Forest Ecology and Management 236 (2006) 286294

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Phenology of Caryocar brasiliense in the Brazilian cerrado region

Germano L.D. Leite a,*, Ronnie Von dos S. Veloso a, Jose C. Zanuncio b,
Luiz Arnaldo Fernandes a, Chrystian I.M. Almeida c
a

Setor de Fitotecnia, Nucleo de Ciencias Agrarias (NCA), UFMG, Av. Osmane Barbosa, s/n, B. JK, Cx. Postal 135,
CEP 39404-006 Montes Claros, Estado de Minas Gerais, Brazil
b
Departamento de Biologia Animal, Setor de Entomologia, UFV, CEP 36571-000 Vicosa, Estado de Minas Gerais, Brazil
c
Departamento de Horticultura, Faculdades de Ciencias Agrarias, UNESP-Botucatu, Botucatu, CEP 18.630-970 Estado de Sao Paulo, Brazil
Received 26 August 2005; received in revised form 13 September 2006; accepted 13 September 2006

Abstract
The objective was to study the phenology of Caryocar brasiliense Camb. (Caryocaraceae), a type of fruit of the Brazilian cerrado, in three areas
(cerrado, pasture and in the campus of the Federal University of Minas Gerais) of the municipality of Montes Claros, State of Minas Gerais, Brazil,
during 3 years. The plants of C. brasiliense presented linear growth with about 50 cm of height and 35 cm of crown width per year. Soils with loamy
texture, dystrophic, rich in aluminum and with low pH favor the production of this species. C. brasiliense produced higher quantity of flowers in the
middle third, followed by the apical and basal parts of the crown, while the fruit production was higher in the basal third, followed by the middle and
apical parts of the canopy of this species. C. brasiliense bloomed during the dry period and produced fruits in the rainy season. Its fruits presented
about 6, 8 and 6 cm of width, length and height, respectively, and 160 g of gross weight. The extrativism without control has strong impact on the
propagation of C. brasiliense because only about 7.96 and 10.65% of plants up to 1.0 m height and 59.58 and 44.73% higher than 3.0 m
(reproductive phase) was observed in the cerrado and pasture, respectively. This indicates that fruit collectors, practically, remove all fruits of this
tree and thus they reduce, considerably, the propagation of C. brasiliense in the cerrado areas of Brazil.
# 2006 Elsevier B.V. All rights reserved.
Keywords: Phenology; Caryocar brasiliense; Conservation

Cerrado, savannah-like vegetation, has many physiognomic

forms, ranging from the tall cerradao (trees to 20 m height
which is similar to woodland savannah of other continents), a
more common cerrado sensu stricto (a species-rich dense scrub
of shrubs and trees, 810 m height, with a dense understory) to
the grassland open forms (Miranda et al., 1997; Durigan et al.,
2002; Fidelis and Godoy, 2003).
The cerrado occupies about 21% of the Brazilian territory
with a great fauna and flora diversity (Aguiar and Camargo,
2004). The cerrado represents the main area of grain
producing and cattle farm in Brazil. Only 20% of cerrado
area is considered original or with low disturbance, which
means that 80% of the Brazilian cerrado was converted to
antropic areas (Aguiar and Camargo, 2004). The percentage
* Corresponding author.
E-mail addresses: gldleite@ufmg.br (G.L.D. Leite),
ronnievond@yahoo.com.br (R. Von dos S. Veloso), zanuncio@ufv.br
(J.C. Zanuncio), larnaldo@ufmg.br (L.A. Fernandes), iezid@fca.unesp.br
(C.I.M. Almeida).
0378-1127/$ see front matter # 2006 Elsevier B.V. All rights reserved.
doi:10.1016/j.foreco.2006.09.013

of the cerrado that is protected by law is, only, 4.40% of this

area and is not included among the biomass considered, by
the Federal Constitution of Brazil, as a national patrimony.
The scientific knowledge on the cerrado is still very
incipient, but it is possible to estimate its species richness
in at least 30% of the total for Brazil. However, the Brazilian
cerrado is being destroyed at a higher speed than the capacity
of the scientific community to produce the necessary
knowledge for its protection and conservation (Aguiar and
Camargo, 2004).
The fire, sazonality of raining and poor and acid soils are
determinants for the cerrado vegetation (Miranda et al., 1997;
Pinto et al., 2002; Fidelis and Godoy, 2003). The surface fire in
the cerrado is of short duration and it basically consumes the
fine fuel of herbaceous plants (Aguiar and Camargo, 2004), but
successive fires can reduce new sprouting and the plants do not
reach a minimum thickness of 68 mm of bark that causes
tissue death (Aguiar and Camargo, 2004). Besides, the
intensive fire causes repellence water to soil in the pasture,
that affects nutrient cycling (Johnson et al., 2005).

