全 文 ：Journal of Forestry Research (2009) 20(1):15–22
DOI 10.1007/s11676-009-0003-5
-





Aboveground nutrient components of Eucalyptus camaldulensis and E.
grandis in semiarid Brazil under the nature and the mycorrhizal inocu-
lation conditions

Marcela C. Pagano1*, Antonio F. Bellote2, Maria R. Scotti1
1Microorganism-Plant Interaction Laboratory, Institute of Biological Sciences, Federal University of Minas Gerais, Av. Antônio Carlos, 6627, Pampulha,
CEP: 31270-901, Belo Horizonte, MG, Brazil.
2Embrapa Forestry, Estrada da Ribeira, km 111, Caixa Postal 319 - Colombo, PR - 83411-000- , Brazil.

Abstract: A study was conducted to evaluate the aboveground biomass, nutrient content and the percentages of mycorrhizal colonization in
Eucalyptus camaldulensis and Eucalyptus grandis plantations in the semiarid region (15° 09 S 43° 49 W) in the north of the State of Minas
Gerais in Brazil. Results show that the total above-ground biomass (dry matter) was 33.6 Mg·ha-1 for E. camaldulensis and 53.1 Mg·ha-1 for
E. grandis. The biomass of the stem wood, leaves, branches, and stem bark for E. camaldulensis accounted for 64.4%, 19.6%, 15.4%, and
0.6% of the total biomass, respectively (Table 2); meanwhile a similar partition of the total above-ground biomass was also found for E.
grandis. The dry matter of leaves and branches of E. camaldulensis accounted for 35% of total biomass, and the contents of N, P, K, Ca, Mg,
and S in leaves and branches accounted for 15.5%, 0.7%, 12.3%, 22.6%, 1.9%, and 1.4% of those in total above-ground biomass, respec-
tively. In the trunk (bark and wood), nutrient accumulation in general was lower. Nutrient content of E. grandis presented little variation
compared with that of E. camaldulensis. Wood localized in superior parts of trunk presented a higher concentration of P and bark contained
significant amounts of nutrients, especially in E. grandis. This indicated that leaving vegetal waste is of importance on the site in reducing
the loss of tree productivity in this semi-arid region. The two species showed mycotrophy.
Keywords: Eucalyptus; biomass; nutrient components; semi-arid region; mycorrhizal symbioses; Brazil



Introduction

Eucalyptus, native from Australia, with more than 600 species,
has been used as a monoculture in afforestation programs. In
many countries these plantations are used in cellulose industries,
pharmaceutics and hygiene. The Eucalyptus species present
characteristics suitable for commercial use, such as fast growth,
high cellulose production and resistance to environmental stress
and diseases (Santos et al. 2001). In Brazil, for a sustainable
production in intensive systems of wood extraction, to keep or
improve the nutritional soil levels is essential. Studies showed a
general upward trend in N fertilizer requirements in commercial

Foundation project: This study was financed by CAPES (Coordenação de
Aperfeiçoamento de Pessoal de Nível Superior)
Received: 2008-03-28; Accepted: 2008-10-30
© Northeast Forestry University and Springer-Verlag 2009
The online version is available at http://www.springerlink.com
Biography: Marcela Claudia Pagano (1966-), *Corresponding author,
female, Major: Botany; Postdoctoral researcher at Federal University of
Minas Gerais, Brazil . e-mail: marpagano@gmail.com
Responsible editor: Chai Ruihai

eucalypt plantations (Gonçalves et al. 2004; Laclau et al. 2005;
Corbeels et al. 2005) and the N inputs are a major cost in Brazilian
silviculture not to mention the potential risks of pollution and
leaching of nitrates in tropical forest soils (Fisher and Binkley
2000).
Gonçalves (1995) reported the nutrient accumulation in 5–6
year-old Eucalyptus grandis plantations in Brazil and observed
that during harvesting part of nutrients remain in the ecosystem
accumulated in leaves, branches and litter when these are not
removed from the site. Nevertheless, 30% of nitrogen (N),
phosphorus (P) and calcium (Ca), and 43% of potassium (K) are
removed when wood is extracted. The loss of N, P, K and Ca
increases 40%, 60%, 65%, and 48%, respectively, when bark is
extracted with wood. Mineral fertilization is a common practice to
improve productivity; however, management policies need fur-
ther studies. Most biomass and nutrients accumulated by planted
E. grandis occurred between two and five years of age, when the
leaf area is expanding. E. camaldulensis is very widely distributed
in inland Australia along river banks. This fast growing tree
species can tolerate moderate salinity, alkalinity, extended dry
seasons and waterlogging, and is extensively planted throughout
the world for purposes such as shade, shelter, agroforestry, fur-
niture and industrial wood production (Midgley et al. 1989;
Marques Júnior et al. 1996). The rotation length is about 3–5 years
for E. camaldulensis and 7 years for E. grandis (Campinhos
RESEARCH PAPER
Journal of Forestry Research (2009) 20(1):15–22

