gs are 0.0023 and 0.0064 m s−1 in the wet season, and 0.0110 and 0.0020 m s−1 in the dry season.

level. Unfortunately, there are not enough data at Sinop for
a formal statistical analysis (only 2 points of data in the wet
season), so the significance for gs and ga cannot be calcu-
lated at this site. Despite this limitation, there is a strong
reduction in gs in both seasonally dry forests sites.
[35] The seasonal change in ET is a combination of the four
drivers, Rn, VPD, ga, and gs. In the seasonally dry forests they
behave in three different ways: (1) Rn and ga show little
variation (<10%) from season to season in both sites, causing
weak patterns of seasonal change; (2) gs decreases by about
half, contributing to a decrease in ET; and (3) VPD increases
in both sites by over 80%, contributing to an increase in ET.
The increase in VPD partially balances the decrease in gs,
avoiding a strong reduction in ET.
[36] A combined analysis of the four controlling variables
(Rn, VPD, ga and gs) across the four sites indicates a clear
distinction between the wet equatorial and the southern sea-
sonally dry tropical sites with respect to the control of ET in
Amazonia. Two important research findings support this
statement: first, the strong dependence of ET on Rn for the
wet equatorial sites, a characteristic not seen in the seasonally
dry tropical sites. Second, the observation that while wet
equatorial sites do not seem to be water‐stressed, the southern
tropical sites exhibit different degrees of water stress pro-
portional to the intensity of the dry season. In other words, ET
in the wet equatorial sites is almost totally environmentally
controlled, while in the seasonally dry tropical sites the bio-
logical response of the plants to water stress (e.g., a stomatally
mediated reduction in surface conductance) plays an impor-
tant controlling role.
5. Conclusions
[37] The question originally posed by this study was the
following: is evapotranspiration in wet equatorial and sea-
sonally dry tropical rain forests controlled by different fac-
tors? The answer is yes, in the sense that in wet equatorial
sites evapotranspiration in the dry season is higher than that in
the wet season, and surface net radiation is the main controller
of evapotranspiration in these sites. Our analyses also indicate
that wet equatorial rain forest and the southern seasonally dry
tropical rain forest in the Amazon are characterized by dif-
ferent drivers of the seasonality of evapotranspiration. Our
key finding is that while evapotranspiration in wet equatorial
forests is driven by abiotic environmental factors, in southern
seasonally dry tropical forests, seasonal surface conductance
(the biotic response of plants to water stress and therefore the
biological driver of evapotranspiration) varies by a factor of
2. Identification of these different drivers of evapotranspira-
tion is a major step forward in our understanding of the water
dynamics of tropical forests and has significant implications
for the future development of vegetation‐atmosphere models
and land use and conservation planning in the region.
[38] One potential application of these results is to
improve the structure and calibration of climate models of
the Amazon rain forest. The realization that wet equa-
torial and tropical rain forests behave differently requires
a regionally specific evaluation of model results. Further-
more, although the rain forest clearly has mechanisms to
access water deeply stored in the soils, through a deep root
system [Nepstad et al., 1994; Hodnett et al., 1996; Negrón‐
Juárez et al., 2007], hydraulic redistribution [Oliveira et al.,
2005], or shallow water table [Vourlitis et al., 2008], it is not
correct to assume in models that rain forests are not water
stressed, as the southern seasonally dry tropical rain forests
do show a relevant degree of water stress. If the wet equa-
torial forests have similar rooting systems to their tropical
counterparts, they would also be water stressed if the dry
season were as intense near the equator as it is at 10°S. This
sensitivity of wet equatorial rain forests to water stress has

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G04021 COSTA ET AL.: CONTROLS OF AMAZONIAN EVAPOTRANSPIRATION
already been demonstrated by rainfall exclusion experiments
[Nepstad et al., 2002; Meir et al., 2009].
[39] Our research also raises an important question that
will need to be answered by future studies: are regional
differences in the biological control of evapotranspiration a
consequence of dry season duration and intensity only, or
does composition and structure of the forest influence
observed patterns of seasonality? In other words, are wet
equatorial and seasonally dry tropical rain forests different
in structural attributes such as root systems, so that the trees
of seasonally dry forests are better adapted to water stress
than those in wet equatorial forests? If this indeed turns out
to be the case, it may have significant implications for the
future trajectory of Amazonian vegetation under anthropo-
genic climate change and, by extension, regional conser-
vation prioritization and policy.
