Egypt. J. Bot., Vol.

52 (2), 199-212 (2012)

Growth Performance of Four Irrigated Plantations in Egypt

Emad Farahat1,* and Hans W. Linderholm2
1Botany and Microbiology Department, Faculty of Science, Helwan University, Cairo, Egypt
2 Regional Climate Group, Department of Earth Sciences, University of Gothenburg, Gothenburg, Sweden

HE objective of this study was to compare and evaluate the spatial variation in crown architecture
traits, related growth and foliar traits and their relation to height (H) for five tree species

(Eucalyptus camaldulensis Dehnh., Khaya senegalensis (Desr.) A. Juss., Dalbergia sissoo (Roxb.),
Cupressus sempervirens L. and Casuarina spp.), irrigated with wastewater and/or ground water in four
desert planted forests in northern Egypt. The studied traits were significantly different for each tree
species among sites. The positive correlation between H and crown architecture traits increased with
decreasing tree heights, and strongly correlated with crown length in all species. Crown diameter and
area, and Leaf mass per area (LMA) appear to be potential indicators for the plastic changes in growth
performance of the species and its adaptability to site conditions. Stem diameter (dbh)-H relationship
was nonlinear and highly significant for all tree species except Cupressus sempervirens and their best
correlations were obtained at small tree heights. Soil sand content was the main operating edaphic factor
and accounted for 40.8% of the species traits-environment relationship among the sites. The growth
performance of trees was not correlated only with tree age or water availability but also with other site
conditions and silviculture practices.
Keywords: Crown architecture, Size traits, LMA, Management, Planted forests, Egypt

Tree architecture is an important determinant of height extension, light capture, and mechanical stability of
trees, and it allows species to exploit the vertical height gradient in the forest canopy as well as the horizontal
light gradients at the forest floor (Poorter et al. 2006). The concept of plant architecture suggests that each plant
species has its own growth form (Bell 1984). Westoby and Wright (2006) stated that addressing the questions
of how and why plant traits and architectures vary among species and sites is the most important single step
towards understanding land-ecosystem properties in general. Inter-specific variation in height growth rates is
related to that in crown architectural traits (Takahashi & Rustandi 2006). Changes in crown morphology have
been shown to play an important role in the acclimatization capacity of species to different light environments,
and crown-morphological plasticity was found to be important to shade-tolerant trees (Canham 1988, 1989,
Beaudet & Messier 1998). Crown-architecture traits (crown area, diameter, length and shape) are important
factors affecting the light environment in forest canopies. The plasticity in crown traits under different stress

Corresponding author, Email: emad23_1999@yahoo.com

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EMAD FARAHAT and HANS W. LINDERHOLM

and light conditions has been reported by many investigators of forest trees (e.g. Takahashi et al. 2005, Chmura
et al. 2007, 2009).
In general, the plasticity strategies for plants which are to successfully endure in arid regions have
been found to be associated with two main adaptive traits; minimizing water loss and maximizing water uptake.
Foliar traits, such as leaf mass per area (LMA ) and leaf area, are indicators for this strategy, either by
increasing LMA or reducing light absorbance through rolled leaves (Pena-Rojas et al. 2005, Stefanos et al.
2008). Higher LMA is a main adaptation to droughts, and is positively related to the photosynthetic tissue per
unit area and investment in structural tissues, allowing higher tolerance to unfavorable conditions (Marcoco et
al. 2000, Chen & Wang 2009). Differences in leaf morphology and physiology and changes in crown
architecture traits and LMA among tree species are common. These traits changes dramatically with growth
stage, tree height and/or susceptibility to drought stress and they are size-dependent in nature (Koch et al. 2004,
Poorter et al. 2006, Ryan et al. 2006, He et al. 2008, Tanaka-Oda et al. 2010). Leaf trait responses to
environmental conditions often differ among species (Bond 2000, Ryan et al. 2006) and can be used as
indicators of whole-organism physiology, such as drought tolerance and growth performance, under stressful
circumstances.
Crown architecture affects tree growth through control of leaf area and its display for effective light
capture and photosynthesis. So, it may important to quantify crown traits for effective use of intensive
silvicultural practices to improve tree growth in forest plantations (Chmura et al. 2007). It is also very relevant
in studies of the growth of stands due to the close correlation between crown size and stem diameter, and
density of trees in a stand (Hemery et al. 2005). Knowledge of the crown traits-tree size relationship of a
species can be applied to indicate the tolerance of a species to variations in stand density (Dawkins 1963), for
predicting basal areas, devising thinning regimes and, given the inclusion of height data, for developing stand
volume estimates. Our understanding to these relationships will assist in decision which tree can be selected in
a certain site exposed to a range of site conditions (e.g. wind speed) according to the crown parameters and to
inform the design of forest management prescriptions (Hemery et al. 2005, Thorpe et al. 2010).
The impacts of different silviculture practices e.g. thinning, repeated fertilization, control of competing
vegetation, plant density and irrigation method, on the growth and crown traits of the plants were studied by
many investigators (e.g. Chmura et al. 2007, Waterworth et al. 2007, Thorpe et al. 2010, Yanga et al. 2011) .
For instance, Stem growth of Pinus radiata (10-29 years old) responded markedly to irrigation, with a strong
interactive effect when combined with N fertilization. Moreover, irrigation increased growth in the larger trees

