ECOHYDROLOGY

Ecohydrol. 9, 610–630 (2016)

Published online 7 September 2015 in Wiley Online Library
(wileyonlinelibrary.com) DOI: 10.1002/eco.1661

Improving the ability of 3-PG to model the water balance of

forest plantations in contrasting environments
Auro C. Almeida1* and Peter J. Sands2
1
CSIRO, Land and Water, Private Box 12, Hobart, Tasmania, Australia, 7001
2
39 Oakleigh Av, Taroona, Tasmania, Australia, 7053

ABSTRACT
Because water availability is a primary factor that influences growth of forest plantations, quantifying plantation water use is an
important part of the decision-making process when selecting new plantation areas. The 3-PG model (Landsberg and Waring,
Forest Ecology and Management 95: 209–228, 1997) has been used successfully to model forest growth for several species.
However, some studies have demonstrated limitations in 3-PG’s ability to accurately model the soil and plant water balance. This
paper addresses these limitations through the development and validation of a modified water balance submodel that models
canopy interception, soil evaporation, tree transpiration, drainage, run-off, soil water content and available soil water in the root
and non-root zone. The new submodel was tested and validated using detailed, long-term observed growth and water balance
data from contrasting environments in Australia and Brazil and showed improved estimation of available soil water, even when
only monthly weather data were available. However, the new submodel requires more soil texture-dependent parameters than the
original submodel. Sensitivity analysis of these new parameters shows that values of soil field capacity, wilting point and soil
depth may have strong impacts on the prediction of run-off, soil water content and available soil water. Copyright © 2015 John
Wiley & Sons, Ltd.

KEY WORDS 3-PG model; water balance; plantation; forest growth; water use; available soil water

Received 24 May 2015; Revised 19 June 2015; Accepted 21 June 2015

INTRODUCTION 3-PG includes a simple soil water balance submodel

comprising a single soil layer, with evapotranspiration (ET)
Globally, the area of forest plantations is increasing, mainly
driven by stand-level canopy leaf area index (LAI) and
in drier regions where there is competition for water
canopy conductance. This water balance submodel has
resources (Winjum and Schroeder, 1997; Almeida et al.,
been criticized on a number of counts. Firstly, water stress
2010). In order to optimally manage these plantations,
is based on the ASW of the entire soil profile, whereas it is
models that accurately predict their growth and water use
only the water content of the root zone that directly affects
by coupling stand growth and forest soil water balance are
plant water status (Whitehead and Beadle, 2004). Second-
required. Examples are Forest BGC (Running and
ly, tree transpiration and soil evaporation are combined as a
Coughlan, 1988), CABALA (Battaglia et al., 2004) and
single flux of water, whereas in reality, they are separate
3-PG (Landsberg and Waring, 1997; Sands and Landsberg,
fluxes that depend on environmental and stand factors in
2002; Landsberg and Sands, 2010).
different ways. Thirdly, the dynamics of soil water balance
3-PG is a process-based forest growth model widely used
can occur on a much shorter timescale than a month, so the
as a research and management tool (Almeida et al., 2004a;
monthly time step fails to account for these changes (e.g.
Landsberg and Sands, 2010, Section 9.4) and has been
Paul et al., 2006; Almeida et al., 2007).
extensively applied to plantations or even-aged, relatively
In this paper, we present a more detailed water balance
homogeneous forests. It operates on a monthly time step and
submodel that improves the ability of 3-PG to predict the
predicts stem, foliage and root biomass, number of trees per
stand-level forest soil water balance, while maintaining the
hectare, and available soil water (ASW) in response to
simplicity of 3-PG and its ability to accurately predict forest
changing weather conditions and management interventions
growth. To address the aforementioned criticisms and
such as irrigation, thinning and fertilization.
deficiencies, the new submodel uses water availability in
the root zone to determine the effects of soil water stress,
*Correspondence to: Auro C. Almeida, CSIRO, Land and Water, Private
Box 12, Hobart, Tasmania, Australia, 7001. considers transpiration and soil water evaporation as distinct
E-mail: Auro.Almeida@csiro.au processes that depend on stand characteristics and environ-

Copyright © 2015 John Wiley & Sons, Ltd.

 MODEL WATER BALANCE OF FOREST PLANTATIONS IN CONTRASTING ENVIRONMENTS
mental factors in different ways and allows the use of shorter
time steps than a month, including a daily time step.
However, this approach requires more soil texture-
dependent parameters than does the original submodel.
Also, rigorous validation of the model requires long-term
observed data on the separate fluxes in the water balance,
which may not always be available. We used data from
plantations grown in contrasting environments that together
provide observed data for all the fluxes in the water
balance. To test the model for different species and
contrasting soils, climates and water availability, we used
detailed forest growth and water balance measurements
from Eucalyptus grandis hybrids grown on the east coast of
Brazil (Soares and Almeida, 2001; Almeida et al., 2004b,
2007) and from Eucalyptus globulus plantations in Victoria
and South Australia (Benyon et al., 2006) and in Western
Australia (WA) (Mendham et al., 2011).
We test and compare the performance of the original
and new water balance submodels using three different
time steps: (i) monthly; (ii) ‘pseudo-daily’, where the time
step is the average time between precipitation events; and
(iii) daily. Sensitivity analyses were conducted to quantify
the sensitivity of water balance and predicted tree growth
to the new soil-specific parameters.
MODEL DESCRIPTION
Overview of 3-PG
3-PG is driven by monthly climatic data and requires
knowledge of the local soil texture, soil water holding
capacity and soil fertility. Simple relationships describe the
processes underpinning tree growth, mortality and water use
on a monthly timescale. Many of the parameters in these
relationships are species specific. Values are also required at
some initial age for stand stocking, the stem (including
bark and branches), foliage and root biomass per hectare,
and the ASW.
The parameterization of 3-PG has been described and
illustrated for a range of individual species or genotypes within
a species. 3-PG is used worldwide for different tree species
such as E. globulus (Sands and Landsberg, 2002; Fontes et al.,
2006; Miehle et al., 2009; Smettem et al., 2013), Eucalyptus
nitens (Rodriguez et al., 2009; Pérez-Cruzado et al., 2011;
Gonzalez-Garcia et al., 2015), E. grandis (Esprey et al., 2004;
Stape et al., 2004; Almeida et al., 2004b, 2007), Eucalyptus
cladocalyx (Paul et al., 2007), Pinus patula (Dye, 2001), Pinus
radiata (Coops, 1999; Rodríguez et al., 2002; Feikema et al.,
2010) and more recently to mixed species (Forrester and Tang,
2015). The model has been applied at stand and spatial scales
(Coops and Waring, 2001; Tickle et al., 2001; Sands, 2004;
Almeida et al., 2010) for different purposes, e.g. to estimate
potential productivity or the impact of climate and silvicultural
managements on forest yield and as a research tool for forest
Copyright © 2015 John Wiley & Sons, Ltd.
611
management decision-making (Landsberg, 2003; Almeida
et al., 2004a).
A full description of 3-PG is given by Landsberg and
Sands (2010), which we abbreviate in this paper to L&S.
Here, we focus on the water balance submodel. The
original water balance submodel in 3-PG, called WB1 in
this paper, assumes the soil profile is homogeneous and of
a fixed depth and that roots always have access to all water
stored in the profile. The plant ASW in the full profile is the
focus of the model and mediates the effects of water stress
on growth. Water availability in the profile is specified by
the maximum ASW capacity and two parameters cθ and nθ
that characterize the soil water-dependent growth modifier
fθ. WB1 does not predict actual soil water content (SWC).
The monthly change in ASW is the monthly rainfall less the
sum of a rainfall interception loss, surface run-off and stand
transpiration ET (which combines canopy transpiration and
soil evaporation). A fraction of rainfall is intercepted by the
foliage and subsequently returned to the atmosphere (through
evaporation). The remaining rainfall is added to the ASW
pool, and any excess over the maximum ASW capacity of the
profile is deemed lost as surface run-off and deep drainage.
Stand transpiration is determined using an expression based on
the Penman–Monteith equation. This requires a stand-level
canopy conductance gC, which increases linearly with
increasing canopy LAI up to some maximum value. When
LAI is zero or small, gC is non-zero because ET includes soil
evaporation. Conductance is also modified by functions of
temperature, vapour pressure deficit (VPD), ASW and
atmospheric CO2 (L&S, Section 9.2.7; Almeida et al., 2009).
The new water balance submodel
The three deficiencies of WB1 we address here are as follows:
(i) not all the water in the soil profile is available for
transpiration until roots have fully occupied the profile; (ii)
soil evaporation and canopy transpiration depend on stand
characteristics and environmental factors in different ways
and should not be aggregated; and (iii) the monthly time step
ignores the fact that rainfall occurs in discrete events, whose
temporal distribution affects the soil water balance. The new
water balance model, called WB2 in this paper, addresses
these issues by (i) subdividing the profile into an upper
horizontal zone containing roots and a lower root-free zone,
with the effects of water stress based only on the water content
of the root zone, (ii) modelling canopy transpiration and soil
evaporation as distinct processes and (iii) explicitly consid-
ering time steps that are less than a month. WB2 predicts the
SWC of the profile, and ASW is determined from this, the
current root depth and texture-dependent soil characteristics.
The principles of WB2 are described here; mathematical
details are in Appendix A.
The hydrological balance in WB2 is as described in L&S
(Section 7.1), except that recharge from a water table and
Ecohydrol. 9, 610–630 (2016)
612 A. C. ALMEIDA AND P. J. SANDS
lateral flow are not considered [Equation (A2)]. Input of water
is assumed to be from rainfall, and possibly irrigation.
Contributions from snow are not explicitly considered.
However, such contributions could be included by using a
simple model of snow melt to allocate additional daily
amounts of equivalent rain.
Rainfall is partially lost through canopy interception and
subsequent evaporation, and the remaining throughfall is
added to the root zone. It is assumed that all input water comes
from rainfall, possibly including irrigation. Snow contribu-
tions have not been modelled because our experimental sites
are not subjected to snow. The fraction of rainfall lost by
interception is determined by canopy LAI up to a maximum
fraction, which is a parameter of the species [Equation (A5)].
Once the profile is saturated, any further throughfall is lost as
run-off. Free drainage occurs from the root zone into the root-
free zone, and from the root-free zone out of the bottom of the
profile. Soil evaporation and transpiration remove water from
the root zone, and recharge of the root zone can occur from the
root-free zone through hydraulic pressure.
Root and root-free zones. The soil profile in WB2 is still
homogeneous but is divided into a root zone, from which
water is lost by transpiration and soil evaporation, and a
non-root zone from which water cannot be immediately
accessed by trees. The soil water-dependent growth
modifier (fθ) is defined as in WB1 but is now based on
the water content of the root zone only.
The ASW, i.e. the amount of water above wilting point, is
calculated only for the root zone and is used to determine the
effects of water stress on growth and canopy conductance.
This is performed using a growth modifier fθ based on
relative ASW θr in the root zone [Equation (A4)]. This
modifier varies from zero when θr = 0 to 1 when θr = 1 and is
applied to both growth and canopy conductance.
The depth zR (mm) of the root zone is assumed to be
proportional to current root biomass, up to some site-
specific maximum depth zRx (Appendix A.7). Below this
limit, each kilogram of root is assumed to exploit a volume
σ zR (m3 kg
1) of soil, where σ zR is called the specific root
volume. As root biomass increases, the volume of the root
zone increases at the expense of the root-free volume, and
the water contents of these two volumes are adjusted to
take this change into account. Provision is made for the
possibility that SWC is observed only down to some depth
zO, less than the profile depth zS, but the full profile is
always used when determining the water balance.
Drainage from the root zone into the root-free zone and
from the latter out of the bottom of the profile is
considered, and the rate of free drainage is proportional
to the amount of water above field capacity (Appendix
A.6). Recharge of the root zone is based on a simple
representation of the movement of water across a boundary
between two regions of soil with different volumetric
Copyright © 2015 John Wiley & Sons, Ltd.
SWCs. The recharge flux is assumed to be proportional to
the difference between the volumetric SWCs (θ) of the two
zones (Appendix A.7), and water flows from the region
with the higher θ to that with the lower θ.
WB2 requires several texture-dependent soil character-
istics. These include the volumetric SWCs of the soil at
wilting point, field capacity (above which free drainage
occurs) and saturation (θwp, θfc and θsat; m3 m
3).
Additional texture-dependent characteristics are the param-
eters cθ and nθ that characterize fθ, the rate constant (kDrain,
day
1) for drainage of water through the soil and a measure
(kS, m day
1) of soil hydraulic conductivity determining
the rate of root-zone recharge. These can be assigned by
direct measurement, or on the basis of published soils data
(Williams et al., 1983; Saxton and Rawls, 2006). The
fraction pstones of soil volume occupied by stones is also
taken in account.
Transpiration and soil evaporation. Canopy transpiration
and soil evaporation are modelled separately in WB2
(Appendix A.4). Transpiration is modelled as in WB1,
using the Penman–Monteith equation, but because it now
excludes soil evaporation, gC is the conductance of the
canopy alone and increases from zero as LAI increases
from zero, and transpiration is driven by the radiation
absorbed by the canopy, not incident upon it.
Soil evaporation is implicitly confined to a shallow
upper soil layer that acts as a barrier to further evaporation
as the soil dries. Soil evaporation rate (eS) is high when the
soil surface is wet after rain and declines as it dries out.
This is modelled (Appendix A.5) using the so-called two-
phase Ritchie (1972) model. In Phase 1 (after a wetting
event), eS is the wet-surface rate and determined using a
Penman–Monteith equation. Phase 2 commences once
accumulated evaporation exceeds an amount ES1, and eS is
then assumed to decline hyperbolically with increasing
evaporation accumulated above that in Phase 1. The rate of
decline is described by a parameter (ES2), which is the
amount of evaporation in Phase 2 required to reduce eS by
50%. The equations are Equations (A9) and (A10). ES1 and
ES2 are potentially soil texture dependent. WB2 takes into
account the difference between evaporation in full sun in a
partially open canopy and in the shade under the canopy.
Pseudo-daily and daily time steps. Two approaches can be
used to improve the time resolution of the water balance
submodel. The first is to use daily weather data and to run
both the growth and water balance submodels of 3-PG with a
daily time step. The second approach uses traditional
monthly 3-PG input data provided the average number nR
of days on which rain falls in each month is available. If R is
the total rain for the month, the month is divided into nR
periods of equal length, each with an amount R/nR of rain.
This rain is assumed to fall on the first day of each period,
Ecohydrol. 9, 610–630 (2016)
 MODEL WATER BALANCE OF FOREST PLANTATIONS IN CONTRASTING ENVIRONMENTS 613