G.L.D. Leite et al. / Forest Ecology and Management 236 (2006) 286294

The Caryocar brasiliense Camb. (Caryocaraceae) (common

names: pequi, piqui, piquia or piqui-do-cerrado) is present in a
wide distribution in the Brazilian cerrado (Brandao and
Gavilanes, 1992; Almeida et al., 1998). This species presented
1542 plants/ha in the cerrado of the Brazilian Federal District,
but it can reach 180 plants/ha (Almeida et al., 1998) or 43
plants/ha spaced 15 m from each other (Almeida et al., 1998).
Its fruits are used for food, production of cosmetics and
lubricants and in the pharmaceutical industry (Araujo, 1995;
Passos et al., 2003). The oil of these fruits has invigorating
effect, thus being used for bronchitis, influenzas, colds and
against tumors, and the tea of its leaves is a menstrual regulator
(Almeida et al., 1998). C. brasiliense is one of the most cut trees
used to produce charcoal due to its diameter and good
combustion value (Vale and Felfili, 2005).
The flowering of trees of C. brasiliense occurs after the fall
of their old leaves and the producing of new sprouts from
August (dry station) (Almeida et al., 1998; Macedo and Veloso,
2002). Its flowers are important as food for Agouti paca (L.)
(Rodentia: Agoutidae) and Mazama gouazoupira (G. Fischer)
(Artiodactyla: Cervidae), and bats, bees and moths pollinate
them (Oliveira, 1997; Macedo and Veloso, 2002). The
dispersion of its fruits is made by Didelphis albiventris (Lund)
(Mammalia: Marsupialia) and Cyanocorax cristatellus
(Temminck) (Passeriformes: Corvidae) (Almeida et al., 1998).
The multiple uses of the C. brasiliense plant in the cerrado
makes it the main source of income for many communities in
the cerrado areas of Brazil (Ribeiro, 2000), but the economical
potential of this species is, still, poorly known (Araujo, 1995).
This makes it necessary to develop research to preserve C.
brasiliense in natural conditions, to establish commercial

287

Fig. 1. Height and width increment (m) of the crown of Caryocar brasiliense in
the campus of the NCA/UFMG, municipality of Montes Claros and State of
Minas Gerais, Brazil. The symbols represent the averages and the bars the
standard errors.

plantations and to study the phenology of this specie. It is

important to develop studies on the phenology of C. brasiliense,
to plan the rational crop of its fruits by the communities and to
protect native forest species (Falcao et al., 2003), to predict the
emergency and to control pests (Herms, 2002), to plan refuge
areas, to attract beneficial insects and to enhance natural
biological control, besides protecting bees and to predict global
warming trends. Unfortunately, the few publications about the

Table 1
Data of the physical and chemical analyses of the soil during the experimental period in the three areas in the municipality of Montes Claros, State of Minas Gerais,
Brazil (sample size = 108) (average  standard deviation)
Parameters of the soil

Cerrado

Pasture

NCA/UFMG

pH in water
Phosphorus-Mehlich 1 (mg dm3)
Phosphorus-remaining (mg L1)
Potassium (mg dm3)
Calcium (cmolc dm3)
Magnesium (cmolc dm3)
Aluminum (cmolc dm3)
H + Al (cmolc dm3)
Sum of bases (cmolc dm3)
t (cmolc dm3) a
m (%)a
T (cmolc dm3) a
V (%) a
Organic matter (dag kg1)
Gross sand (dag kg1)b
Fine sand (dag kg1)
Silt (dag kg1)
Clay (dag kg1)
Texture

4.85  0.04 B (Bx)

0.80  0.10n.s. (MBx)
40.76  1.16 A
28.25  2.92 C (Bx)
0.20  0.01 B (MBx)
0.10  0.01 C (MBx)
0.68  0.06 B (M)
5.19  0.19 B (M)
0.37  0.01 B (MBx)
1.05  0.07 C (Bx)
63.58  1.61 A (A)
5.56  0.19 b M
6.66  0.30 B (MBx)
1.11  0.01 B (Bx)
20.92  0.54 A
53.92  2.44 A
10.83  1.16 C
14.33  1.20 C
Sandy