GROWTH PERFORMANCE OF FOUR

201

proportionally more than the smaller trees, resulting in an increased diameter distribution compared with the
non-irrigated stands. Cessation of irrigation in combination with thinning after 5 years of treatment immediately
reduced annual basal area growth of Pinus radiata to levels similar to the control for at least 2 years
(Waterworth et al. 2007).
The establishment of planted forests in the world is expected to help in preventing desertification of
agricultural land, as well as providing increased wood supplies and reducing negative impacts of environmental
pollutions (FAO 2006). In Egypt, the establishment of planted forests is an ongoing annual activity by the
Ministry of State for Environmental Affairs to cope with an increase in wastewater production burden (about
2.4 billion m3year-1) (Egypt State of the Environment Report 2009). Considering the scale of recent
establishment of the planted forests, there is not enough literature on growth performance, biomass production,
economic products, efficiency of current silvicultural practices, wood diseases, shoot dieback and tree death.
The objective of this study was to compare and evaluate the spatial variation of tree crown architecture
traits, related growth and foliar traits and their relation to height (H) for five tree species (Eucalyptus
camaldulensis Dehnh., Khaya senegalensis (Desr.) A. Juss., Dalbergia sissoo (Roxb.), Cupressus sempervirens
L. and Casuarina spp.) growing in four desert planted forests in Egypt named: Sadat, Wadi El-Natrun,
Sarapium and El-Tur forest. We tried to answer the following questions: 1. Is there a spatial variation in growth
performances of the studied tree species in desert planted forests? 2. Are the variations in architectural traits
related to tree height? 3. What is the relation between height and diameter among species and sites? 4. Which
environmental variables have most influence on species performance?. This study will provide more
understanding for the performance of tree species in their habitats and help in improving species selection
techniques for successful planting in arid areas.
Methods
Study sites and tree species
This study was carried out in four desert planted forests (plantations) in Egypt. Two forests are located
in the Western Desert, and are commonly known as Sadat Forest (officially: Chinese-Egyptian Friendship
Young People Forest) and Wadi El-Natrun Forest (officially: Japanese Planted Forest). The third forest is
located in the Eastern Desert and is commonly known as Sarapium Forest. The fourth forest is El-Tur
which is located outside El-Tur City, South Sinai Governorate (Fig. 1). The trees planted in the forests are
irrigated with lightly treated sewage wastewater, except in Wadi El-Natrun, which is irrigated with

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EMAD FARAHAT and HANS W. LINDERHOLM

Fig. 1. Location map showing the distribution of studied four planted forests in Egypt

groundwater. The quality of wastewater varied among the forests depending on the efficiency of treatment
method. The forested area ranges from 420 ha (Sarapium) to 63 ha (El-Tur forest). More details about irrigation
systems and their efficiency, and climate characteristics in the different study sites are shown in Table 1.
Silviculture practices include fertilization, tillage; control of competing understorey vegetation and pest control
are the most common practices in Sarapium forest only compared with other forests. In addition, not all of the
sampled tree species are cultivated in all sites, and they vary in age and density within and among forests
(Table 2). More canopy gaps were observed in El-Tur and Wadi El-Natrun forests for most tree species
compared with the other forest sites. The sampled tree species were: Eucalyptus camaldulensis Dehnh. (River
Red gum), Khaya senegalensis (Desr.) A. Juss. (Dry zone Mahogny), Dalbergia sissoo (Roxb.) (Sissoo),
Cupressus sempervirens (Italian cypress) and Casuarina spp. (Australian pine). All trees are cultivated as
single overstory species. The Casuarina spp. consisted of mixed populations, represented by 3 species (C.

GROWTH PERFORMANCE OF FOUR

203

Table 1. General characteristics of the studied planted forest in Egypt

Parameter/Forest

Sadat

Sarapium

Area (ha)

210

420

Wadi
El-Tur
El-Natrun
63
84

Irrigation system

Drip

Drip

Drip

Flooding

Type of water
Efficiency of
irrigation system

Sewage

Sewage

Ground

Sewage

*Poor

*Good

Poor

*Excellent

Air temperature, C
(Min.-Max.)

**15-27.2 13.9-28.7 15-27.2

Relative humidity %
63-72
(Min.-Max.)

52-66

63-72

17.2-29.7
46-59

* Poor = the irrigation network is frequently blocked by water suspended solids and the trees are irrigated
irregularly, Good = the irrigation network is maintained and the trees are irrigated regularly in
considerable amounts, Excellent = large pipes are used for distributing the water of irrigation and trees are
flooded by water during irrigation
** The distance between Sadat and Wadi El-Natrun forest locations is ca. 38.3 km

Table 2. Age (yr.) and density (trees ha-1) of the sampled tree species in 2009 in the studied planted forests in
Egypt
Tree species/Forest

Sadat

Sarapium

Wadi El-Natrun

El-Tur

Age

Density

Age

Density

Age

Density

Age

Density

Eucalyptus camaldulensis Dehnh

1114

1005

11

900

13

750

Casuarina spp.