16
1999).
Sandy soils are characterized by little water and nutrient reten-
tion, which means difficult management and vegetation devel-
opment in these areas, the presence of arbuscular mycorrhiza
being important to increase the capacity of water and nutrient
absorption by plants (Córdoba et al. 2001). It is important then to
know the mycorrhizal condition of species in order to allow for
research on seedlings production and technologies for a successful
reforestation.
Arbuscular mycorrhiza (AM) are substantially involved in the
vegetative state of the mycotrophic plants, defining their ecologic
niches, influencing vegetal communities composition, mainte-
nance and soil fertility, plant fitness, and nutrient turnover
(Jeffries et al. 2003). Some plant species, like Eucalyptus spp.,
have the capacity to form two types of mycorrhizas, arbuscular
and ectomycorrhiza (ECM) (Malajczuk et al. 1981; Zambolim
and Barros 1982). Establishment of AM association in Eucalyptus
has been known for over 20 years, and the benefits of symbiosis
have proved commercially relevant (Zambolim and Barros 1982;
Coelho 1997; Gomes and Trufem 1998; Grazziotti et al. 1998;
Santos et al. 2001). Studies of AM inoculation on eucalypt are
increasing (Standish et al. 2007) and Glomus sp. was used as
inoculum (Arriagada et al. 2007). As regards ECM, Molina et al.
(1992) and Thomson et al. (1996)’s studies suggest that the
majority of fungi, selected for their enhancement of seedling
growth, may persist only for short periods in the field, where they
are replaced by wild populations. Whereas, Pampolina et al. (2002)
showed the significance of ECM in immobilizing P and other
nutrients as well as the impact of P fertilization, as well as, Chen et
al. (2000) and Mason et al. (2000) recommended the inoculation
with tested ECM. There are numerous studies on ECM; however,
works on E. camaldulensis and E. grandis AM/ ECM colonization
in field conditions are to the best of our knowledge few.
The need to evaluate the success of these Eucalyptus spp. at the
Jaíba region motivated this study of plant nutrient content and
mycorrhizal status, aiming at increasing the possible ways of
forest management in the region, and as a productive model for
revegetation of local degraded areas with the objective of mixed
them with native local species. This information is essential to
identify methods for sustainable management.
The objective of the present study was: (1) to evaluate growth
and quantify nutrient content of aboveground dry mass in
28-month-old pure stands of Eucalyptus camaldulensis and E.
grandis, and (2) to examine the mycorrhizal root colonization.