with the remaining days dry. These nR periods are then used
as time steps with 3-PG applied to each period. We call this
approach ‘pseudo-daily’, and it can be used equally with the
original or new soil water balance submodels. Simple
expressions can be written for each water balance compo-
nent over each pseudo-daily time period (Appendix A).
METHODS
Sites and species
Contrasting climatic conditions, soils and species are desirable
to test and validate the performance of the model. We used data
from plantations of E. grandis hybrids grown at Aracruz on the
east coast of Brazil and from E. globulus grown at contrasting
sites in Southern Australia. The E. globulus data come from
eight sites in the Green Triangle area of Victoria and South
Australia and from two sites in WA. All sites had been
extensively monitored for growth and water use over a long
period. Table I lists the experimental sites, their location and
main characteristics. Figure 1 shows the average monthly
weather data of selected study sites in WA, Green Triangle and
Brazil and clearly demonstrates the strong variability between
sites mainly in terms of rainfall distribution and air temperature.
Further details are given in the following.
Procedures for the calibration and validation of the
original 3-PG model for E. grandis and E. globulus have
been published elsewhere (Sands and Landsberg, 2002;
Esprey et al., 2004; Almeida et al., 2004b).
Aracruz site
The Aracruz site is in an experimental catchment that is well
documented and described (Soares and Almeida, 2001;
Almeida et al., 2004b, 2007). Plantation growth and water
balance were intensively monitored for a period of more than
7 years, and rainfall, soil moisture, groundwater level and run-
off were monitored automatically and continuously. Tree
transpiration (Mielke et al., 1999), canopy rainfall intercep-
tion and destructive biomass samples (stem, bark, branch,
foliage and roots) were obtained from intensive seasonal or
annual campaigns. Tree growth was measured monthly in
permanent sample plots (Almeida et al., 2004a).

Table I. Details of experimental sites.

Mean annual Age of Planted Water

precipitation measurement Planted stocking table depth
Site name Species Lat. Long. (mm year
1) (years) date (tree ha
1) Soil texture (m) Sourcea

Aracruz E. grandis
19·861
40·211 1250 1–7 2/1997 1111 Clay loam 15.0 1, 2
hybrid
Bessie E. globulus
38·158 141·818 786 6–9 7/1998 1100 Loamy sand/clay loam 6·0 3
DPI E. globulus
37·847 142·076 679 8–11 7/1996 950 Clay loam/clay 10·0 3
Gummy E. globulus
37·737 140·782 710 4–9 7/1996 1125 Loamy sand/clay 4·3 3
Jack E. globulus
37·373 140·456 664 3–7 7/1998 1175 Loamy sand/clay 3·3 3
Jasper E. globulus
37·836 141·620 706 7–9 7/1998 750 Sandy loam/clay 20·0 3
Jill E. globulus
37·373 140·456 643 4–7 7/1998 1200 Loamy sand 10·5 3
Padthaway E. globulus
36·541 140·652 503 4–6 7/1995 1025 Loamy sand 15·9 3
Vikki E. globulus
38·104 142·146 736 4–9 8/1998 925 Clay loam/clay 8·3 3
Wellstead E. globulus
34·580 118·550 591 2–8 7/1996 1131 Sand n/a 4, 5
Scott River E. globulus
34·260 115·520 984 2–8 7/1996 1145 Sandy clay loam n/a 4, 5
a
Sources: 1, Almeida et al. (2004a); 2, Soares and Almeida (2001); 3, Benyon et al. (2006); 4, White et al. (2009); 5, Mendham et al. (2011).

Figure 1. A comparison of the climate of the three experimental sites: (a) Aracruz in Brazil, (b) DPI in the Green Triangle and (c) Scott River in Western
Australia (S. River). Data shown are monthly precipitation (bars, mm), monthly average maximum (■) and minimum (●) temperature (°C) and monthly

2
1
average global radiation (♦, MJ m day ).

Copyright © 2015 John Wiley & Sons, Ltd. Ecohydrol. 9, 610–630 (2016)
614 A. C. ALMEIDA AND P. J. SANDS
The catchment is located on the coast of Espírito Santo
state. It has an area of 286 ha with 190 ha of E. grandis
hybrid plantations at spacing of 3 m × 3 m located in the flat
part of the catchment favouring water infiltration rather than
run-off and 86 ha of preserved Atlantic rainforest in the
valley and riparian zone. Mean annual precipitation is
1250 mm, average temperature is 23·5 °C and solar radiation
averages 17 MJ m
2 day
1. Meteorological data were col-
lected in three automatic weather stations at 15-min, hourly
and daily intervals. Daylight VPD was calculated as in
Almeida and Landsberg (2003). Daily climate data were
used in the Aracruz site in order to compare the model
performance using daily or monthly time steps.
The soil at the Aracruz site is classified as Agrisol with a
predominant clay-loam texture; a detailed soil description
is in Almeida et al. (2004b). Soil moisture under the
plantation was measured at weekly or fortnightly intervals
from age 2 to 7·5 years using a neutron probe in access
tubes at 0·2-m intervals of depth to a maximum of 2·6 m
(Almeida et al., 2007; Smethurst et al., 2015). These data
were calibrated against volumetric SWC of samples
collected in the dry and wet seasons at each site, and the
results presented as total SWC. ASW was calculated as the
difference between measured SWC and the minimum SWC
over the measurement period. Fertility rating (FR) was
assigned as in Almeida et al. (2010), and the sites from
which data were collected were well managed (good weed
control and fertilizer applied).
Green Triangle sites
The eight Green Triangle sites used in this study were E.
globulus plantations planted between 1995 and 1998 and
described in detail by Benyon and Doody (2004) and
Benyon et al. (2006, 2007, 2008). Mean annual precipi-
tation varied from 500 to 800 mm, and potential ET was
980 mm year
1 (Wang et al., 2001).
For all Australian sites, daily maximum and minimum
air temperature, rainfall and solar radiation data were
obtained from the Scientific Information for Land Owners
(SILO) Data Drill (Jeffrey et al., 2001) from planting until
the last growth measurement at the site or the end of the
established rotation. SILO data are constructed by spatially
interpolating (on a regular 0·05° grid, with latitude,
longitude and elevation as independent variables) daily
data collected by the Australian Bureau of Meteorology
(Jeffrey et al., 2001). Daily SILO data were used to
calculate monthly average maximum and minimum air
temperature and solar radiation, monthly rainfall and
number of rain and frost days per month (i.e. minimum
air temperature < 0 °C). Soil texture had not been directly
measured in the Green Triangle sites but was taken from
the Victorian soils database (DNRE, 2002). Soils varied
from sandy loam/clay to clay loam/clay, depth ranged from
3·9 to 6·0 m, and water table levels varied from 3·3 to 20 m
Copyright © 2015 John Wiley & Sons, Ltd.
below the surface. Measurements of rainfall, canopy
interception, soil evaporation, tree transpiration and changes
in SWC (Benyon et al., 2008) were made approximately
monthly. SWC was measured at 0·3-m intervals of depth
using a neutron probe in tubes to a maximum depth of
5·85 m, and the changes in soil moisture between measure-
ments were evaluated for a period of at least 2 years. Details
of the measurements are given by Benyon et al. (2008).
The FR in the Green Triangle and WA sites was based
on a decision tree using soil type and previous land use
(Paul et al., 2007). Site FR was therefore generally
assumed to be between 0·7 and 0·9, or between 0·4 and
0·6 if soils were particularly infertile or growth was
impeded by salinity, shallow soil depth or poor drainage.
WA sites
In this study, we selected two contrasting sites in WA from
those described in detail by White et al. (2009) and
Mendham et al. (2011): Scott River and Wellstead. At
both, initial planting density was 1250 trees per hectare,
and growth, LAI and soil moisture had been measured.
Mean annual rainfall was 591 and 984 mm for these sites,
respectively. The soils in the Scott River and Wellstead
sites are deep (up to 8 and 18 m, respectively) with a sandy
A-horizon and show evidence of laterite in the top 2 m with
clay sub-soils (White et al., 2009). Measurements in these
plots were from September 1998 to September 2004.
Model calibration and validation
The original version of 3-PG was calibrated for E. globulus
grown in Australia (Sands and Landsberg, 2002; Booth et al.,
2006; Smettem et al., 2013), and Fontes et al. (2006) adapted
these parameters for E. globulus grown in Portugal. Almeida
et al. (2004b, 2007) provided a parameterization for E. grandis
grown in Brazil that predicted foliage, stem and root biomass
with remarkable accuracy. Esprey et al. (2004) found a very
similar set of parameters for E. grandis grown in South Africa.
These species-specific parameter sets were used as the basis for
the applications reported here, with a minor re-calibration for
the new water balance model required because of the interaction
between ASW and tree growth. This is described later.
An important parameter is the specific root volume σ zR,
i.e. the volume of soil explored by unit biomass roots, because
this determines the extent of the soil zone from which water is
taken up by the stand. Data were available for the Brazilian site
for root biomass and root depth as a function of stand age and
for a number of different clones of E. grandis (Almeida, 2003).
These data were used to assign σ zR = 2·5 for E. grandis.
Possible variation in σ zR is discussed later. The same value was
used for E. globulus because no alternative data were available.
A minor re-calibration for the new water balance model
because of the interaction between ASW and tree growth is
described in the following.
Ecohydrol. 9, 610–630 (2016)
 MODEL WATER BALANCE OF FOREST PLANTATIONS IN CONTRASTING ENVIRONMENTS 615

