A process-based model to evaluate

site quality for Eucalyptus nitens in

the Bio-Bio Region of Chile

Downloaded from http://forestry.oxfordjournals.org/ at U. of Florida Health Science Center Library on December 4, 2013
ROLANDO RODRÍGUEZ1*, PEDRO REAL2, MIGUEL ESPINOSA2 and
DAVID A. PERRY3

1 Corporación Nacional Forestal, Barros Arana 215, Concepción, Chile

2 Facultad
de Ciencias Forestales, Universidad de Concepcion, Casilla 160-C, Concepción, Chile
3 Department of Forest Science, Oregon State University, 321 Richardson Hall, Corvallis, OR 97331-5752, USA

*Corresponding author. E-mail: rrodrigu@conaf.cl

Summary
A simplified, physiologically based (3-PG) model was used to evaluate the effects of spatial
variation in climate and soils on the potential productivity of Eucalyptus nitens Deane and Maiden
(Maiden). The model was parameterized based on the physiology of E. nitens to predict potential
productivity and leaf area index as influenced by environmental factors. Data obtained from
conventional weather stations were utilized by the 3-PG model to predict productivity site classes.
A final plantation suitability grid was mapped to show areas of the region with productivity classes
predicted to be very high (>52 m3 ha⫺1 year⫺1), high (47.5–52 m3 ha⫺1 year⫺1), moderate (45–47.5
m3 ha⫺1 year⫺1), low (35–45 m3 ha⫺1 year⫺1) and very low (<35 m3 ha⫺1 year⫺1). The lowest
potential productivity was attributable to soil water and nutrients limitations. A process-based forest
growth model that can be widely extrapolated using geographic information system is particularly
useful to screen areas as prospective plantation sites.

Introduction three significant limitations. First, it is based on a

In a particular region, potential site productivity,
expressed as wood production, is linearly related
to leaf area index (LAI), a variable that expresses
the amount of foliar surface available for pho-
tosynthesis (Waring et al., 2005). The maximum
potential LAI on a given site in turn depends on
the structural properties of particular tree species,
climate and site resources (Waring and Running,
1998).
Potential productivity has been traditionally cal-
culated by modelling height growth for a selected
group (dominant height) of trees in the stand.
This approximation, called site index, has at least
© Institute of Chartered Foresters, 2009. All rights reserved.
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single variable that is not sufficient to represent
the stand’s real growth potential (Bernier et al.,
2003). Second, it requires the presence of trees for
measurement, which strongly limits its use in areas
where trees of a given species have not been previ-
ously established (Ford and Bassow, 1989). Third,
it gives no direct information about LAI and con-
sequently cannot be used to construct mechanistic
models (Ford and Bassow, 1989). Process-based
models simulate productivity by directly modelling
the effect of environmental factors on physiologi-
cal processes (Landsberg and Gower, 1997). The
3-PG model developed by Landsberg and Waring
(1997) is used in natural and planted forests in
Forestry, doi:10.1093/forestry/cpn045
Advance Access publication date 15 January 2009
150 FORESTRY
the US (Landsberg et al., 2000; Coops and Waring,
2001), South Africa (Dye, 2001), New Zealand
(White et al., 2000), Australia (Coops et al., 1998;
Sands and Landsberg, 2002) and Chile (Rodríguez
et al., 2002). The model incorporates basic physi-
ological characteristics of many species as well as
the genus Eucalyptus, making it possible to predict
growth and LAI after crown closure with accept-
able precision (Landsberg et al., 2003).
Additionally, the 3-PG model can be extrapo-
lated to large areas through the use of geographic
information system (GIS). Based on its reliabil-
ity in the diagnosis of environmental factors that
limit productivity, Bernier et al. (2003) recom-
mended the use of 3-PG or other process-based
models to estimate potential productivity on sites
where tree species of interest are not growing.
In this study, we used the 3-PG model together
with local information on climate and soils to
evaluate site quality for Eucalyptus nitens and to
develop potential productivity maps for the Bio-
Bio Region of Chile. Environmental factors limit-
ing productivity were also identified.
Methodology
Study area
The Bio-Bio Region comprises 36 929 km2 extend-
ing north–south from latitude 36° 00′ to 38° 30′
south and east–west from longitude 71° 00′ west
to the Pacific Ocean. The Andes Mountains to the
east and the Coastal Mountain Range to the west
define four agro-climatic areas located longitudi-
nally: the coastal dry sector, the inner dry sector,
the central valley and the Andean foothills. The
climate and soils of each agro-climatic area deter-
mine potential land uses and productive capacity
(Del Pozo and Del Canto, 1999). The National
Growth and Yield Simulation Project (Facultad
de Ciencias Forestales, Universidad de Concep-
cion and National Forest Companies) provided
information on 46 widely scattered plots as well
as physiographic information (Table 1).
3-PG
model
To assess the appropriateness of a simplified
process-based model to predict potential forest
growth over the study area, we applied a GIS
with spatial layers of climate and soils to the
3-PG model. The accuracy of model predictions
was assessed by comparison with existing plot-
based data.
As described by Landsberg and Waring (1997),
the 3-PG model:
1 Estimates gross primary production (PG)
based on the utilizable photosynthetically ac-
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tive radiation (φp.a.u) and the canopy quantum
efficiency (αC). The value of φp.a.u is obtained
by reducing the photosynthetically active
radiation (φp.a.) through non-dimensional
modifying factors whose values vary between
0 and 1. The modifying coefficients reflect
constraints imposed on the utilization of φp.a.
by vapour pressure deficit (D), drought and
stand age. Soil fertility determines allocation
of carbon to above- and below-ground com-
ponents. Drought is defined for soils of dif-
ferent textures by the ratio of water in the
root zone to the maximum available (θ). The
model also includes suboptimal temperature
and frost modifiers.
2 Estimates net primary production (PN) from PG.
The model uses the proportion PN /PG = ⊂ pp ,
which has been found to be 0.47 ± 0.04 for
diverse types of forests and geographic loca-
tions (Waring et al., 1998).
3 Estimates carbon in root biomass using two
basic relations: (1) the inverse relation be-
tween stem growth and the PN fraction allo-
cated to soil (Beets and Whitehead, 1996) and
(2) the effects of drought and nutrition on the
annual allocation of carbon to root biomass
(Santantonio, 1989; Beets and Whitehead,
1996).
4 Calculates changes in stand density over time
using a submodel derived from the self-thin-
ning law coupled with stem growth rates.
5 Apportions carbon among aboveground tree
components using allometric relations:
wˆ i = aiwˆ N i ,
where ŵ is the tree’s total biomass and i is the
tree component. The parameter Ni reflects the
species’ genetic characteristics.
6 The growth rate of trees decline with age. To
this respect, the hydraulic limitation theory
proposes that the decline of forest productiv-
 PROCESS-BASED MODEL TO EVALUATE SITE QUALITY 151