Materials and methods

Study area, design, determination of biomass and nutrient quan-
tification

The study area is located in the semiarid region (15°09 S 43°49
W) in the north of the State of Minas Gerais in Brazil, and is
characterized by annual pluviometric rates of 800 mm concen-
trated in the spring-summer months from November to January,
there being about 10 dry months (Prado 2003). According to
Köppen, the climate of the region is BSh type (semiarid)
(Carvalho 2003). Mean annual air temperature ranges from
24.2–28.1ºC, with mean annual temperatures of 34ºC in the
hottest month (January) and 14.8ºC in the coldest month (July).
Annual precipitation is 749 mm. Precipitation in December and
July are 217 mm and 1 mm, respectively (Mocambinho Agro-
climatic Station). Predominant soil types are Quartzarenic Neosoil
with high infiltration rate. Furthermore, they are moderately acid
and have small amounts of soil organic matter.
The present study focuses on Eucalyptus monocultures within
the scope of a broad project dealing with the introduction of mixed
and monocultures of Eucalyptus in a 10-ha experimental area,
after the woody Caatinga had been cut and occupied by degraded
vegetation named “Carrasco”, with the aim of providing wood
supply and minimising exploratory actions in biological reserves
in the northwest of Minas Gerais state, Brazil. The experimental
site (1.5 ha) was cleared of “Carrasco” vegetation (with retards the
natural succession of forest) and Eucalyptus camaldulensis Dehnh
and E. grandis Hill ex Maiden were cultivated in monocultures.
Seedlings were transplanted during the rainy season in 2001,
using a randomized block design with 42 plants, which were
randomly distributed in each of the three blocks per site. Each
block was composed by one plot of 378 m2 (21m×18 m) with
single plantations and 42 plants per block, with a spacing of
3m×3m. These plots were irrigated for about 10 months (Duarte et
al 2006, Pagano et al. 2008).
For the present study we selected two experiments: 1- E.
camaldulensis inoculated with AMF, and 2- E. grandis inoculated
with AMF. The AMF inoculum consisted of a mix of three species
Gigaspora margarita, Scutellospora heterogama and Glomus
brohultii from the ICB-UFMG laboratory collection. Endomy-
corrhizal inoculation was accomplished by placing 1 ml of sus-
pension composed of 50 spores of each AMF species.
Fertilization consisted in triple superphosphate (400 kg·ha-1),
KCl (305.6 Kg·ha-1), MgSO47H2O (40 kg·ha-1), ZnSO47H2O
(37.4 kg·ha-1), Mo7O24H2O (1.4 kg·ha-1), urea (177.6 kg·ha-1)
corresponding to 80% of complete fertilization following So-
masegaran and Hoben (1985), and was applied at the beginning of
the experiment. Plantation was then carried out and, when nec-
essary, ant colonies were exterminated with formicide. In subse-
quent years, plantations were rid of ants, weeds and low branches.
Soil samples were collected from the top 20 cm of 3
spots/block/experiment (3 subsamples x 3 blocks x 2 experiments).
Composite samples (2) were analyzed for chemical and physical
properties. The soil analysis was performed by Embrapa - Bra-
zilian Agricultural Research Corporation (1979). After carefully
removing the surface organic materials and fine roots, the soil was
sieved through a 2-mm mesh screen. Soil pH (H2O), cation
exchange capacity (CEC) determined by the ammonium acetate
1N method at pH 7.0, and percent BS (base saturation) were
determined. O.M. (organic matter) was extracted according to
Walkley and Black, as described by Nelson and Sommer (1982).
Mean diameter at breast height (1.3 m) over bark (D) (BHD, cm)
and mean height of E. camaldulensis and E. grandis were calcu-
lated for each treatment. We selected three trees at random. They
were chainsawed and sampled for subsequent nutrient analysis.
Branches were separated from the trunk and all the leaves were
Journal of Forestry Research (2009) 20(1):15–22

17
collected at field. Then, total fresh weight of leaves, branches,
bark and trunk wood of the sampled trees were determined at
EPAMIG Company, Mocambinho. Sampling was done in April,
between summer and autumn, following Bellote (Personal com-
munication).
Dry mass production and nutrient concentration of Eucalypus
spp. cultivated in monoculture were evaluated at 28 months. The
growth model was based on individual trees. Trunk samples
consisted in 3.0 cm thick discs removed at each trunk, at base and
25%, 50%, 75% and 100% of height, including BHD, as proposed
by Shimoyama (1990). For each disc, the bark was separated from
the wood. This measurement served as a reference for wood
density (wd) determination. These samples (weighed after im-
mersion) were dried in a circulation oven at temperatures ranging
from 65°C, until constant weight. Finally, samples were weighed
with an analytical balance in order to obtain dry weight (dw) and
the dry biomass of the components in each tree was calculated
proportionally.
Total dry weight of leaves was obtained against composite
samples of fresh weight, by using the following equation:
SFWSDWTFWTDW /)( ×= (1)
where, TDW is the total dry weight (g), TFW the total fresh
weight (g), SDW the sample dry weight (g), and SFW is the
sample fresh weight (g).
Discs were used to calculate wood density of each subcom-
partment (bark and trunk wood), in each section of trunk, by the
hydrostatic balance method, according to Bellote (1990).
)/( IWFWDWbd −= (2)
where bd is the basic density (g/cm3), DW the dry weight, FW the
saturated fresh weight (g), and IW is the immersed weight (g).
Bark was removed from stem sections used to determine wood
density. Dry mass of the sample was determined after oven drying
for 48 h at 105°C.
Samples were taken of BHD, D5 (disc 5) and D6 sections for
determination of mineral nutrient levels at EMBRAPA-Maize and
Sorghum Soil Analyses Laboratory. Samples were dried at 75°C,
digested in nitric-perchloric mixture and analyzed for macronu-
trients. Phosphorus was analyzed by the ammonium phospho-
molybdate method. Calcium, potassium and magnesium were
analyzed by atomic absorption and potassium by flame pho-
tometry. After sulfuric digestion, Nitrogen (N) was determined by
Kjeldahl method. These procedures are in according with Sarruge
and Haag’s methodology (1974). Nutrient concentrations of D2
sample were used to calculate wood (under bark) and bark nutrient
concentrations (Young and Carpenter 1976).
Total tree volume (bark and trunk wood) was calculated by the
Smalian equation which expressed the sum of segments of length
calculated, which make up the trunk)
hdDV ×+×= )4/4/(2/1 22 ππ (3)
where, V is the log volume (m3), D the log major diameter (cm), d
the log minor diameter (cm), and h is the log height (m).
Based on basic density and volume, total dry weight (W) was
determined by using the following equations:
VbdW ×= (for bark and trunk wood) (4)
where, W is the weight (g), bd the basic density (g·cm-3), and V the
log volume (m3).
To find out tree nutrient content, the model considered differ-
ences in concentration in stem compartment, that is, in bark and
wood and also in leaves and branches. Bark volume was obtained
through the difference between volumes with bark and without
bark. The sum of leave and branches provided 12 samples, which
yielded 12 determinations of nutrient levels. Discs (18 samples)
provided 16 density determinations (2 samples were contami-
nated), besides allowing for calculating nutrient content in each
tree log and each compartment analyzed.
Macronutrient stock (kg·ha-1) in the aboveground biomass was
calculated on the basis of the dry mass estimation (kg·ha-1) and the
macronutrient concentrations (g·kg-1) obtained in the present
study. The sum of the values for each component provided the
total nutrient content (kg·ha-1) of aboveground dry mass.