(m day
1)
The new input data required to run the model are the

kScond

0·2
0·5
0·5
1·0
0·5
0·5
0·5
1·0
0·5
5·0
0·5
following soil characteristics: soil profile depth (m);
volumetric SWC at saturation, field capacity and wilting
point (m3 m
3); saturated hydraulic conductivity (cm h
1);
drainage ratio (month
1); and parameters ES1 and ES2

(day
1)
kDrain

0·10
0·20
0·50
0·20
1·00
0·80
0·80
0·66
0·50
0·50
0·50
characterizing soil evaporation. These are presented in
Table II for each experimental site. The specific root
volume was estimated from root depth and biomass at

Esoil2

0·50
0·60
0·30
0·60
0·60
0·50
0·25
0·30
0·30
0·30
0·30
different ages (Almeida, 2003).
Water balance is dependent on the LAI of a stand, so any
error in the prediction of growth would impact on the

Esoil1

0·20
0·20
0·10
0·10
0·30
0·10
0·10
0·05
0·10
0·10
0·10
performance of the water balance submodel. Accordingly,
because the Aracruz site had a long series of observed LAI,
we tested the model’s performance when predicted LAI

nθ

5
5
5
5
8
7
8
8
5
9
5
was replaced with observed LAI in order to reduce
potential errors in the water balance due to errors in the

0·5
0·50
0·50
0·50
0·65
0·60
0·65
0·65
0·50
0·70
0·50
cθ
growth predictions. This approach did not produce any

Table II. Values of soil characteristics at each experimental site.

major changes in the results, largely because the original

(m3 m
3)
model produced excellent predictions of LAI (Almeida

SWwilt

0·240
0·270
0·380
0·350
0·125
0·235
0·090
0·060
0·290
0·120
0·170
et al., 2004b; Smettem et al., 2013).
Model efficiency (EF) and the root mean square error
(RMSE) were calculated to quantify the model accuracy. For
EF, observed and predicted values of selected variables were

(m3 m
3)
SWfcap

0·29
0·32
0·40
0·42
0·26
0·36
0·20
0·09
0·46
0·27
0·30
assessed using Equation (1) (Soares et al., 1995). EF can
vary from
1 (low accuracy) to +1 (high accuracy); when
positive, the model predicts better than average and, when
negative, worse than average. The RMSE [Equation (2)] is a
(m3 m
3)
SWsat

0·33
0·34
0·48
0·45
0·32
0·42
0·26
0·39
0·47
0·30
0·32
measure of the precision of the estimates in the same units as
the dependent variable.

2
∑ni¼1 Y i
Y^ i
0·8
0·8
0·7
0·8
0·8
0·8
0·9
0·9
0·8
0·6
0·6
FR

EF ¼ 1
n
2 (1)
∑i¼1 Y i
Y
SWC measured
Depth at which

sffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi

2
(m)

2·6
5·7
5·7
2·7
2·7
5·0
1·6
4·5
5·7
8·0
8·0
∑n Y i
Y^ i
RMSE ¼ i¼1
(2)
n

where Yi and Ŷi are the observed and predicted values,

depth (m)
Max root

respectively, Ῡ is the mean of the observed values and n

2·0
4·0
6·0
3·0
3·0
3·0
1·6
4·0
5·7
8·0
5·0

represents the total number of measurements.

Eucalyptus grandis hybrids grown in Brazil

depth (m)

An additional set of biomass observations (at age 7 years) is

Soil

2·6
6·0
6·0
4·3
3·9
5·0
1·8
5·0
6·0

5·0
15·0

now available following Almeida et al. (2004b), and the

original parameters for E. grandis grown in Brazil were
applied with the new model to simulate this extended set of
observations. Root biomass at 7 years was slightly under-
class
Soil

SCL
CL

CL

CL
LS

LS
LS
SL
LS
LS

predicted; the only change required to simulate the extended

data was to reduce root turnover from 0·02 to 0·015
(month
1). Simulation of SWC was improved by changing
Scott River
Padthaway

Wellstead

maximum canopy conductance, attained at an LAI of 3·0,

Gummy
Aracruz

from 0·020 to 0·024 (m s
1). In Almeida et al. (2004a), the 3-

Bessie

Jasper

Vikki
Jack
DPI
Site

Jill

PG parameter MaxAge in the age-related growth modifier

Copyright © 2015 John Wiley & Sons, Ltd. Ecohydrol. 9, 610–630 (2016)
616 A. C. ALMEIDA AND P. J. SANDS
fage was quoted as 9 years, but fage is in fact not used, and a
large value (>100 years) should be assigned.
Eucalyptus globulus grown in Australia
Few changes to parameters given by Sands and Landsberg
(2002) were required to adequately predict the growth of E.
globulus at the Green Triangle and WA sites: the power in the
stem mass allometric relationship was changed from 2·4 to
2·25; the minimum allocation of net primary production (NPP)
to roots from 0·25 to 0·1; maximum litterfall rate from 0·027 to
0·015 (month
1); and maximum canopy conductance, attained
and LAI 3·3, from 0·020 to 0·028 m s
1 (Schulze et al., 1994,
White et al., 1999); and the influence of fertility on canopy
quantum efficiency was set at 0·6, as used elsewhere. The sites
used by Sands and Landsberg (2002) all had FR = 1. These
changes were based on observed data from these sites and
related studies (Benyon et al., 2006; White et al., 2009).
Sensitivity analysis
A sensitivity analysis provides information on how
sensitive an output is to changes to, or uncertainties in,
species-specific parameters, site and soil characteristics or
other inputs. We used the concept of relative sensitivity
defined by Brylinsky (1972) and used by Battaglia and
Sands (1998) and Esprey et al. (2004).
Brylinsky (1972) defined the relative sensitivity λ(X, p)
of X with respect to p, as the change δX in X produced by a
change δp in p relative to the original values of X and p:
p ∂X
λðX ; pÞ ¼ (3)
X ∂p
Relative sensitivity is zero if X is independent of p and is
positive or negative depending on whether an increase in p
results in an increase or a decrease, respectively, in X.
Large values of λ indicate that X is highly sensitive to p,
while a value of λ close to zero indicates X is essentially
independent of p. In particular, if λ = 1, then a 10% change,
for example, in p will lead to a 10% change in X.
Esprey et al. (2004) also provide a simple computational
formula for determining λ from data obtained by running
the model with the parameter p increased and decreased by
a small amount δp. We determined λ for a range of model
outputs with respect to a range of soil characteristics, using
δp = 0·1p (i.e. 10% changes in p).
RESULTS
Growth and water balance at the Brazilian site
The model accurately predicted forest growth at the
Aracruz site. Figure 2 shows predicted and observed stem
and root biomass, diameter at breast height (DBH) and LAI
Copyright © 2015 John Wiley & Sons, Ltd.
forest growth from age 1 to 7·5 years. WB2 predicted 12%
lower stem mass than did WB1 at the end of the rotation.
Most of the difference occurred from age 5 to 7 years.
Soil water content was predicted using pseudo-daily and
daily time steps in WB2 and is compared with observed
data in Figure 3. Prediction accuracy (R2) of SWC was
86% with a pseudo-daily time step and 90% with a daily
time step. WB1 does not predict SWC. The standard error
(SE) of predicted SWC with WB2 was about 14% of the
maximum ASW at this site (172 mm) with a pseudo-daily
time step and 10% with a daily time step.
Available soil water was predicted using pseudo-daily and
daily time steps in WB2 and also with WB1 and is compared
with observed data in Figure 4. Prediction accuracy (R2) of
ASW was 85% with a pseudo-daily time step and 91% with a
daily time step, but only 67% with WB1 (which uses a
monthly time step). The SE of predicted ASW with WB1 was
about 15% of the maximum ASW at this site. For WB2, the
SE of predicted ASW was 11% with a pseudo-daily time step
and 7% with a daily time step.
WB2 was particularly effective at predicting soil
moisture changes after rainfall events and the reduction
of soil moisture by transpiration. In particular, the new
model, using both pseudo-daily and daily time steps,
predicted the low ASW observed early in the rotation and
from age 5 years onwards better than WB1 did. This led to
higher water stress and reduced growth at the end of the
rotation, as was observed. The ability of WB2 to predict
ASW early in the rotation (Figure 4a) was because root
depth depends on root biomass, which results in a more
realistic behaviour of root distribution in the soil profile,
and hence of water use. The volume of soil occupied by
roots is small after planting but increases as roots develop
over time up to the maximum root depth, achieved at age
3 years. A synthesis of the predicted water balance
components obtained with both WB1 and WB2 using
monthly, pseudo-daily and daily simulations is presented in
Table III for the Aracruz site. This table shows rainfall, ET
(i.e. ET, the sum of transpiration, soil evaporation and
rainfall interception), drainage and run-off accumulated
over a period of 7·5 years.
Daily estimates of SWC and ASW at the Aracruz site are
presented in Figures 3a and 4a, respectively. The predictions
were compared with 206 measurements made in the stand from
age 2 to 7·5 years. Values of ASW were inferred from SWC
measured (at 20-cm intervals with a neutron probe) down to the
root depth estimated by the model. It was assumed that the
minimum observed value of SWC corresponded to the wilting
point. The maximum root depth of 2·0 m was reached 3 years
after planting, and the corresponding maximum soil water
holding capacity is 172 mm. The use of the new model with
either a pseudo-daily or daily time step improved the estimation
of ASW when compared with the original model (which uses a
monthly time step). It also allows the direct prediction of SWC.
Ecohydrol. 9, 610–630 (2016)
 MODEL WATER BALANCE OF FOREST PLANTATIONS IN CONTRASTING ENVIRONMENTS 617