10

1213

1100

10

1100

13

850

Cupressus sempervirens L.

10

1213

825

10

857

Khaya senegalensis(Desr.) A. Juss.

10

1020

1089

Dalbergia sissoo(Roxb.)

10

716

957

glauca Sieb., C. equisetifolia L., C. cunninghamiana Miq.) and one hybrid, cultivated irregularly in the stands.
Consequently, we dealt with Casuarina. spp. as one species in this study to avoid mis-identification during
sampling process.
Data collection
Sampling and data collection were conducted during the main growing season of the trees in 2010
(spring and summer). In all the four forests, and for each tree species, we conducted the measurements through
North-South transects (lengths ranging from 300 to 700 m). The number of transects was depending on the area
occupied by the tree species in each forest. The positions of transects were decided to represent all the apparent
variations in growth and height of trees. Twenty five trees were measured across their whole height range. The

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EMAD FARAHAT and HANS W. LINDERHOLM

selected trees were healthy, upright standing, and had undamaged crowns and with no forks or obvious weak
vigor according to the criteria suggested by Poorter et al. (2006). Total height (H), crown length (CL), average
crown diameter (Cdia), projected crown area (Carea), and stem diameter at breast height (dbh at 1.3 m) were
measured in the field for each tree. Height and crown lengths were measured with a clinometer (model HEC-R,
Haglf, Sweden), while the dbh was measured with a diameter tape. The base of the crown was defined as the
lowest leaf-bearing branch. At the end of the main growing season for all trees (in summer) and before leaf
senescence in autumn, the average crown diameter was calculated as the mean of two perpendicular diameter
measurements, one of which was the maximum crown diameter. The crown diameter was measured in north
south and eastwest directions, and for smaller trees in the direction of the longest width and perpendicular to
it. Crown area was calculated as an ellipse according to the shape of the trees, using the equation 0.25 d 1 d 2 , in
which d 1 and d 2 are the two crown diameter measurements per tree, and crown length as the height of the tree
minus the height of the trunk beneath the lowest leaf-bearing branch (Beaudet & Messier 1998, Poorter et al.
2006). The crown shape (CS) of each tree was described as the ratio of crown length to mean crown diameter
(see Beaudet & Messier 1998).
Leaf mass per area (LMA) was calculated for each tree species (10 tree individuals, 10 fully expanded
leaves/ individual). The sampled leaves were collected during the main growing season for trees from all the
canopy directions at a height range 1.5-3.0 m above soil surface. In the case of Cupressus sempervirens, ten
small branches with their carrying scale-like green leaves were sampled for this analysis. The sampled leaves
were scanned, and then leaf area was measured from the scanned image. Measurements of leaf area was
performed, where the public domain ImagJ 1.44 image program (available on the internet at website:
http://imagej.nih.gov/ij/download.html) was used to estimate the fresh leaf area. The leaves were weighed after
oven-drying at 60 C for at least 48 h. Petioles of leaves were excluded from the measurements of the leaf traits
(leaf area and leaf dry mass), because their function is the support of leaf blade. To determine the growth rate of
each tree per life time, as a function of both stem height (HGR) and dbh (DGR), we assumed that the initial
stem height and dbh for the trees at the time of transplanting was 0.4 m and 1.0 cm respectively, based on what
was reported by the workers in the forests (personal communications). These values were subtracted from each
tree height and dbh to calculate its growth rate per life time. We divided the tree height by its life time (i.e. age
in years) to obtain rough estimate for the growth rate of the trees in each forest.
Soil and water analysis

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Soil samples were collected from five locations in each forest, under the tree canopies, and each
location was represented with one composite sample (N=3). Soil samples were collected through soil profiles
including the topsoil layer at a depth of 0-50 cm. The samples were used as representative samples for each
forest. The soil samples were brought to the laboratory in plastic bags shortly after collection, spread over paper
sheets; air dried, passed through a 2 mm sieve to remove gravel and debris, and then packed in paper bags
ready for physical and chemical analysis. Soil texture was determined by a set of stainless steel test sieves
confirming to ASTM standards, whereby the percentage of clay, silt and sand were calculated. Soil extracts
were prepared to meet the requirements for different determinants, 1:5 (w: v) soil (g): distilled water (ml)
extract. This extract was used to measure soil conductivity (EC), pH, bicarbonates, calcium, magnesium,
sodium and potassium. The pH values of soil samples were determined using a glass electrode pH meter
(Model 9107 BN, ORION type). Salinity was evaluated by a conductivity meter (dS m-1). Bicarbonates were
estimated by titration against 0.01N HCl using methyl orange as indicators (Allen et al. 1989, Maff 1986).
Soluble soil cations (Ca, Mg, K and Na) were determined by atomic absorption spectrometer in the Central
Laboratory of Ein-Shams University, Egypt.
Water samples were collected in plastic bottles directly from the main pipe feeds in each forest and
brought to the laboratory shortly after collection. pH and EC were determined directly after collection, after
which the samples were kept at 4 C for further analysis. Bicarbonates, calcium, magnesium, sodium and
potassium were estimated using the same methods as in the soil analysis.
Data analysis
To test the significance of the spatial variations in crown architecture, size and leaf traits, we compared
the obtained data for each tree species among the forests using the analysis of variance (ANOVA). The same
technique was applied on growth rates as functions of height and dbh. Pearsons correlations coefficients were
calculated for the architecture traits of each tree species and height in all locations using STATISTICA 7.0
Software (StatSoft. Inc. 2004).
Analysis of allometry
To check the relationships between height and dbh, we first regressed height data against dbh, and
polynomial fitted second-order equations for each relationship were conducted using Microcal Origin 5.0
Software (Microcal Software Inc. 1997). Bivariate trait relationships were analyzed using Pearsons
correlations on untransformed trait values. To assess multivariate trait associations among species, principal