Table 1: Information on the locations of plots, physiographic setting, rainfall, available soil water-holding
capacity and MAI at 10 years for 10-year-old Eucalyptus nitens in the Bio-Bio Region

Available soil MAI at

Physiographic Rainfall water-holding 10 years
Name of plots setting Soil fertility (mm year⫺1) capacity (mm) (m3 ha⫺1)
Elías Andean foothills Poor 1390 154 44.62
Laurel Andean foothills Poor 1815 102 52.90
Los Angeles (1) Inner dry sector Very poor 1093 190 22.60

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Los Angeles (2) Inner dry sector Very poor 1093 135 44.97
Las Nieves Andean foothills Rich 1354 173 45.29
Santa Rosa Central valley Poor 1815 118 43.39
Santa Catalina Andean foothills Very poor 1390 227 36.45
Santa Catalina (2) Andean foothills Poor 1390 227 47.86
Santa Catalina (3) Andean foothills Poor 1390 237 50.07
Santa Catalina (4) Andean foothills Very poor 1390 231 35.67
Bellavista Central valley Poor 1390 226 45.51
El Carmen Inner dry sector Very poor 1134 175 24.29
El Castaño Andean foothills Rich 1354 201 57.10
El Huachi Andean foothills Poor 1815 170 45.83
El Huachi (2) Andean foothills Poor 1815 155 45.32
El Huachi (3) Andean foothills Poor 1815 154 46.23
El Huachi (4) Andean foothills Poor 1815 155 48.70
El Mirador Inner dry sector Very poor 1390 227 39.43
El Prado Andean foothills Poor 1505 224 45.01
El Prado (2) Andean foothills Poor 1505 225 40.82
El Prado (3) Andean foothills Very poor 1505 224 41.86
El Prado (4) Andean foothills Very poor 1505 225 41.73
El retiro Inner dry sector Very poor 1390 224 40.97
El retiro (2) Andean foothills Rich 1390 223 59.58
Estrella Andean foothills Very poor 1815 155 46.59
Huallenco Andean foothills Poor 1390 400 52.95
Huallenco (2) Andean foothills Poor 1390 226 50.73
Huallenco (3) Andean foothills Poor 1390 226 47.50
La Tranquera Inner dry sector Very poor 1390 223 44.40
La Acacias Andean foothills Poor 1390 226 47.15
La Acacias (2) Andean foothills Poor 1390 226 47.50
La Acacias (3) Andean foothills Very poor 1390 226 36.26
La Acacias (4) Andean foothills Poor 1390 226 46.41
Los Coihues Central valley Poor 1437 250 53.76
Monteverde Coastal dry sector Rich 1390 475 60.62
Pisagua Inner dry sector Very poor 1815 177 12.43
San José Central valley Poor 1390 145 49.01
San José (2) Central valley Poor 1390 160 44.30
San José (3) Central valley Poor 1390 150 47.70
San Ramón Coastal dry sector Poor 1390 293 55.80
San Luis Andean foothills Very poor 1354 233 39.28
San Luis (2) Andean foothills Poor 1354 325 51.73
Santa Sofía Coastal dry sector Poor 1390 345 54.44
Santa Teresa Andean foothills Poor 1390 340 51.51
Santa Cristina Andean foothills Poor 1390 397 54.94
Santa Cristina (2) Andean foothills Poor 1390 400 49.98