Figure 2. Stand growth predicted with WB2 (–––) and WB1 (– – –) for stem and root mass, leaf area index and DBH at the Aracruz site. Dot points (●)
are observed data.

Figure 3. Comparison of observed and predicted soil water content (SWC) data for the Aracruz site. (a) Observed (●) and time course of SWC predicted
by WB2 with pseudo-daily (­­­­­­­) and daily (–––) time steps and correlation between observed and predicted SWC using WB2 with (b) pseudo-daily
and (c) WB2 daily time steps. The 1-to-1 line is shown (······).

Copyright © 2015 John Wiley & Sons, Ltd. Ecohydrol. 9, 610–630 (2016)
618 A. C. ALMEIDA AND P. J. SANDS

Figure 4. Comparison of observed and predicted available soil water (ASW) data for the Aracruz site. (a) Observed (●) and predicted time course of
ASW by WB1 (------) and WB2 pseudo-daily (······) and daily (–––) time steps and correlation between predicted observed ASW using (b) WB2 pseudo-
daily, (c) WB2 daily and (d) WB1. The 1-to-1 line is shown (······).

Table III. Mean annual water balance components (mm) at the Aracruz site.

Evapotran Soil Canopy Rainfall

Rainfall spiration evaporation transpiration interception Drainage Run-off SWC ASW

Individual components of water balance

WB2 pseudo-daily simulation (mm) 7257 5585 1709 3229 647 1160 441
WB2 daily simulation (mm) 7276 5336 1219 3592 524 1029 790
WB1 monthly simulation (mm) 7257 5224 — — 777 — 1961
Components as % of rainfall
WB2 pseudo-daily simulation (%) 100 77·0 23·5 44·5 8·9 16·0 6·1
WB2 daily simulation (%) 100 73·3 16·8 49·4 7·2 14·2 10·8
WB1 monthly simulation (%) 100 72·0 — — 10·7 — 27·0
Annual averages
Annual average 967 771 710 56
– monthly simulation (mm)
Annual average 967 712 721 63
– daily simulation (mm)

Growth and water balance at the Australian sites
The Green Triangle and WA sites were analysed individ-
ually to evaluate model performance. Both WB1 and WB2
were applied to the Green Triangle site DPI because it had a
long series of observed data. Figure 5 compares nine outputs
from the two models with corresponding observations, and
this comparison shows WB2 provides a better fit to the
observed data.
Figure 6 compares the observed and predicted ET
(i.e. sum of interception, transpiration and soil evaporation)
for four sites in the Green Triangle region demonstrating a
reasonable agreement between observed and predicted except
for Vikki site. Predictions were made with WB2 using a
pseudo-daily time step. Figure 7 compares observed and
predicted SWC for six sites in the Green Triangle and WA at
which SWC had been observed. Predictions were with WB2
using a pseudo-daily time step. We compared measured SWC

Copyright © 2015 John Wiley & Sons, Ltd. Ecohydrol. 9, 610–630 (2016)
 MODEL WATER BALANCE OF FOREST PLANTATIONS IN CONTRASTING ENVIRONMENTS 619

Figure 5. Temporal variation of observed and predicted stand volume, diameter at breast height, leaf area index, evapotranspiration, soil water content,
canopy interception, available soil water (predicted only), soil evaporation and canopy transpiration for the DPI site in the Green Triangle region.
Observations are ●, predictions by WB1 are (­­­­­­­), and by WB2 are (–––).

with the predictions made by WB2 for similar periods using
data collected at dates close to the estimation periods.
An evaluation of the quality of the predictions of forest
growth, as represented by DBH, is presented in Table IV
for the Australian sites. This shows the precision and
accuracy of predicted DBH using the basic statistics of the
observed versus predicted regression lines, model effi-
ciency (EF) and the root mean square residual (RMSE).
Sensitivity analysis
We ran a sensitivity analysis for the Aracruz site using both
daily and pseudo-daily time steps to test the sensitivity of the
model to the new site-specific inputs (soil saturation, field
capacity and wilting point, drainage, hydraulic conductivity
and soil evaporation phases 1 and 2). The calculation of
sensitivity was based on 10% changes in each input, and
results were very similar for both time steps. Table V shows
the sensitivity of the biomass pools and each component of
the soil water balance accumulated over the entire rotation to
each of these new input data. Sensitivity may depend on the
stand age at which the analysis is made (e.g. Minunno et al.,
2013; Song et al., 2013). For instance, heavy rain at the age at
which the sensitivity analysis is determined can have
pronounced effects on transient fluxes of soil water.
Accordingly, it is generally advisable to base the sensitivity
analysis on variables that are accumulated over the entire
history of the stand, and our results are for variables
accumulated up to the end of the rotation. They clearly
indicate that the SWC at saturation, field capacity and wilting
point has strong effects on the water balance and, because this
affects soil water stress, also on stand biomass components.
DISCUSSION
Water use by Eucalyptus plantations has been a controver-
sial issue in various regions (Calder, 1992; Soares and

Copyright © 2015 John Wiley & Sons, Ltd. Ecohydrol. 9, 610–630 (2016)
620 A. C. ALMEIDA AND P. J. SANDS

Figure 6. Comparison of evapotranspiration predicted by WB2 using pseudo-daily time step with observed data for four experimental sites in the Green
Triangle region. The 1-to-1 line is shown (------).

Almeida, 2001; Dye, 1996; Dye and Versfeld, 2007;
Dvorak, 2012), particularly aggravated by the expansion
of plantations into drier areas. The impacts of this expansion
on water resources are often unknown, and inappropriate
generalizations and extrapolations of the results of other
studies have been made without considering local and
regional biophysical characteristics. In many cases, these
generalizations are driving policies or imposing restrictions
to expansion of plantations without scientific basis. Soil
physical characteristics can vary drastically in relatively
small areas and have a strong influence on water availability.
The lack of detailed maps of soil physical characteristics is
obviously a limitation for accurate spatial prediction of
plantation water balance (Feikema et al., 2010).
Models such as 3-PG are often used to predict plantation
growth and water use, and it is imperative that they provide
reliable predictions of plant and soil water balance. The
modifications (WB2) that we incorporated into the original
3-PG water balance submodel (WB1) were inspired by the
need to improve the water balance predictions.
The new water balance model assumes a single-layer
homogeneous soil profile, rather than multiple layers as
described by Soares and Almeida (2001). However, it does
subdivide the profile into a horizontal zone containing roots,
below which is a root-free zone. There is also an implicit
surface layer from which soil water is evaporated. Our
objection is to the use of a true multi-layered soil profile.
The model does not explicitly consider access to a water
table, e.g. by tap roots. In particular, the simplicity of 3-PG
does not allow the prediction of the vertical distribution of root
biomass. Smettem et al. (2013) suggested increasing the
maximum ASW at a site if it is suspected that a stand has access
to groundwater. However, we suggest setting a non-zero
minimum ASW such that if the root-zone ASW falls below this
value it is assumed the stand draws water from the water table
so as to maintain water stress above this minimum amount.
Disaggregation of the transpiration flux of WB1 into
canopy transpiration and soil evaporation as presented in
WB2 provides a more realistic estimation of the compo-
nents of the water balance because they are now separately
affected by the environment and are more realistically
modelled. These two fluxes can be compared with
measurements such as sap flow from individual trees and
mini-lysimeters used to measure soil evaporation.
Based on the results presented by applying WB1 and WB2
to the same sites, we conclude that the modifications leading
to WB2 improved the estimation of the SWC and ASW in
plantations of two species growing in contrasting environ-

Copyright © 2015 John Wiley & Sons, Ltd. Ecohydrol. 9, 610–630 (2016)
 MODEL WATER BALANCE OF FOREST PLANTATIONS IN CONTRASTING ENVIRONMENTS 621

Figure 7. Comparison of soil water content predicted by WB2 with a pseudo-daily time step with observed data at six experimental sites in Green
Triangle and Western Australia.

Table IV. Observed versus predicted regression analysis, model efficiency and accuracy for predicted DBH (cm) at the Australian sites.

Bessie DPI Gummy Jack Jasper Padthaway Jill Vikki Wellstead Scott River

Slope 0·65 0·92 0·73 0·61 0·76 1·39 0·71 1·07 1·45 1·14
Intercept 6·22 1·50 4·06 6·14 4·14
4·2 3·87 2·54
6·01
2·41
R2 0·86 0·95 0·98 0·90 0·94 0·98 0·95 0·99 0·99 0·98
N 39 37 67 73 24 17 74 57 7 7
EF 0·78 0·98 0·76 0·74 0·64
0·68 0·80 0·24 0·81 0·74
RMSE 0·50 0·14 1·05 0·79 0·42 0·57 0·64 1·45 1·49 0·98

WB2 with a pseudo-daily time step was used.

Copyright © 2015 John Wiley & Sons, Ltd. Ecohydrol. 9, 610–630 (2016)
 622

Table V. Relative sensitivity to various soil characteristics of the stand biomass components (WS, WF and WR), total and available soil water(SWC and ASW) and water balance
fluxes (ET, Transp, Esoil, Rainint, Run-off, Drainage) accumulated over the entire rotation.