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EMAD FARAHAT and HANS W. LINDERHOLM

component analysis (PCA) was done on the grouped traits data of height and crown architecture for each
species in all sites, using PC-ORD v.3.0 (McCune & Mefford 1997).
Environmental variable-trait relationships
To check the relationship between the measured tree traits and environmental variables (soil and
water) for all species, canonical corresponding analysis (CCA) was used. All soil and water parameters were
included in this analysis and all the default settings for the CCA in the PC-ORD software were used (McCune
& Mefford 1997). To assess the importance of all environmental variables, the intra-set correlations from CCA
were used, where the significance of the correlations with the CCA axes was tested by calculating Pearsons
correlation coefficients for the variable at p < 0.05.
Results
Spatial variations in crown architecture, foliar traits and growth rate of trees
The highest average values for size traits (H, dbh), Cdia and Carea of E. camaldulensis were obtained
in El-Tur forest, while most of the lowest values for the traits were found in either Sadat or Wadi El-Natrun
forests (Fig. 2). The highest values of dbh, LMA and crown shape for Cupressus sempervirens were found in
Wadi El-Natrun forest, while the values of the other traits were lower than that in Sadat and Sarapium forests.
The size traits and Carea growth of Dalbergia sissoo were greater in Sarapium forest compared to Sadat site. In
contrast LMA was greater at Sadat than Sarapium at which the species had its highest mean LMA value. The
growth performance of Khaya senegalensis was higher in Sarapium than Sadat forest for most traits with
significant differences in H values. Generally, E. camaldulensis and Casuarina spp. had significant Cdia, Carea
and dbh in El-Tur forest greater than other sites while all the species showed more H and dbh growth rates in
Sarapium forest.
Significance of the spatial variations in traits
The results of the one-way ANOVA tests showed that most of the measured traits were significantly different
(at p < 0.05) for individual tree species among the forests (Table 3). The stem H and dbh of a species generally
varied among the forests but the species maximum H did not necessary coincide with maximum dbh at the
same site, and this was independent of tree age. For instance, the highest dbh values of Casuarina spp. (at ElTur forest, flood irrigation, age >13 years) and C. sempervirens (at Wadi El-Natrun forest, drip irrigation, age
H10 years) did not coincide with the obtained highest stem H values for species (Table 3). Despite the age
difference between K. senegalensis plantations in Sadat and Sarapium forests (H 2 years), there was no

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207

significant difference in stem dbh values (at p < 0.05). It is appeared from the results of K. senegalensis that the
relationship between the two size traits was very weak for the species (r = 0.2).
Leaf mass per area (LMA) was significantly different at p < 0.05 for all species among forests, with
maximum values in Wadi El-Natrun forest (for E. camaldulensis and C. sempervirens) or Sadat forest (for D.
sissoo and

Fig. 2. Spatial variation of the architecture traits, size traits, foliar traits and growth rates for each tree species
among forests. H = height, dbh = stem diameter at breast height, LMA = leaf mass per area, Cdia =
crown diameter, Carea = crown area, CL = crown length, CS = crown shape, HGR = growth rate using
stem height, DGR = growth rate using dbh data. Error bars represent the standard error of mean.

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EMAD FARAHAT and HANS W. LINDERHOLM

Table 3. The results of One-Way ANOVA for **

the measured tree traits in different locations. * = significance at
probability level < 0.05 or p < 0.01, = significance at probability level < 0.0001. Within each row,
mean values with the same letter do not differ significantly between treatments at p < 0.05.
Wadi ElEl-Tur
F
Trait
Species
Sadat
Sarapium
Natrun
a
ab
ac
d
Height (m)
E. camaldulensis
13.6
14.6
12.6
17.3
15.4**

dbh (cm)

LMA -2
(mg cm )

Casuarina spp.

9.0a

13.5b

9.7ac

10.7c

25.5**

C. sempervirens

10.2a

10.0a

7.3b

17.8**

D. sissoo

7.7a

10.1b

44.4**

K. senegalensis

5.5

12.0

115.3**

E. camaldulensis

17.7a

19.8ab

15.1ac

25.8d

17.3**

Casuarina spp.