Data were provided by the Instituto de Investigaciones de Recursos Naturales (1964) and Instituto de
Investigaciones Agropecuarias (Del Pozo and Del Canto, 1999). Data of MAI were provided by National
Growth and Yield Simulation Project.
152 FORESTRY

ity with age is a consequence of the loss of

Landsberg (1986); Waring (2000)

MJ⫺1; φp.a.u, photosynthetically active solar radiation utilized, MJ m⫺2 month⫺1; Top, optimum temperature for photosynthesis, °C; Tmi, minimum temperature
These differ from the original model (Landsberg and Waring, 1997), following Sands (2000) for 10-year-old Eucalyptus nitens. αC, canopy quantum efficiency,
whole-plant and leaf-specific hydraulic con-

for growth, °C; Tma, maximum temperature for growth, °C; L, LAI, m2 m⫺2; dia, average stem diameter (mm); m, soil fertility rank (m is 1 fertilized soil and
Landsberg and Waring (1997)
Sands and Landsberg (2002)
ductance with tree height caused by increased
friction. Based on this theory, the 3-PG model

decrease to 0, 1 in non-fertilized soils). To obtain maximum potential MAI, nutritional status was not limiting photosynthetic production and m is 1.
References
estimates the effect of age on tree growth

Battaglia et al. (1998)

Tickle et al. (2001)

Tickle et al. (2001)

Tickle et al. (2001)

using the linear relation between hydraulic
conductivity and PN, in which stem conduc-

Sands (2000)

Sands (2000)
Sands (2000)
Sands (2000)
tance declines with age and thereby induces a
lower stomatal conductance (gc) (Mencuccini

Downloaded from http://forestry.oxfordjournals.org/ at U. of Florida Health Science Center Library on December 4, 2013
and Grace, 1996). Table 2 shows the model
functions and parameters used in our study.

Predicting potential productivity

Battaglia et al. (1998) developed a simple ana-

φp.a.u
lytic model for the relation between LAI and dry
matter production in E. nitens. In their model,

MJ⫺1
‘increasing LAI increased light interception and

Values of functions and parameters

Top was set at 20°C, Tmi 2°C and Tma 32°C

ηr = (0.8 × 0.3)/(0.23 + (0.8 − 0.23)mφp.a.u

hence dry matter production, but simultaneously

Foliage mass, kg = 0.00005 × dia (mm2.65)

Maximum αC ranges from 1.8 to 4.2 g C

Stem mass, kg = 0.00007 × dia (mm2.65)

increased canopy respiration. Consequently, for
a given light utilization coefficient, there was
a value of LAI that maximized NPP’ (Battaglia
et al., 1998). For this reason, for a given value
of mean annual temperature (T) and water stress

1 − (1.38 exp(−0.5 × L))

index (W), optimal LAI could be estimated using
the equation of Battaglia et al. (1998):

2% month⫺1
LAI = 1 / k ln[(sεΩk)/ro N F +γ], (1)