4.87  0.19 B (Bx)

0.59  0.33n.s (MBx).
17.64  1.42 C
62.92  7.00 B (M)
0.71  0.55 B (Bx)
0.37  0.09 B (Bx)
1.06  0.14 A (M)
10.93  0.71 A (MA)
1.23  0.59 B (Bx)
2.30  0.46 B (Bx)
47.75  2.53 B (M)
12.17  0.22 A (B)
11.08  5.56 B MBx
8.77  0.22 A (B)
5.75  0.90 C
30.33  1.34 B
24.83  2.24 B
39.00  1.14 A
Loamier

6.74  0.07 A (A)

1.07  0.11n.s. (MBx)
31.17  0.61 B
101.50  5.99 A (B)
6.85  0.11 A (MB)
1.53  0.06 A (MB)
0.22  0.08 C (MBx)
2.79  0.89 B (M)
8.63  0.17 A (MB)
8.85  0.10 A (MB)
3.75  4.49 C (Bx)
11.42  0.84 A (B)
76.33  1.74 A (B)
1.99  2.72 B (Bx)
8.83  0.21 B
25.00  1.55 B
41.33  0.83 A
24.83  1.48 B
Average texture

Means followed by the same letter (AC) in the line do not differ between them by the test of Tukey at 1% probability. MBx = very low, Bx = low, B = good,
M = mean, A = high, MB = very good and MA = too high.
a
t; capacity of cationic exchange, m; aluminum saturation in the capacity of cationic exchange, T; cation exchange capacity at natural pH 7.0 and V; percentage of
soil base saturation of the capacity of cationic exchange a pH 7.0.
b
Gross sand (2  0.2 mm) (dag kg1), fine sand (0.2  0.02 mm) (dag kg1), silt (0.02  0.002 mm) (dag kg1) and clay (<0.002 mm) (dag kg1).

288

G.L.D. Leite et al. / Forest Ecology and Management 236 (2006) 286294

phenology of C. brasiliense did not include different soil types

of the cerrado (Almeida et al., 1998; Macedo and Veloso,
2002).
The objective was to study the phenology of C. brasiliense in
three areas (cerrado, pasture and in the campus of the Nucleo de
Ciencias Agrarias da Universidade Federal de Minas Gerais
(NCA/UFMG) of the municipality of Montes Claros, State of
Minas Gerais, Brazil).
1. Material and methods
This work was developed in the municipality of Montes
Claros, State of Minas Gerais, Brazil, from June 2001 to June
2004 in a region with tropical savanna climate Aw, according to
the classification of Koppen, with dry winter and rainy summer,
in three areas: (1) cerrado vegetation strictu sensu anthropomorphized, longitude 438550 7.300 W, latitude 168440 55.600 S,
altitude 943 m and dystrophic yellow red latosoil of sandy

texture; (2) pasture (originally cerrado vegetation), longitude

438570 31.400 W, latitude 168460 16.100 S, altitude 940 m and red
dystrophic yellow latosoil of loamy texture; (3) campus of the
NCA/UFMG, longitude 438500 26.800 W, latitude 168400 54.500 S,
altitude 633 m and dystrophic red latosoil of medium texture.
The chemical and physical characteristics of the soils of the
three areas are presented in Table 1.
The number of flowers per bunch and per tree of C. brasiliense
was evaluated monthly by categories to its development,
F1 = closed floral button; F1A = closed floral button with
chalice-corolla developed; F1B = opened flower, and F1C = fecundated flower without corolla. The number of fruits/tree was
evaluated in 10 plants in the cerrado, native plants with height of
4.07  0.18 m AB (Tukeys test, sample size = 900) and crown
width 2.87  0.13 m AB; and 10 in the pasture, native plants with
height of 5.20  0.18 m A and crown width 3.96  0.14 m A;
besides in five others in the NCA/UFMG, planted in these area
with height of 3.79  0.15 m B and crown width 1.66  0.13 m

Fig. 2. Flower production in different stages of Caryocar brasiliense in the three areas from June 2001 to June 2004. The symbols represent the averages and the bars
the standard errors. Means followed by the same letter do not differ between them by the test of Tukey at 5% probability. Development of flowers of C. brasiliense:
FI = closed floral button, F1A = closed floral button with chalice-corolla developed, FIB = opened flower and F1C = fecundated flower without corolla.