10.2a

14.0b

13.8b

22.3d

41.3*

C. sempervirens

11.8a

12.9a

14.8b

4.5*

12.9

25.6**

10.8

D. sissoo

9.1

K. senegalensis

11

Crown shape

Height
growth rate
(m yr-1)

dbh growth
rate
(cm yr-1)

0.01

11.1

11.0

18.1

11.2

10.2**

C. sempervirens

23.9a

21.6a

34.8b

35.8**

D. sissoo

15.9

11.0

22.9**

K. senegalensis

8.5a

6.1b

7.5*

3.2a

4.8b

3.9ab

7.2c

15.1**

3.5a

3.8a

5.5b

5.6b

13.2**

Casuarina spp.

Crown length (m)

E. camaldulensis

Crown diameter (m) E. camaldulensis

Crown area (m2)

C. sempervirens

1.9

2.3

1.2

8.7*

D. sissoo

4.4

5.1

0.8

K. senegalensis

4.2

5.8

E. camaldulensis

8.9

18.8
a

14.4

14.8**

43

10.4

12.0

25.0

25.0

13.3**

C. sempervirens

2.9a

4.4a

1.3b

7.8**

D. sissoo

17.0

22.3

0.9

K. senegalensis

15.3

30.3

8.1

5.3

21.1**

11.2 a

8.8ab

10.4a

1.3c

80.7**

C. sempervirens

8.7

8.1

7.9

0.5

D. sissoo

7.2

6.7

0.5

8.0

45.8**

10.9

Casuarina spp.

3.3

3.1

E. camaldulensis

3.2
c

Casuarina spp.

K. senegalensis

4.5

E. camaldulensis

3.9 a

0.7b

2.3c

1.4d

17.1**

Casuarina spp.

3.3 a

2.4b

1.9c

1.3d

18.5**

C. sempervirens

5.0 a

4.0ab

7.7ac

5.1**

D. sissoo

1.8

1.4

2.9

K. senegalensis

1.1a

1.7b

4.0

1.6

0.8

0.8

92.8*

0.9 a

2.2b

0.9ac

0.8ad

202.2**

E. camaldulensis

0.9

Casuarina spp.

ac

1.6

0.7

109.0*

0.7 a

1.2b

134.2**

K. senegalensis

0.5 a

1.6b

159.8**

E. camaldulensis
Casuarina spp.
C. sempervirens
D. sissoo

1.0 a
0.9aa
1.1
0.8 a

1.6b
2.2bb
2.0
1.5b

1.3c
1.3cc
1.4
-

1.6bd
1.6d
-

12.3**
47.8***
34.3
70.5**

K. senegalensis

1.0 a

1.4b

9.6*

C. sempervirens

1.0

D. sissoo

GROWTH PERFORMANCE OF FOUR

209

K. senegalensis). LMA value for C. sempervirens in Wadi El-Natrun forest was 2- to 3- fold higher than the
other tree species.
The variations in crown-architecture traits were more profound than the size and foliar traits.
Significant variations in Cdia and Carea were found for C. sempervirens, with the highest values obtained in ElTur forest for E. camaldulensis and Casuarina spp. and in Sarapium forest for the other species (Table 3). The
highest values for CL and CS were obtained in Sadat forest for all species except K. senegalensis and C.
sempervirens, while the spatial variations of these traits were highly significant (at p < 0.05) for E.
camaldulensis and Casuarina spp. and size-dependent more than that found for other tree species.
Height growth rate (HGR) and diameter growth rate (DGR) for each species were significantly different in all
forests (at p < 0.05). The highest growth rates for all species, irrespective of their ages, were found in Sarapium
forest (Table 3). Non-significant variations (at p < 0.05) in HGR and DGR around the mean value were
observed between species in the same forest (data not shown).
Traits associations
The relationship between H and dbh was highly significant, at p < 0.0001 or p < 0.01, for all species
except C. sempervirens (Fig. 3). The equations indicated a nonlinear relationship between both variables for all
species and among forests. The best values for the coefficient of determination (R2) between the two variables
were obtained for D. sissoo (R2 = 0.55) and E. camaldulensis (R2 = 0.37). Although K. senegalensis had more
or less equal mean values for size traits compared with D. sissoo, the coefficient of determination was very low
(R2 = 0.16) indicating a weak H-dbh relationship.
The correlation between H and crown traits (diameter, area and length) was often size-dependent; the positive
sign of the correlation coefficient (r) increased with decreasing tree heights (Fig. 4). The highly significant
positive correlations between H and crown traits were observed in the canopies of D. sissoo, while the lowest
correlations were found in E. camaldulensis canopies. Crown length was strongly correlated with height in all
species (r range: 0.36-0.96), while the correlation between height and crown shape was not significant for any
species (at p < 0.05). The correlation pattern between H and crown traits was similar for each species in all
forests, with non-significant variations (data not shown).

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EMAD FARAHAT and HANS W. LINDERHOLM

Fig. 3. Polynomial regression for the stem height (log H) and diameter (log dbh) for tree species in all forests.
R2 value is shown when the relation is significant at probability level, p < 0.0001 or p< 0.01.

Fig.4. Pearsons correlation coefficients (r) between height and crown architecture traits of the sampled tree
species (crown diameter = Cdia, crown area = Carea, crown length = CL and crown shape = CS). The
trees are plotted on x-axis from shorter to taller species.