Biomass pools Soil water contents Water fluxes

WS WF WR SWC ASW ET Transp Esoil Rainint Run-off Drainage

SWC at saturation 0·27
0·09 0·11 0·15 0·62 0·12 0·23
0·09 0·13
16·5 6·5
SWC at field capacity
0·05
0·03 0·15 0·66 2·9 0·01 0·02 0·02
0·09 7·9
10·5
SWC at wilting point
0·48 0·23
0·38
0·01
4·1
0·29
0·52 0·14
0·24 0·39 1·62

Copyright © 2015 John Wiley & Sons, Ltd.

Rate of drainage (kDrain)
0·03 0·01
0·01
0·02
0·08
0·01
0·02 0·01
0·01
3·7 0·21
Stage 1 soil evaporation (E1)
0·03
0·02
0·02 0·00
0·01 0·00
0·03 0·09
0·02
0·02
0·02
Stage 2 soil evaporation (E2)
0·12
0·12
0·08
0·01
0·03 0·01
0·14 0·34
0·08
0·05
0·05
Soil conductivity (kScond) 0·00 0·00 0·00 0·00 0·01 0·00 0·00 0·00 0·00 0·09
0·05

Data are based on runs with a daily time step. Sensitivities regarded as being exceptionally high are shown bold, while moderately high sensitivities are shown in italic.

Table VI. End-of-rotation predictions for growth, average soil water content and accumulated water balance fluxes for all study sites.

Site Soil evap Trans Rain

A. C. ALMEIDA AND P. J. SANDS

Age Stems WF WR WS Stand volume Average LAI SWC ASW Rainfall ET oration piration Drainage interception Run-off

years tree ha
1 t ha
1 t ha
1 t ha
1 m3 ha
1 m2 m
2 mm monthly average mm accumulated over rotation

Bessie 12 1100 6·8 16·4 91·2 204·1 2·6 1662 60 7966 7689 2290 4057 203 1342 0
DPI 12 950 5·3 14·1 65·0 145·4 1·9 2239 28 6816 6417 2126 3425 300 866 0
Gummy 12 1125 6·5 16·0 97·2 217·6 2·7 1040 51 7605 7658 2159 4152 168 1347 0
Jack 12 1175 6·6 17·7 97·2 232·0 3·0 454 106 7146 7912 2304 4285 120 1323 0
Jasper 12 750 5·5 15·6 75·0 167·8 1·9 1384 83 7218 6918 2588 3439 0 891 0
Padthaway 12 1025 4·1 9·9 52·6 117·7 1·8 180 41 5122 5000 1726 2678 163 596 0
Jill 12 1200 6·8 12·1 87·9 196·7 2·5 298 22 7369 7031 2195 3134 322 1702 0
Vikki 12 925 5·9 17·9 87·6 195·9 2·4 2109 166 7564 7784 2372 4194 248 1218 0
Wellstead 12 1250 5·8 21·5 88·9 198·8 3·0 1349 152 6487 8085 2041 4945 39 1099 0
Scott River 12 1250 5·2 53·3 91·3 170·5 2·4 1347 303 10 695 10 254 2245 6635 62 1374 0
Aracruz 7·5 1111 1·9 17·1 147·8 292·1 2·1 716 60 7257 5585 1709 3228 1160 647 441

Ecohydrol. 9, 610–630 (2016)

 MODEL WATER BALANCE OF FOREST PLANTATIONS IN CONTRASTING ENVIRONMENTS
ment: there is a better correspondence between the predicted
and observed forest growth and soil water balance. However,
we have no explanation as to why the site Vikki has such poor
predictions of ET and SWC (Figures 6 and 7) whereas
predictions of DBH at this site were good (Table IV).
This new submodel, however, requires more detailed
information about soil water characteristics and parameters
than did WB1. The required soil parameters can be derived
from soil texture data with a reasonable degree of confidence
(Clapp and Hornberger, 1978), but we cannot avoid the fact
that more texture-dependent soil data are required.
As expected, the use of a daily time step improved the
prediction of the water balance (Figures 3 and 4). Our results
are consistent with those of Feikema et al. (2010), who also
tested a modified version of 3-PG using a daily time step.
With the exception of one site (Vikki), our estimations of ET
have a similar accuracy to those reported by Feikema et al.
(2010). The latter also found that simulations of ASW using
a monthly time step are higher than using a daily time step.
However, our model explicitly takes into account a changing
root distribution during early stand development.
Daily climatic data are not always available. Also, their use
for spatial applications could be a limiting factor as increasing
the number of climatic surfaces adds complexity and
computational demands. If daily data are not available or if
its use is inappropriate for other practical reasons, either version
of the model can be run with a pseudo-daily time step. This
treats each month as a series of dry spells of equal length, with
rain falling on the first day of each spell. The use of a pseudo-
daily time step results in a marked improvement in the ability of
the model to predict finer details of the temporal variation of
soil water status, even with the original water balance model.
Prediction of water balance components for the 11 study
sites shows considerable differences between some sites.
This is shown in Table VI, which also shows an example of
the outputs of plantation growth and water balance for the
full rotations of the study sites. The only site that produced
run-off was at Aracruz. Over a full rotation (7·5 years for the
Brazilian site and 12 years for the Australian sites),
transpiration varied from 41% to 60% of the incoming
Figure 8. Analysis of specific root volume. (a) Observed values for Eucalyptus grandis at the Brazilian site as a function of stand age. (b) The effect of
varying specific root volume at this site on the stem biomass (–♦–) and total accumulated ET (–♦–) at the end of the rotation.
Copyright © 2015 John Wiley & Sons, Ltd.
623
rainfall, drainage from 0% to 16%, soil evaporation from
21% to 34% and canopy interception from 9% to 17%. This
variation in fluxes would be primarily due to the differences
in soil, species characteristics and climate at each site.
As the areas of new plantations increase, there is a clear
demand for better understanding of and ability to quantify
water use by plantations and the effects of water deficit on
forest growth. Understanding and quantifying water avail-
ability at the plot and catchment scales allow better
management of wood production by the forests and of the
water resources and help avoid potential conflicts caused by
the use of water. The practical implication of the new model
for forest management appears to be its potential to meet this
demand for better understanding and more accurate
quantification of water used by trees and water production
from catchments.
The results of the sensitivity analysis (Table V) indicate
that the water balance is highly sensitive to some of the soil
parameters required for the new model. Wilting point and
field capacity strongly affect ASW: field capacity and the
drainage parameter (kD) have respectively negative and
positive impacts on the prediction of run-off, and field
capacity also influences drainage and ET. The effects of the
soil characteristics on tree growth were less pronounced
and would be mediated through their effects on ASW, and
hence on the soil water growth modifier.
The specific root volume σ zR determines the extent of the
soil zone from which water is taken up by the stand. Data
for root biomass and root depth as a function of stand age
and for a number of different clones of E. grandis
(Almeida, 2003) were used to estimate σ zR, and the
resulting values are plotted in Figure 8a for the clone used
in this study. It is clear that σ zR declines with stand age, but
there is considerable variation at a given stand age. (Similar
values were obtained for other clones but are not shown.) It
is clear σ zR may vary with stand age, and we would expect
possible variations with soil texture and between species.
We explored the possible effects of changes in σ zR by
running the model for the Brazilian site using radically
different values. Figure 8b shows the variation in stem
Ecohydrol. 9, 610–630 (2016)
624 A. C. ALMEIDA AND P. J. SANDS
biomass and accumulated ET at the end of the rotation with
σ zR. It is clear that in this case that values of σ zR >1 are
equivalent, but there are strong effects for σ zR < 1. Small
values of σ zR mean the roots are slower in exploring the soil
profile and hence water stress effects are larger later in
stand growth. In our case, the value used for σ zR was that
observed for a young stand and hence the predicted root
depth matched observed. Once the roots attain maximum
depth, the subsequent value of σ zR is largely irrelevant.
Our objective was to improve the prediction of water
balance without losing the simplicity of the 3-PG model;
we demonstrated that the new model achieves this
objective. There are advantages and disadvantages in using
the new submodel, and the decision about which submodel
will be more appropriate should be based on the questions
to be answered, and obviously on the availability of data.
We believe that there is no advantage in using the WB2
submodel if the soil information is very limited or
inadequate. On the other hand, if a more precise water
balance is required, this new submodel proves to be a better
approach and provides more accurate predictions of the
growth and water use by plantations. Finally, if daily data
and detailed soils data are both not available, the original
soil water balance model can be improved by using a
pseudo-daily time step in lieu of a monthly time step.
CONCLUSIONS
Analyses of the predicted growth and water balance in three
contrasting regions show that the new 3-PG water balance
submodel was able to improve the predictions of the water
balance components at widely contrasting sites, and for two
species. Moreover, use of the pseudo-daily time step still
provides improved simulation of water balance when using
the original water balance submodel because it better captures
the dynamics of rainfall distribution. The new submodel
requires more detailed information of the soil characteristics,
and the water balance is sensitive to these parameters. It is our
intention to make the new model also freely available.
ACKNOWLEDGEMENTS
Richard Benyon and Tanya Doody provided the Green
Triangle data, Don White provided the WA dataset and Fibria
(formerly Aracruz Celulose) supported the data collection
from the Aracruz site as published in Almeida (2003). We
also appreciate Keryn Paul and Anders Siggins for their
contributions to data collation and Phil Polglase for his
incentive to write this paper. We thank Joe Landsberg,
Anthony O’Grady and Jacqui England for reviewing the
manuscript and Sadanandan Nambiar for his comments. We
acknowledge the farmers in WA, VIC and SA who allowed us
access to their properties for data collection. We also thank
Copyright © 2015 John Wiley & Sons, Ltd.
Dick Waring and an anonymous reviewer for their comments
on an earlier draft of the manuscript.
APPENDIX A
WATER BALANCE SUBMODEL EQUATIONS
A.1 Hydrological balance
The basic equation governing the hydrological balance
of a soil profile specifies the change in the SWC Θ in some
time interval Δt in terms of the rainfall R, possibly
including any irrigation, and various losses that occur in
that interval. Thus (L&S, Section 7.1),
Θðt þ Δt Þ ¼ Θðt Þ þ R
I R
E C
ES
qD
qR (A1)
where all units are millimetres (or kilograms per square
metre) and the losses are calculated over the time interval
Δt. The losses are the amount IR of rainfall intercepted by
the canopy and subsequently evaporated, canopy transpi-
ration EC, evaporation ES from the soil, the amount qD of
water that drains out of the bottom of the profile and
surface run-off qR. Lateral flow is not considered in 3-PG.
If the soil profile depth is z (m), then the volumetric
SWC θ (m3 m
3 or mm m
1) of the profile is given by
θ = Θ/z. The relative plant ASW θr is the ratio of the current
plant ASW to the maximum ASW:


θ
θwp
θr ¼
 (A2)
θf c
θwp
where θwp and θfc are the volumetric SWCs (mm) of the
profile at wilting point and field capacity, respectively.
A.2 Soil water stress and the soil water growth modifier
Water stress is determined by the water content of a zone
of soil around the roots. The size of this zone changes
dynamically as plants grow and the root system occupies
more of the soil profile. All transpiration is drawn from the
root zone, which consequently dries out far more rapidly
than the bulk soil. Recharge of the root zone is modelled
(Appendix A.7) by movement of water at a rate
proportional to the difference in volumetric SWC in the
two zones and depends on the saturated hydraulic
conductivity of each soil.
The soil water growth modifier (fθ) determines the
effects of soil water stress on growth and canopy
conductance and is based on relative plant ASW (θr) in
the root zone, not in the entire soil profile. It is defined so
that it varies between zero and one while soil water varies
between no plant available water (θr = 0) to unlimited soil
Ecohydrol. 9, 610–630 (2016)
 MODEL WATER BALANCE OF FOREST PLANTATIONS IN CONTRASTING ENVIRONMENTS
water (θr = 1). Thus,
1
ð1
θ r Þn θ
f θ ðθ r Þ ¼
1 þ ð1
2cθ nθ Þ½ð1
θr Þ=cθ nθ
where the parameters cθ and nθ are soil texture-dependent
parameters (L&S, Section A2.1).
A.3 Rainfall interception
The fraction of rainfall intercepted by the canopy is
assumed to be proportional to canopy LAI L, up to some
maximum fraction iRx that occurs at an LAI of LRx (L&S,
Section 7.2.2), where LRx and iRx are species-specific
parameters. The rainfall interception loss IR is
I R ¼ iRx minf1; L=LRx gR
A.4 Canopy transpiration
Daily canopy transpiration E C (kg m
2 day
1 or
mm day
1) is modelled as in the original 3-PG water
balance submodel using the Penman–Monteith equation
(L&S, Section 7.2.1):


gC sφna þ gb ρa cpa D
EC ¼ h
λððγ þ sÞgC þ γg aC Þ
Here, h (s day
1) is the day length, φna (W m
2) is the daily
average net absorbed radiation, D (Pa) is the atmospheric
VPD above the canopy, gC (m s
1) is the canopy conductance
and gaC (m s
1) is the aerodynamic boundary layer
conductance of the canopy. Other quantities are the latent
heat λ = 2·45 MJ kg
1 of vaporization of water, the density
ρa = 1·204 kg m
3 and specific heat cpa = 1004 J kg
1 K
1 of
dry air, the psychrometric constant γ = 66·1 Pa K
1 and the
derivative s = 145 Pa K
1 (at 20 °C) of the saturated vapour
pressure with respect to temperature.
Because EC is canopy transpiration and does not include
evaporation from the soil, φna is the radiation absorbed by
the canopy, not incident upon it. Almeida et al. (2003)
found
φna ¼ a þ bφi

2
where φi (W m ) is the average daily intercepted radiation,
and a and b have values close to 0 and 0·8, respectively.
From Beers law, φi is related to daily insolation Q
(MJ m
2 day
1) by
Q
 and that prior to canopy closure part of the soil is in direct
φi ¼ 106 1
e
kL ;
h sun. The analogue of gC is very large, but aerodynamic
Copyright © 2015 John Wiley & Sons, Ltd.
625
where L is canopy LAI. Because soil evaporation is
separately modelled in WB2, canopy conductance gC
declines to zero as L declines to zero and is the product of
(A3)
the optimal conductance gCopt (m s
1) and 3-PG growth
modifiers, as in L&S [Equation (9.16)], with
 
L
g Copt ðLÞ ¼ min ; 1 g Cx (A8)
LgCx
where LgCx is the value of L at which conductance attains its
maximum value gCx (m s
1). LgCx and gCx are species-
specific parameters.
A.5 Soil evaporation
(A4)
Soil evaporation is modelled using the so-called two-
phase Ritchie (1972) model. Soil evaporation is assumed to
be confined to a shallow upper soil layer that acts as a
barrier to further evaporation as the soil dries. Evaporation
is high when the soil surface is wet after rain and declines
as it dries out. Evaporation from soil in full sun and that
from soil shaded by the canopy are treated separately.
In Phase 1 (immediately after a wetting event), the rate
eS (mm day
1) of soil evaporation is the wet-surface rate, e0
(mm day
1). Phase 2 commences once accumulated
(A5)
evaporation ES (mm) since the last wetting event exceeds
an amount ES1, and eS is then assumed to decline
hyperbolically with increasing evaporation accumulated
above that in Phase 1. The rate of decline is described by a
parameter ES2, which is the amount of evaporation in Phase
2 required to reduce eS by 50%. Thus,
8
ES ≤ ES1
dE S <
e0
eS ¼ ¼ e0 (A9)
dt : E S1 < E S
1 þ ðE S
E S1 Þ=E S2
which can be integrated to give
(
e0 t t ≤ t S1
ES ðt Þ ¼
pffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffi 
ES1 þ E S2 1 þ 2ðe0 =E S2 Þðt
t S1 Þ
1 t > t S1
(A6)
(A10)
where tS1 = ES1/e0 is the duration of Phase 1 evaporation
(L&S, Section 7.2.4). The wet-surface rate e0 is calculated
using the Penman–Monteith equation [Equation (A5)] and
is driven by radiation incident on the soil and the VPD
above the soil, taking into account the presence of a canopy
(A7)
Ecohydrol. 9, 610–630 (2016)
626 A. C. ALMEIDA AND P. J. SANDS
conductance gas above the soil is smaller than canopy
aerodynamic conductance, e.g. gas ≈ 0·01gac. There are
three new parameters: ES0 and ES1, which we assume are
texture-dependent soil properties, and the aerodynamic
conductance gAs.
The water balance submodel keeps track of the
accumulated net evaporation ES from the soil, as this
determines subsequent soil evaporation. Because of this,
there is no need to explicitly include a separate surface soil
layer in the model. When a rainfall event occurs, ES is
reduced by the amount of throughfall (subject to ES ≥ 0).
This forms the initial condition ES0 for the subsequent
period of soil evaporation, and Equation (A10) is inverted
and solved for an equivalent initial time t0. If the time to the
next rainfall event is tR, soil evaporation in the intervening
period is ES(tR + t0)
ES0. For the daily time step model,
tR = 1. For monthly or pseudo-daily time steps, tR is days in
the month/number of rain days, and soil evaporation is
computed separately for each rainfall event and then
summed over all events.
Although the Ritchie model was originally empirical, it
can be derived from physical principles after making some
simple approximations. The parameters ES0 and ES1 are
then expressed explicitly in terms of various textural
properties of the soil.
A.6 Run-off and drainage
If rainfall saturates the soil, i.e. volumetric water
content θ exceeds saturation (θsat), the excess above
saturation is deemed to be surface run-off. The remaining
soil water is available for transpiration and evaporation.
If θ exceeds field capacity (θfc), this excess is available
for drainage, which proceeds at a rate proportional to the
excess, where the rate constant kDrain (day
1) is a texture-
dependent soil property. Drainage qD over t days is then
given by


 V rz
qD ðt Þ ¼ Θ0
V Soil θf c 1
e
k Drain t (A11)
where Θ0 is the initial SWC, VSoil is the volume of soil
and θfc is the volumetric soil content at field capacity.
Drainage is first determined from the root zone into the
root-free zone and then from the root-free zone out of
the bottom of the profile. For a daily time step, t = 1; for
the pseudo-daily time step, t = number of days between
rainfall events, i.e. days in month/rain days; and with a
monthly time step, t = number of days in the month.
A.7 Root-zone growth and recharge
The root-zone concept introduced in this paper is simple
and pragmatic. The soil profile is divided into a root-free
Copyright © 2015 John Wiley & Sons, Ltd.
zone and a root zone. The root zone is that volume of soil
from which the current stock of roots can access soil water.
Its depth zR (m) is determined from the root biomass WR
(t ha
1) by
zR ¼ minf0:1σ zR W R =ð1
pstones Þ; zRx g (A12)
where σ zR (m3 kg
1) is the volume of soil explored by 1 kg of
root biomass and the 0·1 converts tons per hectare to
kilograms per square metre. As roots grow, the root-zone
volume increases at the expense of the root-free volume. The
SWCs of these two volumes are adjusted to take this into
account.
Water also moves from the root-free zone into the root
zone along the water pressure gradient induced by
differences in their volumetric SWCs. If two regions have
volumetric SWCs θ1 and θ2, with θ1 < θ2, the flux of water
from regions 2 to 1 is assumed to be
F V ¼
k S ðθ1
θ2 Þ (A13)
where FV (m3water m
2 day
1) is the volume of water
crossing unit area in a day and kS (m day
1) is a texture-
dependent measure of the hydraulic conductivity of the
soil. Root-zone recharge is modelled by assuming that
during recharge the volumes of the two zones are constant,
the total amount of water in the two zones is constant and
they share a common area A (m2). Application of Equation
(A13) then shows that the temporal variation of the SWC
Θrz(t) of the root zone is given by
Θrz0 V nr
Θnr0 V rz
t=tS0
Θrz ðt Þ ¼ e
V rz þ V nr
(A14)
þ ðΘrz0 þ Θnr0 Þ
V rz þ V nr
Here, Θrz0 and Θnr0 are the SWCs of the root and non-
root zones at time t = 0, Vrz and Vnr are the volumes of the
root and non-root zones and tS0 is the time constant for
water movement and is given by
t S0 ¼ V rz V nr =k S AðV rz þ V nr Þ (A15)
Equations (A14) and (A15) determine the SWC of the
root zone after t days.
Ecohydrol. 9, 610–630 (2016)
 MODEL WATER BALANCE OF FOREST PLANTATIONS IN CONTRASTING ENVIRONMENTS 627

APPENDIX B
3-PG PARAMETERS FOR E. GLOBULUS AND E. GRANDIS HYBRIDS APPLIED IN THIS STUDY
Meaning/comments Name Units E. grandis E. globulus