Root colonization

Roots of Eucalyptus spp. were collected by excavating from the
trunk to the lateral root system of each tree. Four root samples
were harvested around the tree, and mixed together. Samples were
collected from three trees at each block and were fixed in FAA
solution (formalin:alcohol:acetic acid) until samples could be
processed. Roots were stained and assessed for mycorrhizal
infection as follows. Roots were taken from the 1/2 FAA, washed
several times in tap water and bleached in 10% (w/v) KOH
(Phillips and Hayman 1970) overnight and then heated to ap-
proximately 90ºC in a water bath for 1 h. The cooled root samples
were washed and stained with 0.05% trypan blue according to
Phillips and Hayman (1970). Roots were cut into 1-cm segments
and thirty 1-cm-root fragments were examined per sample for
their arbuscular mycorrhizal (AM) status under a compound
microscope (100 X). If at least one root segment was found to
contain fungal mycelia, arbuscules or vesicles, then the sample
was considered as an AM plant, recorded as ‘‘+’’. Plants were
recorded as non-mycorrhizal (‘‘-’’) when neither arbus-
cules/vesicles nor fungal mycelia were detected in their root
cortical cells. Quantification of mycorrhiza colonization was
according to McGonigle et al. (1990), and results were expressed
as percentage of colonized segments. Roots colonized by ecto-
mycorrhizal fungi (presence of Hartig net) were included when
calculating the percentage of root length colonized by ECM. Dual
mycorrhizae (AM and ECM) were observed and individually
recorded for calculation. These data were arcsin (x/100)1/2 trans-
formed. The data were subjected to one-way ANOVA using
MINITAB software version 13.2 and means were compared by
Tukey test (P < 0.05).

Results and discussion

The soil at the research site showed a pH ranging between 6.1 (E.
camaldulensis) and 5.8 (E. grandis) in the topsoil (Table 1).
Journal of Forestry Research (2009) 20(1):15–22

18
Textural composition presents 84% sand. Organic matter levels
were found to be low in both E. camaldulensis and E. grandis
monocultures, compared to those of foredunes (0.8%) in Brazil
(Córdoba et al. 2001). The CEC was low and the percentage of
BS (base saturation) was medium in E. camaldulensis and in E.
grandis monoculture. In general, soil presented a high content of
Zn, Cu and Fe (Table 1)，which is in accordance with other reports
that showed an improved acquisition of Zn and Cu by arbuscular
mycorrhizal (Rillig 2004).

Table 1. Chemical analysis of the soil from the sampling site of Eucalyptus camaldulensis and E. grandis monocultures inoculated with arbuscular
mycorrhizal fungi (AMF) soils at 2 years (Jaíba, Minas Gerais, Brazil)
Exchange Iron (cmol (+)·kg-1) Species pH (H2O)
1:1
Soil organic
matter
(mg·g–1)
Available. P
(mg·L-1)
Available. K
(mg·L-1) . Al3+ . Ca 2+ . Mg 2+
CEC
(cmol(+)·kg-1)
Base
saturation
(%)
E. camaldulensis 6.1 0.7 3.96 80 0 2.27 0.57 4.9 61
E. grandis 5.8 0.7 2.89 68 0.1 2.05 0.45 4.8 55
Texture (%)a Species Total
porosity
Zn
(cmol (+)·kg-1)
Cu
(cmol (+)· kg-1)
Mn
(cmol (+)· kg-1)
Fe
(cmol (+)·kg-1) Coarse sand Fine sand Clay Silt
E. camaldulensis 44.46 1 0.5 318 170 48 36 16 0
E. grandis 41.08 1.9 0.5 388 176 52 32 13 3
a Mean of two measures from one composite sample. Particle size distribution: coarse sand 2–0.2 mm, fine sand 0.2–0.02 mm, silt 0.02–0.002 mm and clay < 0.002
mm. mg L–1 = milligram per liter, CEC = cation exchange capacity.