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211

The associations of architectural traits and height for all tree species in all locations were explored with
PCA (Table 4). The first PCA axis explained 50.6% of the variation in traits, and had high positive correlation
with Cdia (0.92) and Carea (0.89). In contrast, CS had high negative correlation with the first PCA axis (-0.82).
On the other hand, the second PCA axis explained 33.9% of the variation in architecture traits, with highly
negative correlation with height (-0.86) and CL (-0.73).

Table 4. Eigenvalues and Euclidean distance for the measured traits and its correlation to the first three axes of
PCA analysis. The last two columns indicate the Pearsons correlation between the traits and the first
and second axes of PCA. Significant correlations are shown in bold.

Eigen-value
% of variance
Height (m)
Crown diameter (cm)
Crown area (m2)
Crown length (m)
Crown shape

1
2.28
52.60
0.20
0.60
0.57
-0.18
-0.48

PCA axis
2
1.28
30.04
-0.66
-0.09
-0.11
-0.67
-0.27

3
0.78
8.96
0.53
-0.07
-0.02
-0.71
0.44

PCA 1

0.03
0.97
0.92
-0.30
-0.78

PCA 2

-0.82
-0.11
-0.14
-0.82
-0.34

Soil and water characteristics and CCA correlations

The analysis of soil samples showed an increase in the soil salinity (expressed as EC) in both Wadi ElNatrun and Sadat forests compared with the other forests (EC = 7.7 and 5.2, respectively) (Table 5). Likewise,
soil soluble Mg, K and Na cations were higher in these two forests than in El-Tur and Sarapium forests. On the
contrary, the highest values for Ca (1235.6 ppm) and Na (1377.7 ppm) were obtained in El-Tur and Wadi ElNatrun forest, respectively. There were no distinct variations in the quality of water among the forests despite
the differences in the quality of treatments. The irrigation water in Wadi El-Natrun forest (ground water) only
differed from the others with its high EC (5.1 dSm-1) and Na concentration (130.2 ppm).
The canonical corresponding analysis for environmental variables (soil and water) and species
measured traits showed weak eigenvalues with the three canonical axes. The total variance in the species data
was 0.25, indicating a weak variability in the environmental data among forests (ter Braak 1990). The first
canonical axis explained 40.8% of the variance, while the second canonical axis explained only 5% of the
variance (Table 6). As expected from the total variance in species data, these results suggested a weak
association between measured traits and the environmental variables. CCA axis 1 was negatively correlated
with soil cations and water K and positively with soil sand (at p < 0.05). On the other hand, CCA axis 2
correlated positively with soil clay content, Mg, K, Na and water Mg and Na, and negatively with soil pH and
Ca in water and soil.

Egypt. J. Bot. 52 (2) (2012)

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EMAD FARAHAT and HANS W. LINDERHOLM

Table 5. General soil and irrigation water characteristics of the studied planted forests. All the measured
parameters are significantly different between forests at p < 0.05.
Parameter

Sadat

Sarapium

Wadi ElNatrun

El-Tur

1-Soil: Physical properities

Soil type

Sandy

Sandy

Sandy

Calcareous

Sand

79.4

85.0

80.1

80.3

Silt

10.1

11.5

11.3

16.5

Clay

10.5

4.6

8.6

4.1

7.8

7.6

7.9

8.1

5.2

2.1

7.7

4.5

Ca

700.0

81.5

269.2

1235.6

Mg

165.4

77.2

132.7

46.1

330.7

138.9

212.6

121.8

Na

1332.2

234.3

1377.7

345.4

175.1

293.1

112.4

207.4

7.5

7.2

7.8

7.3

EC (dSm )

3.7

2.1

5.1

3.9

Ca

128.3

151.4

44.8

191.6

Mg

36.3

38.6

14.9

17.5

Soil particles (%)

pH(1:5)
-1

EC (dSm )
Soluble Cations (ppm)

Soluble anions
HCO 3 (ppm)
2- Water properities
pH
-1

Ppm

23.4

17.9

4.9

14.0

Na

105.8

76.3

130.2

68.7

HCO 3

110.1

85.1

154.9

120.3

Discussion
Intra- and inter-specific spatial heterogeneity of traits
The cultivated species in our study were worldwide and nationally tested for their drought resistance, salinity
and waterlogging tolerance (El-Lakany 1983, Aswathappa & Bachelard 1986, Van Der Moezel et al. 1988,
1989, Gilman & Watson 1993). The crown architecture, size and foliar traits revealed plastic changes among
forests. The growth performance of E. camaldulensis was more pronounced in El-Tur and Sarapium forest for
most measured traits. Meanwhile it had general poor performance in Wadi El-Natrun forest which could be
attributed to general poor silviculture operations and site conditions. On the other hand, the greater value of
LMA for C. sempervirens in Wadi El-Natrun forest, which is classified as a soil salinity tolerant species

GROWTH PERFORMANCE OF FOUR

213

Table 6. Intra-set correlations (of ter Braak 1986) of the soil and water variables with the first three axis of the
canonical correspondence analysis (CCA), together with eigenvalues and traitsenvironment
correlation coefficients. * = significance at p < 0.05.
CCA axis
1