Biomass partitioning and turnover

Allometric relationships and partitioning
Foliage : stem partitioning ratio at D = 2 cm pFS2 — 0·7 1·0
Foliage : stem partitioning ratio at D = 20 cm pFS20 — 0·100 0·150
Constant in the stem mass-versus-diameter relationship aWS — 0·049 0·095
Power in the stem mass-versus-diameter relationship nWS — 2·822 2·25
Constant in the stem mass-versus-diameter relationship aWR — 0 0
Power in the stem mass-versus-diameter relationship nWR — 0 0
Maximum fraction of NPP to roots pRx — 0·6 0·8
Minimum fraction of NPP to roots pRn — 0·07 0·20
Fraction of total root allocation going to fine roots pRfine — 0·01 0·5
Volume of soil accessed by 1 kg of root dry matter spRootVol m3 kg
1 root 2·5 2.5
Litterfall and root turnover
Maximum litterfall rate gammaF1 month
1 0·12 0·015
Litterfall rate at t = 0 gammaF0 month
1 0·00169 0·00100
Age at which litterfall rate has median value tgammaF months 13 12
Average monthly branch and bark litterfall rate gammaS month
1 0 0
Average monthly coarse root turnover rate gammaR month
1 0·02 0·015
Average monthly fine root turnover rate gammaRf month
1 0·25 0·25
NPP and conductance modifiers
Temperature modifier (gmTemp)
Minimum temperature for growth Tmin °C 8·0 8·5
Optimum temperature for growth Topt °C 25·0 16·0
Maximum temperature for growth Tmax °C 36·0 40·0
Frost modifier (gmFrost)
Days production lost per frost day kF days 0 0
Fertility effects (gmNutr)
Value of m when FR = 0 m0 — 0 0
Value of fNutr when FR = 0 fN0 — 0·6 0·6
Power of (1
FR) in gmNutr fNn — 1 1
Atmospheric CO2 effects
Ratio of alpha at 700 and 350 ppm gmCalpha700 — 1·4 1·4
Ratio of canopy conductance at 700 and 350 ppm gmCg700 — 0·7 0·7
Salinity effects (gmSalt)
Salinity below which no effects of salt on growth EC0 dS m
1 999 999
Salinity above which growth ceases EC1 dS m
1 999 999
Power of EC in gmSalt ECn — 1 1
Age modifier (gmAge)
Maximum stand age used in age modifier MaxAge years 20 50
Power of relative age in function for fAge nAge — 4 4
Relative age at fAge = 0·5 rAge — 0·95 0·95
Stem mortality and self-thinning
Mortality rate for large t gammaN1 % year
1 0 0
Seedling mortality rate (t = 0) gammaN0 % year
1 0 0
Age at which mortality rate has median value tgammaN years 0 0
Shape of mortality response ngammaN — 1 1
Maximum stem mass per tree at 1000 trees per hectare wSx1000 kg tree
1 300 300
Power in self-thinning rule thinPower — 1·5 1·5
Fraction mean single-tree foliage biomass lost per dead tree mF — 0 0
Fraction mean single-tree root biomass lost per dead tree mR — 0·2 0·2
Fraction mean single-tree stem biomass lost per dead tree mS — 0·5 0·2
Canopy structure and processes
Specific leaf area
Specific leaf area at age 0 SLA0 m2 kg
1 11 11
Specific leaf area for mature leaves SLA1 m2 kg
1 8 4
Age at which specific leaf area = (SLA0 + SLA1)/2 tSLA years 2·5 2·5
Light interception and VPD attenuation
Extinction coefficient for absorption of PAR by canopy k — 0·5 0·5
Age at canopy cover fullCanAge years 3 3
(Continues)

Copyright © 2015 John Wiley & Sons, Ltd. Ecohydrol. 9, 610–630 (2016)
628 A. C. ALMEIDA AND P. J. SANDS

APPENDIX B (Continued)
Meaning/comments Name Units E. grandis E. globulus

LAI for 50% reduction of VPD in canopy cVPD0 5 5

Rainfall interception
Maximum proportion of rainfall evaporated from canopy MaxIntcptn — 0·15 0·20
LAI for maximum rainfall interception LAImaxIntcptn — 3 3
Production and respiration
Canopy quantum efficiency alphaCx molC/molPAR 0·068 0·060
Edge tree growth % enhancement edgeEffect — 0 0
Ratio NPP/gross primary production Y — 0·47 0·47
Conductance
Maximum stomatal conductance gSx m s
1 0·008 0·008
Radiation for gS = gSx/2 IgS W m
2 100 100
LAI for maximum canopy conductance LAIgcx — 3·00 3·33
Minimum canopy conductance MinCond m s
1 0 0
Maximum canopy conductance MaxCond m s
1 0·024 0·026
Defines stomatal response to VPD CoeffCond mbar
1 0·045 0·050
Canopy aerodynamic conductance gAc m s
1 0·2 0·2
Soil aerodynamic conductance gAs m s
1 0·01 0·02
Wood and stand properties
Branch and bark fraction (fracBB)
Branch and bark fraction at age 0 fracBB0 — 0 0·45
Branch and bark fraction for mature stands fracBB1 — 0 0·15
Age at which fracBB = (fracBB0 + fracBB1)/2 tBB years 0 2
Basic density
Minimum basic density – for young trees rho0 t m
3 0·48 0·40
Maximum basic density – for older trees rho1 t m
3 0·52 0·45
Age at which rho = (rhoMin + rhoMax)/2 tRho years 5 3
Stem height
Constant in the stem height relationship aH — 0·42 0·39
Power of DBH in the stem height relationship nHB — 1·50 1·41
Power of stocking in the stem height relationship nHN — 0 0·00001
Stem volume
Constant in the stem volume relationship aV — 0 0
Power of DBH in the stem volume relationship nVB — 0 0
Power of stocking in the stem volume relationship nVN — 0 0
Conversion factors
Intercept of net-versus-solar radiation relationship Qa W m
2
8·85 0
Slope of net-versus-solar radiation relationship Qb — 0·82 0·8
Molecular weight of dry matter gDM_mol gDM mol
1 24 24
Conversion of solar radiation to PAR molPAR_MJ mol MJ
1 2·3 2·3

REFERENCES
Almeida AC. 2003. Application of a process-based model for predicting
an explaining growth in Eucalyptus plantations, PhD Thesis, The
Australian National University, 232 pp.
Almeida AC, Landsberg JJ. 2003. Evaluating methods of estimating
global radiation and vapor pressure deficit using a dense network of
automatic weather stations in coastal Brazil. Agricultural and Forest
Meteorology 118: 237–250. DOI:10.1016/S0168-1923(03)00122-9.
Almeida AC, Landsberg JJ, Sands PJ, Ambrogi MS, Fonseca S, Barddal
SM, Bertolucci FL. 2004a. Needs and opportunities for using a process-
based productivity model as a practical tool in Eucalyptus plantations.
Forest Ecology and Management 193: 167–177. DOI:10.1016/j.
foreco.2004.01.044.
Almeida AC, Landsberg JJ, Sands PJ. 2004b. Parameterisation of 3-PG
model for fast-growing Eucalyptus grandis plantations. Forest Ecology
and Management 193: 179–195. DOI:10.1016/j.foreco.2004.01.029.
Almeida AC, Soares JV, Landsberg JJ, Rezende GD. 2007. Growth and
water balance of Eucalyptus grandis hybrid plantations in Brazil during
a rotation for pulp production. Forest Ecology and Management 251:
10–21. DOI:10.1016/j.foreco.2007.06.009.
Almeida AC, Sands PJ, Bruce J, Siggins AW, Leriche A, Battaglia M,
Batista TR. 2009. Use of a spatial process-based model to quantify
forest plantation productivity and water use efficiency under climate
change scenarios. 18th World IMACS / MODSIM Congress, Cairns,
1816–1822.
Almeida AC, Siggins A, Batista TR, Beadle C, Fonseca S, Loos R. 2010.
Mapping the effect of spatial and temporal variation in climate and soils
on Eucalyptus plantation production with 3-PG, a process-based growth
model. Forest Ecology and Management 259: 1730–1740.
DOI:10.1016/j.foreco.2009.10.008.
Battaglia M, Sands P. 1998. Application of sensitivity analysis to a model
of Eucalyptus globulus plantation productivity. Ecological Modelling
111: 237–259. DOI:10.1016/S0304-3800(98)00114-8.
Battaglia M, Sands P, White D, Mummery D. 2004. CABALA: a linked
carbon, water and nitrogen model of forest growth for silvicultural
decision support. Forest Ecology and Management 193: 251–282.
DOI:10.1016/j.foreco.2004.01.033.