G.L.D. Leite et al. / Forest Ecology and Management 236 (2006) 286294

B. These last trees produced fruit for the first time in 2003 when
they were 8 years old. The number of flowers was counted per
category and that of fruits in a bunch per side of the crown (north,
south, east and west) and along the canopy (apical, middle and
basal) of 25 trees. The total production of flowers and fruits/tree
was obtained by multiplying the total number of bunches per tree
by the number of flowers and fruits per bunch. Adult trees of C.
brasiliense (producing fruits) were sampled randomly in each
collection, except in the campus of the NCA/UFMG where only
five trees were found. The evaluation of height and crown width
increment was made monthly in the five trees of the campus of
the NCA with a tape measurer. This was made because the five
trees were used every time and they were in a transition period
between the vegetative and reproductive stages. The width,
length and height of 10 fruits were obtained with a tape measurer,
and their weight with a digital scale with three decimal precisions
from 25 trees in each evaluation. These fruits were collected and
transported to the laboratory of the NCA/UFMG for evaluations.
No fire was registered during the experimental period in any of
the tree areas.
The propagation of C. brasiliense was evaluated under
extrativism conditions. The number of plants of this species in the
cerrado and in the pasture was evaluated in the last month (June of
2004) in three areas of a hectare and divided in the following
height categories (m): (1) up to 1.0; (2) from 1.0 to 1.5; (3) from 1.5
to 3.0 (vegetative period) and (4) above 3.0 (reproductive period).

289

The climatic data (temperature, rainfall, relative humidity of

the air, sunshine, direction and wind speed) were obtained from
the main climate station of Montes Claros of the Estacao
Nacional de Metereologia (longitude: 438500 W, latitude:
168410 S and altitude: 646.29 m). A total of 108 soil samples
(020 cm deep) were collected, being 36 in each area (3
samples per season), to evaluate the physical and chemical
characteristics of these samples in the Laboratory of Analysis
of Soils of the NCA/UFMG, according to the methodology of
the EMBRAPA (1997). The samples were collected under the
projection of the canopy of the C. brasiliense trees. The data
were submitted for the analysis of variance (ANOVA)
(P < 0.05), for Tukey (P < 0.05) tests, for a Pearson
correlation (P < 0.05) and/or for a multiple regression analysis
(P < 0.05). The parameters used in the multiple equations were
those significant (P < 0.05) for selection of the variables for the
method Stepwise.
2. Results
The plants of C. brasiliense of the campus of NCA/UFMG
presented linear growth with about 50 cm in height and 35 cm
of crown width per year (Fig. 1). The production of flowers (all
three study areas) of this specie was started in July and August,
with a maximum in September 2002 and 2003 in Montes
Claros, State of Minas Gerais, Brazil (Fig. 2). C. brasiliense

Fig. 3. Total flower production per tree and fruits per branch and total per tree of Caryocar brasiliense in the tree areas from June 2001 to June 2004, municipality of
Montes Claros, State of Minas Gerais, Brazil. The symbols represent the averages and the bars the standard errors. Means followed by the same letter do not differ
between them by the test of Tukey at 5% probability.

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G.L.D. Leite et al. / Forest Ecology and Management 236 (2006) 286294

started to produce fruits (all three study areas) in October 2001

and 2002 and in September and October 2003, with production
peaks in November 2001 and 2002 and October 2003, during
the beginning of the rainy season in the area (Fig. 3; Tables 2
and 3). The fruit production of C. brasiliense lasted for 45
months. The flower production of this plant increased with the
temperature during the dry period when lower relative humidity
and higher sunshine were recorded. Later, the fruit production
increases as a function of rainfall (Table 3).
Plants of C. brasiliense produced larger quantity of fruits per
bunch in the pasture and in the cerrado than in the NCA/UFMG,
but that of flowers per bunch and flowers and fruits per tree was
bigger in the pasture than in the cerrado and in the NCA/UFMG
(Figs. 2 and 3). The largeness of fruits of C. brasiliense was

similar between the three areas: width 6.19  0.20 cm, length
7.71  0.24 cm, height of 6.43  0.15 cm, volume of
314.90  20.93 cm3 and a gross weight of 158.49  8.14 g
with a minimum and maximum of 84 and 270 g, respectively.
The production of flowers/branch of C. brasiliense was
higher in the west face than in other parts of the crown of this
species in the pasture (Table 4) but that of fruits/bunch was
higher in the west, followed by the north, south and east faces
(Table 4). The total number of flowers per bunches was higher
in the middle third, followed by the apical and basal parts of the
crown but that of fruits was higher in the basal third, followed
by the middle and apical parts of the crown of C. brasiliense in
the pasture. This species had higher number of flowers in the
middle and apical thirds, but the number of fruits was higher in