[40] Acknowledgments. This research was supported by NASA
under grant NCC5‐687. Thanks to Richard Ladle, senior research fellow
at the Oxford University Centre for the Environment, for proofreading
the manuscript.
References
Aguiar, R. G., C. von Randow, N. Priante‐Filho, A. O. Manzi, L. J. G.
Aguiar, and F. L. Cardoso (2006), Fluxos de massa e energia em uma
floresta tropical no sudoeste da Amazônia, Rev. Bras. Meteorol., 21(3b),
248–257.
Araújo, A. C., et al. (2002), Comparative measurements of carbon dioxide
fluxes from two nearby towers in a central Amazonian rain forest: The
Manaus LBA site, J. Geophys. Res., 107(D20), 8090, doi:10.1029/
2001JD000676.
Brutsaert, W. (1982), Evaporation into the Atmosphere, 1st ed., 299 pp.,
Kluwer, Boston.
Cohen, W. B., T. K. Maiersperger, D. P. Turner, W. D. Ritts, D. Pflugmacher,
R. E. Kennedy, A. Kirschbaum, S. W. Running, M. Costa, and S. T. Gower
(2006), MODIS land cover and LAI collection 4 product quality across nine
sites in the Western Hemisphere, IEEE Trans. Geosci. Remote Sens., 44(7),
1843–1857.
Costa, M. H., and J. A. Foley (1997), Water balance of the Amazon Basin:
Dependence on vegetation cover and canopy conductance, J. Geophys.
Res., 102(D20), 23,973–23,989.
Costa, M. H., and J. A. Foley (2000), Combined effects of deforestation
and doubled atmospheric CO2 concentrations on the climate of Amazonia,
J. Clim., 13, 18–34.
Costa, M. H., J. D. C. Souza‐Filho, and A. Ribeiro (2004), Comments on
“The regional evapotranspiration of the Amazon,” J. Hydrometeorol., 5,
1279–1280.
Culf, A. D., J. L. Esteves, A. de O. Marques‐Filho, and H. R. da Rocha
(1996), Radiation, temperature and humidity over forest and pasture, in
Amazonian Deforestation and Climate, edited by J. H. C. Gash, C. A.
Nobre, J. M. Roberts, and R. L. Victoria, pp. 57–77, John Wiley, U.K.
da Rocha, H. R., M. L. Goulden, S. D. Miller, M. C. Menton, L. D. V. O.
Pinto, H. C. Freitas, and A. M. S. Figueira (2004), Seasonality of water
and heat fluxes over a tropical forest in eastern Amazonia, Ecol. Appl., 14(4),
S22–S32.
da Rocha, H. R., et al. (2009), Patterns of water and heat flux across a
biome gradient from tropical forest to savanna in Brazil, J. Geophys.
Res., 114, G00B12, doi:10.1029/2007JG000640.
Fisher, J. B., et al. (2009), The land‐atmosphere water flux in the tropics,
Global Change Biol., 15, 2694–2714.
Goulden, M. L., S. D. Miller, H. R. da Rocha, M. C. Menton, H. C. Freitas,
A. M. S. Figueira, and C. A. D. Sousa (2004), Diel and seasonal patterns
of tropical forest CO2 exchange, Ecol. Appl., 14(4), S42–S54.
Hahmann, A. N., and R. E. Dickinson (1997), RCCM2‐BATS model over
tropical South America: Applications to tropical deforestation, J. Clim.,
10, 1944–1964.
Hasler, N., and R. Avissar (2007), What controls evapotranspiration in the
Amazon Basin?, J. Hydrometeorol., 8, 380–395.
Hodnett, M. G., J. Tomasella, A. de O. Marques Filho, and M. D. Oyama
(1996), Deep soil water uptake by forest and pasture in central Amazo-
nia: Predictions from long‐term daily rainfall data using a simple water
8 of 9
G04021
balance model, in Amazonian Deforestation and Climate, edited by
J. H. C. Gash, C. A. Nobre, J. M. Roberts, and R. L. Victoria,
pp. 79–99, John Wiley, U.K.
Hutyra, L., J. W. Munger, S. R. Saleska, E. Gottlieb, B. C. Daube, A. L.