Copyright © 2015 John Wiley & Sons, Ltd. Ecohydrol. 9, 610–630 (2016)
 MODEL WATER BALANCE OF FOREST PLANTATIONS IN CONTRASTING ENVIRONMENTS
Benyon RG, Doody T. 2004. Water use by tree plantations in south east
south Australia. Technical Report No. 148, 1–26, Australia, CSIRO.
Benyon R, Theiveyanathan S, Doody TM. 2006. Impacts of tree
plantations on groundwater in south-eastern Australia. Australian
Journal of Botany 54: 181–192. DOI:10.1071/BT05046.
Benyon R, England J, Eastham J, Polglase P, White D. 2007. Tree water
use in forestry compared to other dry-land agricultural crops in the
Victorian context. Report prepared for the Department of Primary
Industries Victoria to promote scientific knowledge in this area. Ensis
Technical Report No. 159, 50 p.
Benyon RG, Doody TM, Theiveyanathan S, Koul V. 2008. Plantation forest
water use in Southwest Victoria. Technical Report No. 164, CSIRO, 109 pp.
Booth TH., Paul KI, Jovanovic T. 2006. Predicting potential plantation
growth in the South West Goulburn Broken. Final Report May 2006,
Client Report No. 1669.
Brylinsky M. 1972. Steady-state sensitivity analysis of energy flow in a
marine ecosystem. In Systems Analysis and Simulation in Ecology, vol.
2, Patten BC (ed). Academic Press: New York; 81–101.
Calder IR. 1992. Water use of Eucalypts – a review. In Growth and Water Use
of Forest Plantations, Calder IR, Hall RL, Adlard PG (eds). John Wiley &
Sons Ltd: Chichester, New York, Brisbane, Toronto, Singapore; 167–179.
Clapp RB, Hornberger GM. 1978. Empirical equations for some soil
hydraulic properties. Water Resources Research 14: 601–604.
DOI:10.1029/WR014i004p00601.
Coops NC, Waring RH. 2001. The use of multiscale remote sensing
imagery to derive regional estimates of forest growth capacity using 3-
PGS. Remote Sensing and Environment 75: 324–334. DOI:10.1016/
S0034-4257(00)00176-0.
Coops NC. 1999. Improvement in predicting stand growth of Pinus
radiata (D. Don) across landscapes using NOAA AVHRR and Landsat
MSS imagery combined with a forest growth process model (3-PGS).
Photogrammetric Engineering & Remote Sensing 65: 1149–1156.
DNRE. 2002. The 1:125,000 statewide soil attribute coverage documen-
tation. Version 1.1, June 2002, Centre for Land Protection Research,
Department of Natural Resources and Environment, Victoria.
Dvorak WS. 2012. Water use in plantations of eucalypts and pines: a
discussion paper from a tree breeding perspective. International
Forestry Review 14: 110–119. DOI:10.1505/146554812799973118.
Dye PJ. 1996. Response of Eucalyptus grandis trees to soil water deficits.
Tree Physiology 16: 233–238. DOI:10.1093/treephys/16.1-2.233.
Dye PJ. 2001. Modelling growth and water use in four Pinus patula stands
with the 3-PG model. Southern African Forestry Journal 191: 53–63.
Dye P, Versfeld D. 2007. Managing the hydrological impacts of South
African plantation forests: An overview. Forest Ecology and Manage-
ment 251: 121–128. DOI:10.1016/j.foreco.2007.06.013.
Esprey LJ, Sands PJ, Smith CW. 2004. Understanding 3-PG using a
sensitivity analysis. Forest Ecology and Management 193: 235–250.
DOI:10.1016/j.foreco.2004.01.032.
Feikema PM, Morris JD, Beverly CR, Collopy JJ, Baker TG, Lane PNJ.
2010. Validation of plantation transpiration in south-eastern Australia
estimated using the 3PG+ forest growth model. Forest Ecology and
Management 260: 663–678. DOI:10.1016/j.foreco.2010.05.022.
Fontes L, Landsberg J, Tomé J, Tomé M, Pacheco CA, Soares P, Araujo
C. 2006. Calibration and testing of a generalized process-based model
for use in Portuguese eucalyptus plantations. Canadian Journal of
Forestry Research 36: 3209–3221. DOI:10.1139/x06-186.
Forrester DI, Tang X. 2015. Analysing the spatial and temporal dynamics
of species interactions in mixed-species forests and the effects of stand
density using the 3-PG model. Ecological modelling. DOI:10.1016/j.
ecolmodel.2015.07.010.
Gonzalez-Garcia M, Almeida AC, Hevia A, Majada J, Beadle C. 2015.
Application of a process-based model for predicting the productivity of
Eucalyptus nitens plantations grown for bioenergy in Spain. Global
Change Biology - Bioenergy. DOI:10.1111/gcbb.12256.
Jeffrey SJ, Carter JO, Moodie KB, Beswick AR. 2001. Using spatial
interpolation to construct a comprehensive archive of Australian climate
data. Environmental Modelling & Software 16: 309–330. DOI:10.1016/
S1364-8152(01)00008-1.
Landsberg JJ, Waring RH. 1997. A generalised model of forest
productivity using simplified concepts of radiation-use efficiency,
carbon balance and partitioning. Forest Ecology and Management 95:
209–228. DOI:10.1016/S0378-1127(97)00026-1.
Copyright © 2015 John Wiley & Sons, Ltd.
629
Landsberg JJ, Waring RH, Coops NC. 2003. Performance of the forest
productivity model 3-PG applied to a wide range of forest types. Forest
Ecology and Management 172: 199–214. DOI:10.1016/S0378-1127
(01)00804-0.
Landsberg JJ. 2003. Modelling forest ecosystems: state-of-the-art,
challenges and future directions. Canadian Journal of Forest Research
33: 385–397. DOI:10.1139/X02-129.
Landsberg JJ, Sands PJ. 2010. Physiological Ecology of Forest Production.
Principles, Processes and Models. Elsevier: London, 352 pp.
Mendham DS, White DA, Battaglia M, McGrath JF, Short TM, Ogden
GN, Kinal J. 2011. Soil water depletion and replenishment during first-
and early second-rotation Eucalyptus globulus plantations with deep
soil profiles. Agricultural and Forest Meteorology 151: 1568–1579.
DOI:10.1016/j.agrformet.2011.06.014.
Miehle P, Battaglia M, Sands PJ, Forrester DI, Feikema PM, Livesley SJ,
Morris JD, Arndt S. 2009. A comparison of four process-based model
and statistical regression model to predict growth of Eucalyptus
globulus plantations. Ecological Modelling 220: 734–746.
DOI:10.1016/j.ecolmodel.2008.12.010.
Mielke MS, Oliva MA, Barros NF, Penchel RM, Martinez CA, Almeida
AC. 1999. Stomatal control of transpiration in the canopy of a clonal
Eucalyptus grandis plantation. Trees 13: 152–160.
Minunno F, Van Oijen M, Cameron DR, Cerasoli S, Pereira JS, Tomé M.
2013. Using a Bayesian framework and global sensitivity analysis to
identify strengths and weaknesses of two process-based models
differing in representation of autotrophic respiration. Environmental
Modelling & Software 42: 99–115. DOI:10.1016/j.envsoft.2012.12.010.
Paul KI, Polglase PJ, Snowdon P, Theiveyanathan S, Raison J, Grove T,
Rance S. 2006. Calibration and uncertainty analysis of a carbon
accounting model to stem wood density and partitioning of biomass for
Eucalyptus globulus and Pinus radiata. New Forests 31: 513–533.
DOI:10.1007/s11056-005-2740-4.
Paul KI, Booth TH, Jovanovic T, Sands PJ, Morris JD. 2007. Calibration
of the forest growth model 3-PG to eucalypt plantations growing in low
rainfall regions of Australia. Forest Ecology and Management 243:
237–247. DOI:10.1016/j.foreco.2007.03.029.
Pérez-Cruzado C, Muñoz-Sáez F, Basurco F, Riesco G, Rodríguez-Soalleiro
R. 2011. Combining empirical models and the process-based model 3-PG
to predict Eucalyptus nitens plantations growth in Spain. Forest Ecology
and Management 262: 1067–1077. DOI:10.1016/j.foreco.2011.05.045.
Ritchie JT. 1972. Model for predicting evaporation from row crop with
incomplete cover. Water Resources Research 8: 1204–1213.
Rodríguez R, Espionosa M, Real P, Inzunza J. 2002. Analysis of productivity
of radiata pine plantations under different silvicultural regimes using 3-PG
process-based model. Australian Forestry 65: 165–172.
Rodriguez R, Real P, Espinosa M, Perry DA. 2009. A process-based
model to evaluate site quality for Eucalyptus nitens in the Bio-Bio
Region of Chile. Forestry 82: 149–162. DOI:10.1093/forestry/cpn045.
Running SW, Coughlan JC. 1988. A general model of forest ecosystem
processes for regional applications I. Hydrologic balance, canopy gas
exchange and primary production processes. Ecological Modelling 42:
125–154. DOI:10.1016/0304-3800(88)90112-3.
Sands PJ. 2004. 3PGpjs vsn 2.4 – a user-friendly interface to 3-PG, the
Landsberg and Waring model of forest productivity. Technical Report.
No. 140, CRC Sustainable Production Forestry, Hobart.
Sands PJ, Landsberg JJ. 2002. Parameterisation of 3-PG for plantation
grown Eucalyptus globulus. Forest Ecology and Management 163:
273–292. DOI:10.1016/S0378-1127(01)00586-2.
Saxton KE, Rawls WJ. 2006. Soil water characteristic estimates by texture
and organic matter for hydrologic solutions. Soil Science Society of
America 70: 1569–1578.
Schulze ED, Kelliher FM, Korner C, Lloyd J, Leuning R. 1994. Relationships
among maximum stomatal conductance, ecosystem surface conductance,
carbon assimilation rate, and plant nitrogen nutrition: a global ecology
scaling exercise. Annual Review of Ecological Systems 25: 629–660.
Smettem KRJ, Waring RH, Callow N, Wilson M, Mu Q. 2013. Satellite-
derived estimates of forest leaf area index in South-west Western
Australia are not tightly coupled to inter-annual variations in rainfall:
implications for groundwater decline in a drying climate. Global
Change Biology 19: 2401–2412. DOI:10.1111/gcb.12223.
Smethurst PJ, Almeida AC, Loos RA. 2015. Stream flow unaffected by
Eucalyptus plantation harvesting implicates water use by native forest
Ecohydrol. 9, 610–630 (2016)
630 A. C. ALMEIDA AND P. J. SANDS
streamside reserve. Journal of Hydrology Regional Studies 3: 187–198.
DOI:10.1016/j.ejrh.2014.11.002.
Soares JV, Almeida AC. 2001. Modeling the water balance and soil water
fluxes in a fast growing Eucalyptus plantation in Brazil. Journal of
Hydrology 253: 130–147. DOI:10.1016/S0022-1694(01)00477-2.
Soares P, Tomé M, Skovsgaard JP, Vanclay JK. 1995. Evaluating a growth
model for forest management using continuous forest inventory data.
Forest Ecology and Management 71: 251–265. DOI:10.1016/0378-1127
(94)06105-R.
Song X, Bryan BA, Almeida AC, Paul KI, Zhao G, Ren Y. 2013. Time-
dependent sensitivity of a process-based ecological model. Ecological
Modelling 265: 114–123. DOI:10.1016/j.ecolmodel.2013.06.013.
Stape JL, Ryan MG, Binkley D. 2004. Testing the utility of the 3-PG
model for growth of Eucalyptus grandis x urophylla with natural and
manipulated supplies of water and nutrients. Forest Ecology and
Management 193: 219–234. DOI:10.1016/j.foreco.2004.01.031.
Tickle PK, Coops NC, Hafner SD, Team TBS. 2001. Assessing forest
productivity at local scales across a native eucalypt forest using a
process model, 3PG-SPATIAL. Forest Ecology and Management 152:
275–291. DOI:10.1016/S0378-1127(00)00609-5.
Copyright © 2015 John Wiley & Sons, Ltd.
Wang QJ, Chiew FHS, McConachy FLN, James R, de Hoedt GC, Wright
WJ. 2001. Climatic Atlas of Australia: Evapotranspiration. Bureau of
Meteorology: Australia.
White DA, Beadle CL, Sands PJ, Worledge D, Honeysett JL. 1999. Quantifying
the effect of cumulative water stress on stomatal conductance of Eucalyptus
globulus and Eucalyptus nitens: a phenomenological approach. Australian
Journal of Plant Physiology 26: 17–27. DOI:10.1071/PP98023.
White DA, Crombie DS, Kinal J, Battaglia M, McGrath JF, Mendham DS,
Walker SN. 2009. Managing productivity and drought risk in Eucalyptus
globulus plantations in south-western Australia. Forest Ecology and
Management 259: 33–44. DOI:10.1016/j.foreco.2009.09.039.
Whitehead D, Beadle C. 2004. Physiological regulation of productivity
and water use in Eucalyptus: a review. Forest Ecology and
Management 193: 113–140. DOI:10.1016/j.foreco.2004.01.026.
Williams J, Prebble RE, Williams WT, Hignett CT. 1983. The influence of
texture, structure, and clay mineralogy on the soil moisture characteristic.
Australian Journal of Soil Research 21: 15–32. DOI:10.1071/SR9830015.
Winjum JK, Schroeder PE. 1997. Forest plantations of the world: their
extent, ecological attributes, and carbon storage. Agricultural and
Forest Meteorology 84: 153–167.
Ecohydrol. 9, 610–630 (2016)
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