The height (12.2 m) of E. camaldulensis observed in this study
was higher than that (10.4 m) previously reported by Bernardo et
al. (1998) in Cerrado soils in southeastern Brazil, at age 41
months, and the mean diameter was bigger than that in conven-
tional fertilization plantation (Table 2). The above-ground bio-
mass was 33.6 Mg·ha-1 for E. camaldulensis and 53.1 Mg·ha-1 for
E. grandis. Of the total above-ground biomass of E. camaldu-
lensis, the stem wood, leaves, branches, and stem bark account for
64.4%, 19.6%, 15.4%, and 0.6%, respectively (Table 2), and, a
similar partition of the total above-ground biomass was also found
for E. grandis (66.6%, 16.1%, 17.1%, and 0.2% spread in the stem
wood, leaves, branches, and stem bark, respectively). The
aboveground biomass (3.1 Mg·ha-1) of E. grandis in the present
study was higher than that obtained by Faria et al. (2002) (46.4
Mg·ha-1) at 80 months-old stands of this species in a Cerrado
region of Minas Gerais, whereas the stem volume (75.9 m3·ha-1)
was lower than that observed in Cerrado (87.9 m3·ha-1). Almeida
et al. (2007) predicted a lower stem volume (approximately 50 m3
ha-1) for E. grandis on Brazil´s Atlantic coast. Biomass distribu-
tion in descending order was: wood> leaves> branches> bark, for
E. camaldulensis and, wood> branch> leaves> bark for E. grandis
(Table 2).

Table 2. Characteristics of Eucalyptus camaldulensis and E. grandis monocultures inoculated with arbuscular mycorrhizal fungi (AMF), after 28
months of growth, at Jaíba, Minas Gerais, Brazil
Species
Height
(m)
Diameter
(cm)
Above-ground
dry mass
(Mg·ha-1)
Canopy dry
mass
(Mg·ha-1)
Leaves dry
mass (g)
Branches dry
mass (g)
Stem bark
(g)
Wood
(g)
Stem Volume
(m3·ha-1)
E. camaldulensis 12.2 (0.4) 10.5 (0.3) 33.6 (6.5)* 11.7 (3.4) 5916.7 (1208) 4645.5 (2155) 179.7 (54.4) 19475.5 (2833) 42.8 (4.4)
E. grandis 12.9 (0.8) 11.8 (0.8) 53.1 (1.4)* 17.6 (5.1) 7703.1 (1814) 8180.2 (2800) 87.3 (24.3) 31834.1(3639) 75.9 (4.9)
Dates from monocultures with 3 m × 3 m spacing. Standard deviations of means between brackets. N=3. * significant values at 0.05 level

Table 3 shows the results of the content, average amount and
ratio (percentage) of some selected micro- and macronutrients
immobilized in each component of the aboveground biomass of E.
camaldulensis and E. grandis. We observed that 33.2% of nutri-
ents were accumulated in E. camaldulensis leaves, 6.2% in wood,
21.6% in branches, and 38.8% in stem bark.
For E. camaldulensis, the contents of N, P, K, Ca, Mg, and S in
the total biomass were 122.7, 6.4, 99.8, 208.9, 16.1, and 11.2
kg·ha-1, respectively. The dry matter of leaves and branches
accounted for 35% of the total biomass, and the contents of N, P,
K, Ca, Mg, and S in leaves and branches accounted for 15.5%,
0.7%, 12.3%, 22.6%, 1.9%, and 1.4% of those in total
above-ground biomass, respectively. In the trunk (bark and wood),
which represents the remaining 65% of the total above-ground
biomass, 4.5% of N, 0.28% of P, 7.4% of K, 31.3% of Ca, 1.1% of
Mg, and 0.6% of S were accumulated. Thus the canopy (leaves
and branches) concentrates 54.73% nutrients of total aboveground
biomass. Leaves have most tree living cells that tend to accumu-
late larger quantities of nutrients, due to respiration and photo-
synthesis (Kramer and Koslowski 1979).
For E. grandis, the contents of N, P, K, Ca, Mg, and S in the
total biomass were 224.4, 10.7, 103.1, 225.4, 26.5, and 12.6
kg·ha-1, and the contents of N, P, K, Ca, Mg, and S in leaves and
branches accounted for 23.6%, 1.1% , 10.5%, 20.9%, 2.6%, and
Journal of Forestry Research (2009) 20(1):15–22