Dunn, D. F. Amaral, P. B. de Camargo, and S. C. Wofsy (2007), Sea-
sonal controls on the exchange of carbon and water in an Amazonian rain
forest, J. Geophys. Res., 112, G03008, doi:10.1029/2006JG000365.
Ichii, K., H. Hashimoto, M. A. White, C. Potter, L. R. Hutyra, A. R. Huete,
R. B. Myneni, and R. R. Nemani (2007), Constraining rooting depths in
tropical rain forests using satellite data and ecosystem modeling for accu-
rate simulation of gross primary production seasonality, Global Change
Biol., 13, 67–77.
Kumagai, T., T. M. Saitoh, Y. Sato, H. Takahashi, O. J. Manfroi, T. Morooka,
K. Kuraji, M. Suzuki, T. Yasunari, and H. Komatsu (2004), Annual water
balance and seasonality of evapotranspiration in a Bornean tropical rain
forest, Agric. For. Meteorol., 128, 81–92.
Maayar, M. E., J. M. Chena, and D. T. Price (2008), On the use of field
measurements of energy fluxes to evaluate land surface models, Ecol.
Modell., 214, 293–304.
Malhado, A. C. M., M. H. Costa, F. Z. Lima, K. C. Portilho, and D. N.
Figueiredo (2009), Seasonal leaf dynamics in an Amazonian tropical
forest, For. Ecol. Manage., 258, 1161–1165, doi:10.1016/j.foreco.2009.
06.002.
Malhi, Y., E. Pegoraro, A. D. Nobre, M. G. P. Pereira, J. Grace, A. D. Culf,
and R. Clement (2002), Energy and water dynamics of a central Amazo-
nian rain forest, J. Geophys. Res., 107(D20), 8061, doi:10.1029/
2001JD000623.
McWilliam, A. L. C., O. M. R. Cabral, B. M. Gomes, J. L. Esteves, and J. M.
Roberts (1996), Forest and pasture leaf‐gas exchange in south‐west Ama-
zonia, in Amazonian Deforestation and Climate, edited by J. H. C. Gash,
C. A. Nobre, J. M. Roberts, and R. L. Victoria, pp. 265–285, John Wiley,
U.K.
Meir, P., et al. (2009), The effects of drought on Amazonian rain forests, in
Amazonia and Global Change, Geophys. Monogr. Ser., vol. 186, edited
by M. Keller, M. Bustamante, J. Gash, and P. S. Dias, pp. 429–449,
AGU, Washington, D.C.
Negrón‐Juárez, R. I., M. G. Hodnett, R. Fu, M. L. Goulden, and C. von
Randow (2007), Control of dry season evapotranspiration over the Ama-
zonian forest as inferred from observations at a southern Amazon forest
site, J. Clim., 20, 2827–2839, doi:10.1175/JCLI4184.1.
Nepstad, D. C., C. R. de Carvalho, E. A. Davidson, P. H. Jipp, P. A. Lefebvre,
G. H. Negreiros, E. D. da Silva, T. A. Stone, S. E. Trumbore, and S. Vieira
(1994), The role of deep roots in the hydrological and carbon cycles of
Amazonian forests and pastures, Nature, 372(6507), 666–669.
Nepstad, D. C., et al. (2002), The effects of partial throughfall exclusion
on canopy processes, aboveground production, and biogeochemistry of
an Amazon forest, J. Geophys. Res., 107(D20), 8085, doi:10.1029/
2001JD000360.
Nobre, C. A., P. J. Sellers, and J. Shukla (1991), Amazonian deforestation
and regional climate change, J. Clim., 4, 957–988.
Oliveira, R. S., T. E. Dawson, S. S. O. Burgess, and D. C. Nepstad (2005),
Hydraulic redistribution in three Amazonian trees, Oecologia, 145(3),
354–363.
Priante‐Filho, N., L. Sanches, A. C. Dalmolin, and J. S. de Nogueira
(2004), Comparison of the mass and energy exchange of a pasture and
a mature transitional tropical forest of the southern Amazon Basin during
a seasonal transition, Global Change Biol., 10, 863–876, doi: 10.1111/
j.1529-8817.2003.00775.x.
Rottenberger, S., U. Kuhn, A. Wolf, G. Schebeske, S. T. Oliva, T. M.