Eigen values

0.10

0.01

0.01

% of variance explained

40.8

5.0

2.4

Cumulative % of variance
explained

40.8

45.8

48.2

Soil
variables

Sand %

0.80*

0.30

0.032

Silt %

0.02

-1.00

-0.00

Clay %

-0.62*

0.67*

-0.40

pH

-0.48

-0.68*

-0.54

EC (dSm-1)

-0.35

0.09

-0.93*

Ca (ppm)

-0.75*

-0.65*

-0.60*

Mg

-0.51*

0.79*

-0.32

-0.73*

0.66*

-0.17

Na

-0.52*

0.54*

-0.65*

HCO 3

0.45

-0.10

0.88*

pH

-0.22

0.32

-0.92*

EC (dSm-1)

-0.37

-0.09

-0.90*

Ca (ppm)

-0.17

-0.57*

0.80*

Mg

-0.00

0.56*

0.80*

-0.51*

0.29

0.80*

Na

-0.16

0.58*

-0.79*

HCO 3

-0.16

-0.11

-0.97*

Water
variables

(Gilman & Watson 1993), might indicate its adaptability to the local site unfavorable conditions more
than other species.
The significant differences in tree species growth among forest reflect direct responses and plastic
changes in traits according to the prevailing conditions has previously been shown for trees in natural forests
(e.g. Pena-Rojas et al. 2005, Stefanos et al. 2008). It is revealed from the significant correlations between tree H
and dbh, that trees tend to invest more resources in radial growth than in height growth after certain age. This
was more pronounced in El-Tur and Wadi El-Natrun forests compared with Sadat and Sarapium forests. We
can attribute this growth pattern for the presence of considerable canopy gaps for most trees in these forests,
which enhance the plant to invest more resources in radial growth than vertical growth. Furthermore, flood
irrigation in El-Tur seems to be another enhancing factor assist in this growth pattern, while the availability of
water and waterborne-nutrients secure for the trees more resources to invest in horizontal growth (Waterworth

Egypt. J. Bot. 52 (2) (2012)

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EMAD FARAHAT and HANS W. LINDERHOLM

et al. 2007). This uni-direction investment of resources was confirmed by H and dbh data of K. senegalensis.
Similar correlation patterns were reported by Poorter et al. (2006) for tropical trees, in which the tree first
grows mainly in height until it reaches its reproductive size (or the canopy); after which height growth levels
off and diameter expansion continues. This correlation pattern between H and dbh was expected to be clearer in
the planted forests due to absence of interspecific competitions and approximately fixed distance between trees
which minimize the intra-specific competition for light and resources.
In spite of the fact that we have no direct evidence, the differences in LMA among tree species are
more likely related to their susceptibility to unfavorable site conditions in Wadi El-Natrun and Sadat forests.
This may be supported by the obtained high EC in water and soil samples and the observed low frequency of
irrigation and poor silvicultural management practices in both sites. Increasing LMA is a known plastic strategy
for the species to minimize water loss in arid regions (Pena-Rojas et al. 2005, Stefanos et al. 2008), and it has
been shown to be a main adaptation to drought and related to photosynthetic tissue per unit area (Marcoco et al.
2000, Damoura et al. 2008, Chen & Wang 2009). For this clear plastic change in LMA among species, TanakaOda et al. (2010) proposed the likelihood of LMA to be new important criteria for plantation species selection
on the Loess Plateau, China.
Crown diameter and area increased with the height of trees, which increase the amount of
photosynthesizing leaf area and capturing of light (Givnish 1988, Beaudet & Messier 1998). Both crown area
and diameter increased more strongly with tree height than crown length and shape. These were the same
findings for tropical rain forest trees in Bolivia (Poorter et al. 2006) and many other tree communities (Parent &
Messier 1995, Chen et al. 1996). The crown length of the trees in planted forests may be subjected to artificial
partial pruning (e.g. Italian cypress) or natural peeling out and death for bark and lower branches as in Eucalypt
and Italian cypress. Crown shape is an important trait affecting the light environment in forest canopies
(Chmura et al. 2009). In this study, the changes in crown shape may be related to tree genetics, stem density
and age as the most likely factors affecting tree shape in planted forests as reported for natural forests (Ishii &
McDowell 2002).
The intra-specific differences in growth rate per life for tree H or dbh can be interpreted as a direct
response to the local environment and silviculture practices. However, although water is the main key limiting
factor for plants in the desert, the highest values for species growth rates was in Sarapium forest. In other
words, the other biotic (e.g. silviculture practices) and abiotic factors (e.g. soil fertility) beside water are key
factors which determine the growth of tree species in desert plantations. To confirm this hypothesis, a study on