19
1.1% of those in total aboveground biomass (Table 3). In the trunk
(bark and wood), which represents the remaining 66.8% of the
total aboveground biomass, 6.3% of N, 0.34% of P, 4.7% of K,
26.4% of Ca, 1.8% of Mg, and 0.6% of S were accumulated.
The contents of N observed in our study were similar to the
results obtained by Hunter (2001), who reported an average N
content of 16 mg/g for E. grandis at 37 months and 14 mg/g for E.
camaldulensis in a mixed plantation with 2m×2 m spacing. This
corroborates other studies of nutrient contents of forests (Poggiani
et al. 1983; Boerner 1984; Pereira et al. 1984; Timmer and Mor-
row 1984, Hopmans et al. 1993). In general, the concentrations for
N varied as foliage > branches, bole bark, roots > bole wood.
Harrison et al. (2000) showed a higher N concentration in leaf and
bark in E. camaldulensis in Cerrado soils, but the same concen-
trations in branches and wood. Schumacher and Poggiani (1993)
reported higher N, P and K concentrations in leaves and highest
concentrations of Ca and Mg in bark in E. camaldulensis and E.
grandis. Dell et al. (1995) suggest that N leave concentrations for
Eucalyptus grandis × Eucalyptus urophylla must be between 18 to
29 g·kg-1.

Table 3. Results of average content, average amount and ratio of nutrients in the different components of above-ground biomass of Eucalyptus
camaldulensis and E. grandis inoculated with AMF, at Jaíba, Minas Gerais, Brazil
N P K Species cp
g·kg-1 kg·ha-1 % g·kg-1 kg·ha-1 % g·kg-1 kg·ha-1 %
E. camaldulensis Leaves 13.1 86.2 10.7 0.5 3.6 0.4 8* 52.8 6.5
Branches 5.8 30.4 4.8 0.4 2.3 0.3 7.1* 36.8 5.8
Wood 2.4 2.6 1.9 0.1 0.1 0.08 2.3 2.5 1.8
Bark 3.1 3.4 2.5 0.2 0.2 0.2 6.8 7.5 5.5
E. grandis Leaves 17.9 153.3 16.9 0.7 6.2 0.6 5.9* 50.7 5.6
Branches 6.9 63.6 6.6 0.4 4.1 0.4 5.1* 46.8 4.8
Wood 2.2 2.4 2.0 0.1 0.1 0.1 1.6 1.7 1.5
Bark 4.5 5 4.2 0.2 0.2 0.24 3.4 3.7 3.2
Ca Mg S Species cp
g·kg-1 kg·ha-1 % g·kg-1 kg·ha-1 % g·kg-1 kg·ha-1 %
E. camaldulensis Leaves 16.2 106.4 13.2 1.7 11.2 1.4 1 6.7 0.8
Branches 11.6 59.8 9.4 0.6 3.3 0.5 0.7 3.6 0.5
Wood 2.2 2.4 1.8 0.3 0.3 0.2 0.3 0.3 0.2
Bark 36.1 40.1 29.5 1.1 1.2 0.8 0.4 0.4 0.3
E. grandis Leaves 12.3 105.8 11.7 1.9 17 1.8 0.7 8.3 0.7
Branches 9.7 88.5 9.2 0.8 7.4 0.7 0.4 3.6 0.3
Wood 1.3 1.4 1.2 0.2 0.2 0.2 0.3 0.3 0.2
Bark 26.6 29.5 25.2 1.6 1.7 1.5 0.3 0.3 0.2
Cp= component of above-ground biomass; * significant values at 0.05 level