0.005 m s⫺1
4.0 m2 kg⫺1

500 kg m⫺3

0.02 m s⫺1
where LAI is the canopy leaf area index (m2 m⫺2),
Table 2: Functional forms and parameters used in this study

k is the canopy light extinction coefficient, s is the

specific leaf area (m2 kg⫺1 DM), ε is the light utili-
zation efficiency (kg C MJ⫺1), Ω is the annual inci-
dent radiation (MJ m⫺2 year⫺1), ro is maintenance
respiration rate per unit canopy N content (kg C
Fraction of radiation absorbed by the canopy
Light conversion efficiency of photosynthesis

kg⫺1 N year⫺1), NF is the average leaf N concen-

tration (kg N kg⫺1 DM) and γ is rate of carbon
Constraints on light conversion efficiency

Fraction of production allocated to roots

Maximum canopy stomatal conductance

loss as litterfall (kg C kg⫺1 DM year⫺1). We used

Maximum leaf stomatal conductance
Allocation equation for foliage mass

the parameter values of Battaglia et al. (1998) with

Allocation equation for stem mass

the exception of Ω, which was measured in the

associated with temperature

Bio-Bio Region, and ε which, as described below,

we calculated on a site-specific basis.
Light utilization efficiency (ε), therefore the op-
timal LAI, varies with site conditions, particularly
Foliage turn-over