Table 2
Average temperature (8C), total rainfall (mm), relative humidity of the air (%), sunshine (h) and direction and speed (m/s) of the wind in the municipality of Montes
Claros, State of Minas Gerais, Brazil
Experimental period

Average temperature

Total rainfall

Relative humidity

Sunshine

Wind
Direction

Speed

2001
June
July
August
September
October
November
December

23.5
20.7
21.5
23.9
23.8
23.7
23.8

0.0
0.6
0.0
2.4
166.6
333.0
164.1

60
56
48
50
63
78
72

8.8
8.1
8.8
8.1
5.8
4.6
3.8

East
Northeast
Southeast
East
East
Northeast
Northeast

1.9
1.9
2.8
2.6
2.4
1.8
2

2002
January
February
March
April
May
June
July
August
September
October
November
December

24.2
24.0
24.3
23.7
22.5
21.1
21.4
22.8
23.5
25.4
24.5
24.8

199.5
135.2
57.3
7.1
1.7
1.3
0.0
0.4
117.3
34.4
147.9
400.4

76
78
70
65
60
59
56
45
56
48
65
72

6.4
5.4
8.2
9.1
8.7
8.9
8.3
10.3
7.2
9.6
7.1
5.0

Northeast
Northeast
Northeast
Northeast
East
Northeast
Northeast
Northeast
Northeast
Northeast
Northeast
Northeast

2.2
1.7
2.1
1.9
1.9
1.8
1.9
2.6
2.4
2.2
2.2
1.9

2003
January
February
March
April
May
June
July
August
September
October
November
December

24.0
24.8
24.9
24.6
22.4
20.9
20.3
22.2
24.3
25
25.6
25.6

271.8
25.8
39.3
4.3
5.4
0.0
0.0
10.2
0.0
54.0
56.2
204.7

81
64
69
62
60
54
52
54
62
47
58
63

4.6
8.8
6.4
8.6
8.2
9.8
9.1
7.9
7.9
7.9
7.1
7.5

Northeast
Northeast
Northeast
Northeast
Northeast
Northeast
Northeast
Northeast
Northeast
Northeast
Northeast
Northeast

1.7
6.4
1.9
1.9
1.9
1.6
1.9
1.9
2.3
2.3
2.3
2.2

2004
January
February
March
April
May
June

24.0
23.3
23.0
22.6
21.9
19.4

277.3
315.1
184.7
60.8
4.0
26.2

78.0
83.0
82.0
78.0
74.0
72.0

4.1
4.7
6.1
6.6
8.1
7.4

Northeast
Northeast
Northeast
Northeast
Northeast
East

2.1
2.0
1.8
1.7
1.5
1.7

National Metereological Station of the campus of the NCA/UFMG.

G.L.D. Leite et al. / Forest Ecology and Management 236 (2006) 286294
Table 3
Correlation of Pearson and equations of multiple regressions of the variables in
three areas of the municipality of Montes Claros, State of Minas Gerais, Brazil
Correlation of Pearson
Cerrado
F1 with number bunches of flowers/tree
Total flowers/tree with the number of
bunches with flowers/tree
Total flowers/bunch with temperature
Total flowers/bunch with relative
humidity of the air
Total fruits/tree with the number of
total bunches of fruits/tree
Total fruits/bunch with temperature
Pasture
Total flowers/tree with the number of
bunches with flowers/tree
Total flowers/bunch with rainfall
Total flowers/bunch with relative
humidity of the air
Total flowers/bunch with sunshine
Total fruits/tree with the number of
total bunches of fruits/tree
Total fruits/tree with the number
of fruits/bunch
Total fruits/tree with rainfall