Phosphorus accumulation observed in this study was similar to
the results obtained by Hunter (2001). The concentration of P in
branches was somewhat higher than the results showed by Hunter
(2001). By other hand, Harrison et al. (2000) showed higher levels
of P for this species, which varied as foliage > branches, bole bark,
small roots > taproot, coarse roots, bole wood.
Corroborating previous studies (Drechsel and Zech 1991; Har-
rison et al. 2000), K concentrations in leaves were adequate for
satisfactory growth. For all components, E. camaldulensis had the
highest K concentration (33% higher) (Table 3).
Calcium concentration in E. grandis was similar to results
showed by Hunter (2001), whereas Ca and Mg contents in E.
camaldulensis were lower than the data showed by Hunter (2001).
Sulfur is an essential element found in plants mostly in its reduced
form in amino acids cysteine and methionine. Sulfur contents of
the leaves were slightly lower than usual values (1.9 to 3.2 g·kg-1)
proposed by Dell et al. (1995). Silveira et al. (2003) also found a
lower S concentration compared to that proposed in E. grandis
seedlings leaves. The sulfur content in the total biomass (11 to
12.5 kg·ha-1) (Table 3) was higher than that reported by Caldeira
et al. (2002) for a leguminous tree, Acacia mearnsii. Most of the
nutrients were concentrated in the leaves and bark. Similar results
(leaves) were found by Tandon et al. (1988) in Australian planta-
tions of E. grandis, Vezzani (1997) in pure and mixed stands of
Eucalyptus saligna and Acacia mearnsii and in Brazil, by George
and Varghese (1990) in E. globulus. Nutrient concentration in
leaves is influenced by diverse factors such as site conditions, age,
position of leaves and season (Van der Driessche 1984). Bellote
(1990) observed that nutrient concentration in leaves of Euca-
lyptus grandis in Brazil varies with stand age and with the season.
If E. camaldulensis canopy concentrates 54.7% of total above-
ground biomass nutrients, an exploitation system that preserves
these tree components in the site would mean that approx. 403
kg·ha-1 of these nutrients could remain in this site.
If we consider plant height alone, there was no significant dif-
ference between the two species in the monocultures, whereas, in
terms of aboveground dry mass, E. grandis performed better than
E. camaldulensis. The two species thus show a difference in
biomass partitioning and accumulation and differ in their con-
centrations of macronutrients (Table 3). To assess nutrient content
Journal of Forestry Research (2009) 20(1):15–22

20
in trees during sampling, an age of three years old must be chosen
since at this age E. grandis shows larger biomass accumulation
per time unit. This is due to three-year-old trees’ bigger capacity
to absorb soil nutrients and according to Bellote (1990) to the fact
that nutrient content in three-year-old E. grandis mature leaves
shows the highest values.
The inner variability of trees species in longitudinal sense has
more drastic effect on the chemical composition of bark than on
that of wood. Wood of basal log of the two Eucalyptus species
presented higher levels of N than the others logs. More apical logs
present the highest N and P in the bark, and the highest P in the
wood. E. camaldulensis showed the highest levels of Ca, Mg and
S, at the basal log, and for K, the highest content was in the bark of
basal log and wood of apical logs. E. grandis showed also more
Ca, Mg and S content in basal log wood but, in contrast, bark of
apical logs and wood of basal log presents more K (Table 4).

Table 4. Quantification of nutrients of Eucalyptus camaldulensis and
E. grandis inoculated with AMF, 28 months old, at Jaíba, Minas
Gerais, Brazil. (Means, N=3)
Species N (g) P(g) K(g) Ca(g) Mg(g) S(g)
Leaves 77.6† 3.3 47.6 95.8 10.1 6.06
Branches 27.4 2.1 33.1 53.9 3.01 3.3
Bark 0.14 0.01 0.26 1.62 0.05 0.02
Basal logs
Wood 11.03 0.46 11.1 10.25 1.55 1.54
Bark 0.3 0.08 0.13 0.21 0.02 0.009
E. camaldulensis
Apical logs
Wood 6.3 0.7 13.3 7.76 0.9 0.92
Leaves 138 5.6 45.7 95.3 15.3 7.49
Branches 57.2 3.7 42.14 79.7 6.6 3.26
Bark 0.07 0.004 0.06 0.42 0.028 0.005
Basal logs
Wood 18.5 0.8 13.9 11.39 2.16 1.5
Bark 0.08 0.07 0.11 0.45 0.03 0.008
E. grandis
Apical logs
Wood 10.1 0.94 11.6 6.7 1.9 1.17
†= Mean nitrogen content (g) in a plant compartment.