Tavares, and J. Kesselmeier (2004), Exchange of short‐chain aldehydes
between Amazonian vegetation and the atmosphere at a remote forest
site in Brazil, Ecol. Appl., 14(4), S247–S262.
Saleska, S. R., et al. (2003), Carbon in Amazon forests: Unexpected sea-
sonal fluxes and disturbance‐induced losses, Science, 302(5650),
1554–1557.
Sanches, L., G. S. Suli, N. Priante Filho, G. L. Vourlitis, and J. de S. Nogueira
(2005), Índice de área foliar em floresta de transição Amazônia Cerrado,
Ciência Natura, 1, 37–46.
Shuttleworth, W. J. (1988), Evaporation from Amazonian rain forest, Philos.
Trans. R. Soc. London Ser. B, 233(1272), 321–346.
Souza‐Filho, J. D. C., A. Ribeiro, M. H. Costa, and J. C. P. Cohen (2005),
Control mechanisms of the seasonal variation of transpiration in a north-
east Amazonian tropical rain forest (in Portuguese with English abstract),
Acta Amazonica, 35(2), 223–229.
Twine, T. E., W. P. Kustas, J. M. Norman, D. R. Cook, P. R. Houser, T. P.
Meyers, J. H. Prueger, P. J. Starks, and M. L. Wesely (2000), Correcting
eddy‐covariance flux underestimates over a grassland, Agric. For.
Meteorol., 103, 279–300.
G04021 COSTA ET AL.: CONTROLS OF AMAZONIAN EVAPOTRANSPIRATION
Vourlitis, G. L., N. Priante‐Filho, M. M. S. Hayashi, J. de S. Nogueira, F. T.
Caseiro, and J. H. Campelo Jr. (2002), Seasonal variations in the evapo-
transpiration of a transitional tropical forest of Mato Grosso, Brazil, Water
Resour. Res., 38(6), 1094, doi:10.1029/2000WR000122.
Vourlitis, G. L., N. Priante Filho, M. M. S. Hayashi, J. de S. Nogueira,
F. Raiter, W. Hoegel, and J. H. Campelo Jr. (2004), Effects of meteoro-
logical variations on the CO2 exchange of a Brazilian transitional tropical
forest, Ecol. Appl., 14, 89–100, doi:10.1890/01-6005.
Vourlitis, G. L., J. de S. Nogueira, F. de A. Lobo, K. M. Sendall, S. R.
de Paulo, C. A. A. Dias, O. B. Pinto Jr., and N. L. R. de Andrade (2008),
Energy balance and canopy conductance of a tropical semideciduous forest
of the southern Amazon Basin, Water Resour. Res., 44, W03412,
doi:10.1029/2006WR005526.
Werth, D., and R. Avissar (2002), The local and global effects of Amazon
deforestation, J. Geophys. Res., 107(D20), 8087, doi:10.1029/
2001JD000717.
Werth, D., and R. Avissar (2004), The regional evapotranspiration of the
Amazon, J. Hydrometeorol, 5, 100–109.
9 of 9
G04021
Wilson, K., et al. (2002), Energy balance closure at FLUXNET sites, Agric.
For. Meteorol., 113, 223–243.
R. G. Aguiar, Department of Environmental Engineering, Federal
University of Rondônia, Rua Rio Amazonas 351, Jardim dos Migrantes,
Ji‐Paraná, RO, 78961‐970, Brazil.
M. C. Biajoli, M. H. Costa, and A. C. M. Malhado, Department of
Agricultural Engineering, Federal University of Viçosa, Av. P. H. Rolfs
s/n, Viçosa, MG, 36570‐000, Brazil.
A. C. de Araújo, Amazonas State University, Av. Djalma Batista 3578,
Manaus, AM, 69050‐010, Brazil.
H. R. da Rocha, Department of Atmospheric Sciences, University of São
Paulo, Rua do Matão 1226, Cidade Universitária, São Paulo, SP, 05508‐
090, Brazil.
L. R. Hutyra, Department of Geography & Environment, Boston
University, 675 Commonwealth Ave., Boston, MA, 02215, USA.
L. Sanches, Department of Environmental Engineering, Federal
University of Mato Grosso, Av., Fernando Correa da Costa s/n, Cuiabá,
MT, 78000‐000, Brazil.