water availability and temperature (Battaglia et al.,

Specific leaf area

1998). To estimate how environmental conditions

Wood density

affect the distribution of optimal LAI through-

Variable

out the Bio-Bio Region, we used Landsberg’s

(1986) equation relating ε to T and W.
 PROCESS-BASED MODEL TO EVALUATE SITE QUALITY
ε = 0.00195fW fT , (2)
where fT = max{0,(1 ⫺ (T ⫺ 13.2/9.2)2)} and fW =
max[0,(1+0.9 ln W)].
In this modelled relationship, 13.2°C is a value
that maximizes net canopy production and 9.9
is the value of temperature stress. W was cal-
culated as the annual mean of the daily ratio of
actual to potential evaporation (Battaglia et al.,
1998). Actual evapotranspiration was calculated
as the product of potential evaporation and a fac-
tor crop as estimated by Worledge et al. (1998).
Potential evaporation was estimated from Mean
Class A pan evaporation distributed in the Bio-
Bio Region (Del Pozo and Del Canto, 1999).
In the 3-PG model structure, the maximum
rates of water vapour and carbon dioxide ex-
change through leaf stomata approach a con-
stant by the time a forest canopy reaches a LAI
of 3.0 (Kelliher et al., 1995) and that at LAI >3.0,
maximum photosynthesis is a linear function
of ε (Wang et al., 1991). The assumptions reduce
the requirement for accurately assessing LAI
and simplify the calculation of photosynthesis
(Waring, 2000).
We examined the sensitivity of ε to the param-
eters of 3-PG and its temperature dependence,
assuming that water and nutritional status were
not limiting photosynthetic production and then
we ran the 3-PG model until reaching an opti-
mum value of LAI (between 5.8 and 6.2) to ob-
tain maximum potential mean annual increment
(MAI) because LAI is a critical integrator of
water availability and productivity (Gholz et al.,
1990).
Note that the LAI values estimated by the PN-
maximizing model should not be interpreted as
those necessarily present in the plantations (Ludwig
et al., 1965) because trees are under competition
and there has been natural selection. Another im-
portant facet is that there is a linear relationship
between canopy respiration and foliage mass.
In this sense, LAI represents the target for trees
growing under particular conditions and the tar-
get is provisional on the assumption of the model
(Battaglia et al., 1998).
Results produced by the 3-PG model arise from
interactions among carbon, water and nutrients.
Separating the effect of water from that of nutrients
can yield more detailed insights into resource limita-
153
tions. Because the 3-PG model contains a water bal-
ance subroutine, the effects of soil water storage on
growth can be identified. We bounded the effects of
limiting water by predicting MAI when soil water
was at the permanent wilting point. Similarly, we
bounded the effects of limiting nutrients by predict-
ing MAI with a fertility rating (FR) of m = 0.1.
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Analysis of plot information and validation of
the 3-PG model
Comparison between predicted and observed MAI
in 46 permanent, 1000 m2 research forest plots
(Table 1), using the 3-PG model with parameter
values derived from calibration (Table 2) against
research plots, was used to validate the 3-PG model
in the Bio-Bio Region for E. nitens. The plots were
part of the National Growth and Yield Simula-
tion Project and were scattered over the main area
of E. nitens plantations in the Bio-Bio Region.
Plots were hand planted at 1250 trees ha⫺1 and
established following intensive site preparation
that consisted of ripping and mound ploughing
to improve soil rooting volume. Fertilization was
applied in all plots: 120 g per seedling, as a com-
mercial fertilizer, consisting of N (16 per cent), P
(12 per cent), K (8 per cent), S (4 per cent), B (4
per cent) and Mg (1.8 per cent). Following plant-
ing, herbaceous and weed control was necessary
to provide a rooting zone free of broadleaf weeds
and grasses. The plots are measured annually and
have between 6 and 12 measurements, at ages that
vary between 7 and 17 years old. Data collected
on each plot include age, diameter at breast height,
height, volume and density together with climatic
and soil variables that describe site conditions.
The accuracy of the 3-PG model was evalu-
ated with the root of the root mean square error
(RMSE), in measurement units and percentages.
To evaluate bias, the aggregated difference (AD),
in measurement units and percentages, was used
(Cao et al., 1980).
RMSE = ∑ (O − E )
i i
2
/ n,
AD = ∑ (Oi − Ei ) / n,
where Oi is an observed value and Ei the corre-
sponding 3-PG estimated value.
154 FORESTRY
The GIS
Isoproductivity zones were delineated using GIS
software (Chang, 2004). The following databases
were used.
Spatial soil model
The Earth Ordering System (scale 1 : 250 000) de-
veloped by Schlatter (1994) and soil origin data
provided by the Instituto de Investigaciones de Re-
cursos Naturales (1964) were used to construct the
soils’ layer. Estimates of the soil physical properties,
such as bulk density, porosity and field capacity
and permanent wilting point, were obtained from
the Instituto de Investigaciones Agropecuarias (Del
Pozo and Del Canto, 1999). The maximum avail-
able soil water was estimated from retention curves
provided by Carrasco et al. (1993). Data of the soil
chemical characteristics (pH, organic matter, Ca,
Mg, Na, P, SO4, Fe, Mn, Cu, Zn and B) were ob-
tained from the Instituto de Investigaciones Agro-
pecuarias (Del Pozo and Del Canto, 1999). 3-PG
requires a FR (0–1), describing the soil fertility in
the rooting zones. We used qualitative estimates of
soil fertility for physiographic characteristics and
distinguished very poor sites (m = 0.3), poor sites
(0.5) and rich sites (0.7) (Table 1) in the model
validation for MAI prediction. For potential MAI
prediction, a value of m = 1 was used.
Spatial climate model
The spatial distribution of monthly maximum,
mean and minimum temperatures was obtained
from Santibáñez and Uribe (1993). Estimates of
monthly solar radiation values were derived from
maximum and minimum data at direct measure-
ments performed by the Instituto de Investiga-
ciones Agropecuarias (Del Pozo and Del Canto,
1999) using the procedure of Bristow and Campbell
(1984). Maximum potential solar radiation on a
flat surface was obtained by correcting from lati-
tude and elevation (Running and Nemani, 1987).
This potential value was corrected by the monthly
variation of the angle of solar declination over
the earth’s surface (Waring, 2000).
Digital elevation model
The slope and exposure models were derived
from a digital terrain elevation model generated
with information provided by regular Chilean
cartography (Instituto Geográfico Militar, 2001).
Slope and exposure classes were derived using a
triangle irregular network, which is an elevational
data storage system (Crosier et al., 2004).
Map Construction
Each digital coverage layer was created, cleaned
and processed using the Arc Info Work-Station
System version 7.1 running in a UNIX platform.
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GIS results were exported to the 3-PG model.
Finally, the 3-PG estimations and the basic lay-
ers were converted to shape format and manipu-
lated in Arc View 3.2, where the cartography was
built. The resulting maximum potential MAI val-
ues were grouped using the legend tool to define
potential productivity zones. The classification
method used is the Natural Breaks default classifi-
cation method in Arc View. This method identifies
breakpoints between classes using a statistical for-
mula (Jenk’s optimization). Basically, the Jenk’s
method minimizes the sum of the variance within
each of the classes to find groupings and patterns
inherent in the data (Crosier et al., 2004).
We used the program 3PGpjs (Sands, 2000) to
predict potential productivity with a 3-PG inter-
face from the Commonwealth Scientific and In-
dustrial Research Organisation (CSIRO) Forestry
and Forest Products & Cooperative Research
Centre for Temperate Hardwood Forestry (CRC)
for Sustainable Production Forestry. The calibra-
tion of these models for diverse Eucalyptus species
has been described by Landsberg et al. (2003). In
this study, functions and parameters specific to
E. nitens were taken from Morris (1999) and
other published studies (Table 2).
Results
Effects of the annual mean temperature on the
LAI
Figure 1 shows the predicted effect of mean an-
nual temperature on LAI of E. nitens plantations
in the Bio-Bio Region, using Ω equal to 6.5 GJ
m⫺2 measured directly in meteorological stations
within the region and assuming no water stress.
From a low value of 5.7 m2 m⫺2 at 10.9°C, LAI
increases sharply up to 11.5°C, then plateaus at
LAI = 6.1 m2 m⫺2 between temperatures of ~12.2
and 13.2°C (Figure 1). This maximum LAI is
 PROCESS-BASED MODEL TO EVALUATE SITE QUALITY 155