Potassium and Ca are nutrients that could limit productivity in
the next cycle, and this limitation can be reduced, if only wood
was harvested. To support the optimum soil calcium levels is,
therefore, of increasing interest due to the lower levels of this
nutrient present in soils cultivated with eucalypt, and to the fact
that about 58% of total calcium absorbed is exported by bark
removal.
Mycorrhizal colonization varied with Eucalyptus species. As-
sessments of percentage AM and ECM root length for E.
camaldulensis and E. grandis are shown in Table 5. The E.
camaldulensis AM root colonization (hyphae) was 15%, and
vesicles were found in the same percentage. Structures observed
suggest a Glomeraceae AM colonization (Sieverding 1991),
Glomus sp. being relevant for this species at this site (Pagano
2007, Pagano et al. 2008). These results are in accordance with
those obtained by Santos et al. (2001) where E. camaldulensis
presented the highest values of percent AM mycorrhizal coloni-
zation among several Eucalyptus species and with Ad-
joud-Sadadu and Halli-Hargas (2000) who reported a <50% AM
colonization by this same species. By other hand, levels of AM
colonization were lower to those found by Malajczuk et al. (1982)
for E. marginata. Eucalyptus presented a Hartig net (ECM)
confined to epidermal cells (Brundrett 2004).
Eucalyptus grandis did not show AM root colonization (Table
5). This may depend on the time of sampling, but ECM was
preferentially related with this species. Chen et al. (2006) showed
that rooting media greatly affected Scleroderma colonization of
Eucalyptus urophylla roots from spore inoculum in nursery
experiments. Our results of the dual colonization by ECM and
AM fungi in E. camaldulensis support the predominance of ECMs
(Lapyeyrie and Chilvers 1985, Gardner and Malajczuk 1988,
Brundrett et al. 1996, Chen et al. 2000). ECM colonization of E.
grandis was similarly to the results presented by Chen et al. (2006)
for E. urophylla (aprox. 50%), whereas AM colonization (10%)
was similar to E. camaldulensis AM colonization shown in this
work. Sustained levels of root colonization above 50% have been
mentioned as necessary to ensure high survival and productivity
of plantation (Marx et al. 1989) accessing sufficient N and P.
Ectomycorrhizal fungi are common in eucalypt plantations, being
ecologically important in nutrient cycling (Högberg and Högberg
2002, Read and Perez-Moreno 2003). Our observations of abun-
dant basidiocarps (probably Pisolithus), especially in E. grandis
plots, suggest that ECM may enable uptake of immobile phos-
phorus and other nutrients as has been reported by other refer-
ences (Smith and Read 1997) reflecting this in plant nutrient
content. Thus, the higher nutrient content (especially N and P) in
E. grandis biomass in this study may be explained by the presence
of ECM. On the other hand, AMF have an ecological and agro-
nomic importance in the tropics and their presence can be influ-
enced by environmental factors such as: climate conditions,
physical proprieties, soil chemical and physical proprieties, host
vegetal species and their age and variety (Cardoso and Kuyper
2006).

Table 5. Mycorrhizal colonization in the homogeneous plantations of
24-month-old Eucalyptus camaldulensis and Eucalyptus grandis
inoculated with AMF at Jaíba, Minas Gerais, Brazil, in the dry period
Host plant species Colonization
Eucalyptus camaldulensis Eucalyptus grandis
AM hyphae 15† a 0 b
AM vesicles 15 a 0 b
ECM 26.66 a b 50 ba

†% colonization; values followed by the same lowercase letter (row) do not
differ at P<0.05 as determined by Tukey 5%; AM= arbuscular mycorrhiza;
ECM= ectomycorrhiza.

Conclusion

According to the previously described results on Eucalyptus spp.,
it can be concluded that total productivity was of the order E.
grandis > E. camaldulensis and that these species did not reduce
growth and aboveground biomass production when cultivated at
this site, showing nutrient concentrations similar to that informed
in the literature. The highest nutrient concentrations were found in
the leaves and the lowest concentrations of N, P, K, Ca and Mg
were found in the stem wood, those of S being in the stem wood
and bark; wood localized in superior parts of trunk presented
Journal of Forestry Research (2009) 20(1):15–22

21
higher concentration of P and bark contained significant amounts
of nutrients, especially in E. grandis. This points the importance
of leaving vegetal slash (mostly crown) on the site in order to
decrease the loss of tree productivity in this semiarid region. This
work is an approach towards studding Eucalyptus in the Southern
Brazilian region; further research is necessary, especially re-
garding litter accumulation, below-ground biomass, and nutrient
dynamics.

Acknowledgements
The authors are grateful to CAPES (Coordenação de Aper-
feiçoamento de Pessoal de Nível Superior) for scholarships
granted to Marcela C. Pagano. We would also like to thank
EMBRAPA – Maize and Sorghum - and EPAMIG – Mocambinho
for technical support; and Dr Adriana S. Pagano (Federal Uni-
versity of Minas Gerais) for proofreading of the manuscript.

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