Value of r
0.92
0.93

0.0416
0.0352

0.46
0.40

0.0029
0.0080

0.85

0.0354

0.55

0.0005

0.99

0.0000

0.29
0.39

0.0400
0.0096

0.28
0.75

0.0487
0.0407

0.87

0.0001

0.87

0.0000

0.26
0.39
0.64

0.0587
0.0094
0.0093

1.00

0.0000

0.64

0.0093

0.43

0.0405

Equations of the analysis

of the multiple regression
Cerrado
F1B = 0.20 + 0.12F1 + 0.02rainfall
F1C = 0.25 + 6.65F1A + 0.02rainfall
Fruits/bunch = 0.72 + 25.37aluminum + 1.41pH
+ 0.60F1A + 0.04rainfall  0.002
capacity of cationic exchange
Total fruits/tree = 0.0002 + 561.50aluminum
+ 0.75rainfall  510.75t  15.10wind speed

Campus of the NCA/UFMG

Insufficient number of data

its basal third due to lower incidence of wind currents in the

pasture (Tables 3 and 4). The wind direction reduced the
production of flowers and fruits of C. brasiliense in the pasture,
and its speed also affected the plants in the cerrado (Table 3).
The removal of fruits without control reduced the renewal of
C. brasiliense in the cerrado area and in the pasture because
only about 9% of seedlings of this species up to 1.0 m height
were found in these areas (Table 5).
3. Discussion

Campus of the NCA/UFMG

Total flowers/bunch with temperature
Total fruits/bunch with temperature
Total fruits/bunch with the number
of flowers/bunch
Total fruits/tree with the number
of fruits/bunch
Total fruits/tree with the number
of flowers/bunch
Total fruits/tree with the number
of flowers/tree

Pasture
F1 = 0.004 + 8.58sum of bases  7.86height
of the crown  6.55wind speed
F1A = 0.03 + 0.38sum of
bases  0.20wind speed
F1B = 3.47 + 1.67t + 0.06F1
+ 0.03temperature  0.47wind speed
F1C = 4.53 + 1.94t + 0.05temperature
+ 0.03F1  0.58wind speed
Fruits/bunch = 0.17 + 0.65crown
width + 0.04rainfall

Value of P

291

R2

0.80
0.95
0.98

0.0003
0.0000
0.0000

0.97

0.0000

0.64

0.0344

0.79

0.0011

0.96

0.0000

0.92

0.0005

0.62

0.0076

Development of flowers of C. brasiliense: F1 = closed floral button,

F1A = closed floral button with chalice-corolla developed, F1B = opened
flower and F1C = fecundated flower without corolla.

C. brasiliense flowered at the beginning of the dry season.

This was also observed for this species in an area 70 km distant
from Belo Horizonte, State of Minas Gerais, Brazil where it
bloomed from August to October, produced fruit from October
to February with a vegetative period during the rest of the year
(Macedo and Veloso, 2002), and in an area of the Federal
District, Brazil where C. brasiliense flowered after the emission
of new leaves from June to October (before the raining period)
and presented ripe fruits in December and January (Almeida
et al., 1998). The rainfall improved the fruit production of this
species, probably because it flourishes at the beginning of the
rainy period. This can be a reflex of the raining season of the
previous year, because the Brazilian cerrado is an excellent
underground reservoir of raining water (Aguiar and Camargo,
2004; Junior and Haridasan, 2005).
The sprouting, development of leaves and flowers before the
rainy period is common for perennial plants of the cerrado areas
(Almeida et al., 1998, 2006; Felfini et al., 1999). This allows the
plants to have the photosynthetic areas developed up to the
rainy period, which increases its efficiency, and the predation
by insects is lower due to the adverse conditions before the rains
start. Besides there are no heavy raining yet, and the low
quantity of leaves facilitate the pollinators to find the flowers
(Felfini et al., 1999). The direct effect of the soil water deficit on
the ecosystem surface conductance during the dry season in the
cerrado is not very high. In fact, the woody-layer plants have
deep roots to reach perpetually humid layers in the soil
(Miranda et al., 1997). The seasonality in the physiology and
morphology of the ecosystem was also reflected in the changes
of the energy balance in the surface. A smaller fraction of the
energy absorbed was partitioned to latent heat in the dry season
at high solar irradiance (Miranda et al., 1997). This saturation
response of the latent heat flux to incoming solar radiation
consisted of stomatal closure by surviving foliage, in response
to the high vapour pressure deficits prevailing in the dry season
(Miranda et al., 1997).
The height, number of flowers and fruits and the weight of
fruits of C. brasiliense were similar to that reported for plants of
this species with 7 m height. They had bunches with 1030
flowers, 50250 g fruits and with an average weight of 120 g
(Almeida et al., 1998). The number of fruits per bunch was
similar for C. brasiliense in the pasture and in the cerrado, but
the total fruit production per tree was higher in the first area
where its plants were higher and with larger crown than those in
the cerrado and in the NCA/UFMG. The plants of C. brasiliense
in the cerrado were also lower and more shaded than those in