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Figure 1. Effects of mean annual temperature on LAI of Eucalyptus nitens plantations in the Bio-Bio Region.
Symbol ○ is LAI as a function of mean annual temperature; the solid line is not a regression but is used to
emphasize trends.

Figure 2. A water stress index, expressed as the ratio of available soil water to the maximum supply, shows
a general increase in LAI above a threshold of 0.6 as the index approaches 1.0 (i.e. no limitations). Symbol ○
is LAI as a function of water stress index; the solid line is not a regression but is used to emphasize trends.
156
similar to that reported by Pinkard et al. (1999)
for E. nitens in Australia.
Figure 2 shows the effects of water stress on
LAI. There is a sharp increase in LAI between a
water stress indices of 0.6 and 0.65, after which
the effect of increasing water availability (higher
water stress index) is approximately curvilinear.
Model validation for MAI prediction
MAI values observed in the growth plots varied
between 23 and 50 m3 ha⫺1 at age 10 and were
highly correlated with MAI simulated by the 3-PG
model (Figure 3). Additionally, a low bias was re-
corded with AD of 0.26 in absolute terms and 0.85
per cent in percentage terms. The model also had
acceptable precision; RMSE had an absolute value
of 4.55 and a percentage value of 14.6 per cent.
Potential MAI in the Bio-Bio Region
Three main physiographic landscapes dominate the
Bio-Bio Region of Chile. The coastal range along to
the Pacific Ocean and the Andes Cordillera enclose
the Central Valley from north to south and create
Figure 3. MAI simulated by the 3-PG model vs MAI observed in growth plots (m3 ha⫺1 year⫺1) for Eucalyptus
nitens in the Bio-Bio Region. Age in the plots varies between 7 and 17 years old. Symbols ○ are values of observed
MAI. Regression line (y = 0.3975 + 0.9954 × R2 = 0.997) is the predicted MAI as a function of observed MAI.
FORESTRY
a great variety of climates, soils and forest sites
(Toro and Gessel, 1999). Predicted MAI classes
for E. nitens plantations ranged from 17 to 62 m3
ha⫺1 year⫺1 in the Bio-Bio Region. Figure 4 shows
variation in potential productivity mapped across
the Bio-Bio Region. In general, predicted MAI in-
creased in mountainous terrain in the foothills of
the Andes following gradients in precipitation, an-
nual temperature and soil fertility (Table 2; Figure
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4). Not surprisingly, highest predicted MAI were
in areas with highest precipitation, coolest annual
temperatures and the most fertile soils. Elevation
by itself was not a good predictor of MAI, as highly
productive sites occurred at both low and high el-
evations. Soil nitrogen concentrations on the most
productive sites varied widely, but on average, soils
were considerably more fertile and held more water
than sites with lower productivity (Table 3).
Discussion
Combined with spatial information derived from
GIS, the 3-PG model can generate valuable informa-
tion for the development of operational silviculture
 PROCESS-BASED MODEL TO EVALUATE SITE QUALITY 157

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Figure 4. Potential productivity classes of MAI (m3 ha⫺1 year⫺1) for the Bio-Bio Region, as estimated by
the 3-PG model. (The predicted values of MAI in each site are not possible to reach under the operational
silviculture.)