GROWTH PERFORMANCE OF FOUR

215

the biomass production in drip- and flood irrigated forests should be carried out for the same tree (s) species.
The observed non-significant growth rate differences between tree species in the same forest cannot be
understood as an equal performance for these species or the environmental conditions are even for all tree
canopies. This simply could be ascribed to the changes in species growth with age and according to growth
stage; which differ inherently between species (Sillett 2010).
H-dbh and crown architecture association
Non-linear relationships between tree H and dbh has previously been reported for natural forest trees,
where tree height increases steeply with dbh, and starts to cease when species attain their maximal height and
start to reproduce (Thomas 1996, Poorter et al. 2006). In our study, the more correlated H-dbh relation for D.
sissoo compared with other small species (e.g. K. senegalensis) or large species (e.g. E. camaldulensis) may be
due to that Sissoo trees start to reproduce at small dbh. This may foster the species to invest more resources in
dbh growth rather than height, to cope with reproductive needs and carry the developing crown. Moreover, this
could be a strategy for the woody vegetation in hyper-arid lands to provide safe sights for their self regeneration
(Shaltout & Mady 1993). On the contrary, D. sissoo individuals grow in five natural habitats in Nile Delta,
Egypt; tend to expand vertically rather than horizontally when growing in dense populations (Shaltout et al.
2010). The observed negative correlation between crown length and diameter at small heights in this study
agrees with the finding of Beaudet and Messier (1988) and Poorter et al. (2006). The impact of site and height
size did not have any effect on the crown shape of the species which let us to claim that this trait is more
species specific and may be related to other factors (e.g. age and density) more than the other crown traits. This
is in agreement with the findings of Chmura et al. (2009) for young stands of loblolly pine (Pinus taeda L.) in
the southeastern US.
The PCA 1 was highly correlated with light capture traits of the species (Carea and Cdia) while the
PCA 2 was high negatively correlated with the stature of the species (H and CL). Meanwhile, crown shape was
negatively correlated with both PCA axes. This seems logic and accepted correlation pattern in presence of very
weak or absent competition among a tree species, where the tree individuals are almost equally spaced and
watered. This suggests that a tree invests more in building a robust and extended crown to maximize their
capture for light while the need for tall stature is not necessary, which is similar to trees growing in open
canopies (Gasser et al. 2006, Poorter et al. 2006). Crown diameter was found to be the best predictor for the dry
weight of the total aboveground parts and leaves of D. sissoo in Nile Delta, Egypt (Shaltout et al. 2010)

Egypt. J. Bot. 52 (2) (2012)

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EMAD FARAHAT and HANS W. LINDERHOLM

The weak species-environment relationship and presence of high correlation with soil sand content
could be explained by the fact that in drip irrigated soil, the root system concentrates in the wetted zone
developed in the irrigated side of the tree (Sokalska et al. 2009); by presence of more sand content, the root
system will grow more downwardly to reach the permanent water table. In contrast, in case of flood irrigation,
sand help in more growth and aeration around the root system which finally help the tree to grow normally. In
addition, the long root system assists in stability of trees and provides more resistance against the prevailing
wind in our deserts (Sterken 2005).
Management implications
Results from this study improve our knowledge and provide new tools for understanding growth
performance of tree species in planted forests in arid lands. This information can be used to inform the design
of forest management prescriptions. Detailed understanding for the growth performance of crown traits and its
relationship with tree height in different desert regions and at different ages assists in decisions on spacing
when planning the spacing of single and mixed tree stands. Moreover, it may help in deciding which tree can be
selected in a certain site exposed to a range of environmental conditions according to the crown parameters.
Our results improve knowledge about spatial variations in crown architecture of the tree species which may
lead to improved forest productivity overall. The importance of silviculture management practices are
becoming commonplace, but the contribution of these practices in the overall growth of species in desert
plantations should be understandable for the managers in artificial plantations.
Conclusion
In conclusion, the ecological performance of tree species was significantly different between forests.
E. camaldulensis had their best growth in El-Tur and Sarapium forest with general poor performance in Wadi
El-Natrun forest. The greater value of LMA for C. sempervirens in Wadi El-Natrun forest compared to other
species might indicate its adaptability to the poor local site conditions more than other species which in
consistent with many reported results . The highest H and dbh growth rates for the tree species in Sarapium
forest revealed that growth of trees in desert plantations not correlated only with tree age or water availability
but also with site conditions and silviculture practices. The non-linear relationship between H and dbh in our
planted forests coincides with that reported for natural forests which may indicate that this is genetically and
environmentally controlled performance. The investment of small trees to more resources in building strong
stem diameters more than tall stems at certain moment in lifecycle was a strategy for the tree species in the
desert plantations. The crown diameter and crown area, as a reflector to light capturing capacity, were much

GROWTH PERFORMANCE OF FOUR

217

stronger predictors for change in growth performance of the tree species and were more size-dependent (Poorter
et al. 2006); which confirmed by PCA. Crown shape was the least important architecture trait and has no direct
positive correlation with height. The growth of trees was not only affected by the availability of water or the
type of irrigation, but also with many other environmental factors. Soil sand content was the main
environmental factor correlated with variations in species traits among forests as shown by CCA. Our results
are important in management and planning of future man-made forests.
Acknowledgments
This work was supported by the Swedish Research Council; and Swedish International Development
Cooperation Agency SIDA [grant no. 2008-6087]; and partial funding was provided for the first author by
Helwan University. The authors wish to express their thanks to The Ministry of Agriculture and Reclaimed
Lands, Egypt and Dr Saed Khalifa, the Director of Central Administration for Afforestation and Environment
Protection, for their facilities and permit access to the locations of present study.

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(Received 18/12/2011; accepted 26/1/2012)

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