(Mummery and Battaglia, 2001; Almeida et al.,
2004; Dye et al., 2004). The AD values (0.85 per
cent) and RMSE values (14.6 per cent) obtained
for MAI confirm that process-based models have
precision levels similar to traditional inventories.
Our work corroborates the suggestion of Tickle
et al. (2001) that process-based models are a
valuable tool for predicting growth at a regional
level. Additionally, process-based models provide
insights into the role of environmental variables
in determining MAI that are very difficult to ob-
tain with traditional inventories (Waring, 2000;
Almeida et al., 2004).
To better understand the factors that explain
potential productivity of E. nitens plantations in
the Bio-Bio Region, we used soil and climate in-
formation from Del Pozo and Del Canto (1999)
and Carrasco et al. (1993). In addition, typical
values of observed environmental variables are
presented for very high, moderate and very low
MAI classes (Table 3).
Highest productivity sites occur on inceptisols
and on those alfisols that occur in areas with rela-
tively high precipitation and regularly subjected
to freezing temperatures. In the Bio-Bio Region,
inceptisols are derived from volcanic ash and have
high water-holding capacity, good drainage and
high nitrogen concentrations. They also occur
in areas of relatively high precipitation. Alfisols
are marine terrace soils with good drainage. In
addition to high precipitation, those alfisols with
high productivity also have high water-holding
capacity, with relatively high average soil nitro-
gen concentrations (Table 3). High productivity
158 FORESTRY

12.2–14.1
12.5–13.9
13.3–15.6
13.1–14.7
areas span a wide range of elevation, mean an-

T (°C)
nual temperature and annual incident radiation.

mean annual temperature. Data were provided by the Instituto de Investigaciones de Recursos Naturales (1964) and the Instituto de Investigaciones
layer; N is soil nitrogen; WHC is available soil water-holding capacity; Ω is the annual incident radiation; P is the annual precipitation and T is the
MAI is the potential mean annual increment as estimated by 3-PG model; texture: Fr-arc is clay-like layer, Fr-lim is the lime layer and Are is sandy
Consequently, the productivity of E. nitens in
the Bio-Bio Region does not respond directly to
any of these three factors. However, the factors
considered here may affect productivity through

1250–2000
1500–2550
640–1100
750–1250
year⫺1)
P (mm
several distinct mechanisms, factors can inter-
act or factors are related with seasonal variation
(Battaglia and Sands, 1997). Because water is

Downloaded from http://forestry.oxfordjournals.org/ at U. of Florida Health Science Center Library on December 4, 2013
Table 3: Soil and climate variables related to the potential productivity of 10-year-old Eucalyptus nitens in the Bio-Bio Region

considered to be the main factor limiting produc-

tivity, productivity is consistently related to the
Ω (MJ m⫺2

capacity of the soil to store water (Gholz et al.,

year⫺1)
5.5
6.2
5.8
6.6
1990; Dye et al., 2004); this is the case with the
inceptisols derived from volcanic ash and alfisols
derived from marine terraces that have high wa-
ter-holding capacity and can explain the highest
WHC (mm)
410–700
280–570
160–340
52–155

productivity in these sites (Table 3).

The relatively small change in MAI over a large el-
evational span is consistent with the ecophysiology
of E. nitens. Optimal temperatures for photosyn-
thesis span a wide range (14–20°C) for the species
(Battaglia et al., 1996) – commonly occurring tem-
0.05–0.25
0.18–0.35
0.04–0.09
0.01–0.07
N (%)

peratures in the study area. Our results support the

hypothesis of Battaglia et al. (1996) that the broad
photosynthetic response of E. nitens enables it to
acclimate to a wider range of environments than
Fr-arc-Are

many other Eucalyptus species. Eucalyptus nitens

Texture

Fr-lim
Fr-arc

in the Bio-Bio Region occurs in elevations from 0

Are

to 830 m a.s.l., with temperatures varying between

8 and 32°C in the summer and ⫺3.0 to 13°C in the
winter; the higher elevations experience frequent
Altitude (m)

winter frosts. Similar to the results of Battaglia

0–160
300–830
80–130
130–300

et al. (1998) for E. nitens in Australia, frost has

little or no effect on this species’ MAI in Chile.
Moderate levels of MAI were predicted for the
Agropecuarias (Del Pozo and Del Canto, 1999).

Coastal Range. The soils of Coastal Range are

derived from granitic and metamorphic rocks
Soil order

Inceptisol

Entisol

that compose the so-called ‘Batholitic Coastal’

Alfisol

Alfisol

and have been seriously impacted by erosion.

MAI values for the Coastal Range were lowest
in the north and highest in the south following a
gradient of precipitation and plant-available soil
ha⫺1 year⫺1)

47.6–52
MAI (m3

water-holding capacity (Figure 3). Lowest growth

>55.0
>55.0

>35.0

rates occurred in Central Valley at middle eleva-

tions with decreasing precipitation, high evapo-
transpiration rates and a long drought period (~6
months). The sandy soils of the Central Valley
MAI classes

also have low inherent fertility and very low wa-

Very high
Very high
Moderate
Very low

ter-holding capacity (Tables 1 and 3).

The very low productivities occurred in sandy
soils derived from metamorphic and granitic
 PROCESS-BASED MODEL TO EVALUATE SITE QUALITY 159

rocks. These soils have low water-retention ca-
pacity. The very low productivity sites were in
Central Valley on entisols; these are young soils
originating in the Bio-Bio area from various rock
types (e.g. granitic, metamorphic, andesitic and
basaltic). They are sandy soils with excessive
drainage, low soil water-holding capacity (<52
m3 ha⫺1 of available water) and low nitrogen con-
centrations. Compounding the low water-holding
capacity of soils, these areas also have low an-
nual precipitation (750–1000 mm) and high solar
radiation potentials (>6.5 MJ m2 year⫺1), which
would increase evaporative demand (Table 3).
These factors suggest that productivity in these

Downloaded from http://forestry.oxfordjournals.org/ at U. of Florida Health Science Center Library on December 4, 2013

Figure 5. Effect of soil nutrient and water availability in the soil on the MAI (m3 ha⫺1 year⫺1) across the
growth plots for Eucalyptus nitens in the Bio-Bio Region. The upper line shows the MAI without soil and
water limitations (simulated by the 3-PG model), the second line indicates the observed MAI, the third line
and the bottom line indicate the MAI under nutrients and water limitations, respectively (simulated by the
3-PG model).
160 FORESTRY
areas is strongly controlled by water-holding ca-
pacity and perhaps nitrogen limitations. Battaglia
et al. (1998) found that LAI of E. nitens plan-
tations in Australia increased sharply as annual
mean temperature increased from 7 to 11°C and
reached a plateau between 13 and 14°C (the
highest temperature measured for E. nitens). We
found a similar pattern; however, the LAI of E.
nitens plantations in the Bio-Bio Region plateaus
at a lower temperature than those in Australia.
However, although mean annual temperature
had a strong influence of LAI in our study, pro-
ductivity was influenced more strongly by the
constraints on leaf efficiency exerted by water
availability and soil fertility.
Figure 5 shows that the water availability is the
major factor influencing the productivity in the
Bio-Bio Region. This finding is consistent with
the hypothesis of Ollinger et al. (1998) that NPP
is strongly correlated with precipitation and sug-
gests that water is an important factor controlling
regional pattern of productivity in forest types, as
well as in E. nitens, which was demonstrated in our
study. Additionally, E. nitens was very sensitive
to water soil availability (Figure 4) in areas with
low water-holding capacity of soils. These findings
are consistent with the hypothesis of Beadle et al.
(1995) and White et al. (1996) that water stress re-
duces dramatically the growth rate of E. nitens due
to the decrease of the osmotic potential and bulk
elastic modulus in response to water stress.
However, understanding how productivity
varies across complex environmental gradients
requires modelling the combined influences of ra-
diation, temperature, humidity deficits, drought
and soil fertility as they change spatially and tem-
porally.
Conclusions
A process-based forest growth model shows po-
tential to serve as a useful tool to screen areas
as prospective plantation sites. In this study, we
combined the 3-PG model with a GIS to assess
the potential productivity of E. nitens in Chile’s
Bio-Bio Region.
Our results indicate that annual precipitation,
available soil water-holding capacity and low lev-
els of soil nitrogen are the principal factors influ-
encing E. nitens productivity. Our analysis was
able to separate the effects of limiting water from
those of nutrients, and it is reasonable to conclude
that soil water availability is an important fac-
tor controlling regional patterns of productivity.
Highest productivity sites occur in areas with rel-
atively high precipitation and regularly subjected
to freezing temperatures. In our study, E. nitens
reduced growth dramatically in areas with low
water-holding capacity of soils. Additionally, we
Downloaded from http://forestry.oxfordjournals.org/ at U. of Florida Health Science Center Library on December 4, 2013
found that large changes in elevation have only a
small influence of the productivity of E. nitens in
the Bio-Bio Region.
Acknowledgements
We thank Professor Richard H. Waring for his helpful
comments and suggestions to the manuscript. We also
thank National Growth and Yield Simulation Project
(Facultad de Ciencias Forestales, Universidad de Con-
cepcion and National Forest Companies) for providing
sites data. The authors are also grateful to two anony-
mous reviewers and the editor for valuable comments
on the manuscript.
Conflict of Interest Statement
None declared.
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