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Table 4
Distribution of flowers and fruits in the crown of Caryocar brasiliense in the pasture according to the side exposed to the sun (north, south, east and west) or in the
height of this species (apical, middle or basal)
Exposition to the sun a

Number of flowers or fruits/bunch

Flowers/bunch
Fruits/bunch

North

South

West

East

6.12  1.06 B
2.71  0.25 AB

5.90  0.94 B
2.50  0.31 B

8.14  0.58 A
3.28  0.22 A

6.20  0.92 B
2.56  0.33 B

Cannopy of Caryocar brasiliense

Flowers/bunch
Fruits/bunch

Apical

Middle

Basal

9.34  0.94 AB
2.58  0.21 B

11.34  1.03 A
4.13  0.44 AB

5.66  1.34 B
5.34  0.25 A

Municipality of Montes Claros, State of Minas Gerais, Brazil. (Sample size = n) (Average  standard deviation).
Means followed by the same letter (a and b) in the line do not differ between them by the test of Tukey at 5% probability.
a
Sample size on the side exposed to the sun = 1080 and on the canopy = 1440.

the NCA/UFMG, which were 8 years old at the first year of

production. The soil of the area with pasture (originally cerrado
vegetation) was loamier than that of the campus of NCA/
UFMG (silt and compressed texture) and of the cerrado (sandy
texture) (Table 1). Soils of the cerrado are, in general, deep and
loamy (excellent storage capacity of rain water), poor in
nutrients, rich in aluminum and acid pH (Sousa and Lobato,
2004; Junior and Haridasan, 2005), as also found for the soil in
the pasture which had higher levels of aluminum and lower pH
values and sum of bases (calcium, potassium and magnesium)
but not for that of the campus of the NCA/UFMG (Table 1). The
C. brasiliense, a typical tree of the Brazilian cerrado, had to
adapt to these conditions. This species produced larger number
of fruits in the cerrado, where the soils presented higher levels
of aluminum and higher pH besides low capacity of cationic
exchange (Tables 1 and 3). The higher flower production in the
Table 5
Number of Caryocar brasiliense with different height (m) and percent of each
group per height in the cerrado and in the pasture
Numbers of Caryocar
brasiliense/ha

Cerrado

Pasture

Up to 1.00 m
From 1.00 to 1.50 m
From 1.50 to 3. 0 m
Higher than 3.0 m
Total number of C. brasiliense
plants

2.33  0.65 BCn.s.

1.00  1.00 Cb
5.00  1.5 ABn.s.
8.67  2.05 Ab
17.00  3.85 b

4.00  0.80
9.67  2.35
8.00  2.98
20.67  6.32
42.33  9.05

C
Ba
B
Aa
a

Percentage of Caryocar
brasiliense

Cerrado

Pasture

Up to 1.00 m
From 1.00 to 1.50 m
From 1.50 to 3.0 m
Higher than 3.0 m

7.96  2.23 Cn.s.

3.41  3.41 Cb
29.06  8.71 Bn.s.
59.58  14.09 An.s.

10.65  2.13 C
23.31  5.66 Ba
21.31  7.93 B
44.73  13.67 A

Municipality of Montes Claros, State of Minas Gerais, Brazil. (Sample

size = 60) (Average  standard deviation).
Means followed by the same capital letter (AC) in the column or small letters
(a and b) in the line do not differ between them by the test of Tukey at 5%
probability. n.s. = non significant.
Data were not collected in the NCA/UFMG because this is a green area where
the Caryocar brasiliense does not grow.

pasture can be due to a higher sum of bases and a more effective

change capacity of cations of the soil in this area (Table 3).
Higher levels of nitrogen, potassium, calcium, magnesium and
effective cation exchange capacity were also observed in the
pasture than in the soils under the cerrado vegetation, and the
level of phosphorous did not differ between the areas studied
(Lilienfein et al., 2003).
Lower production of fruits in the east and north faces in the
pasture could be due to a higher flower drop in these sides,
because the wind direction predominated from northeast to
east. Besides, the period of blooming (August and September)
of C. brasiliense occurs during the period of stronger winds in
the area. On the contrary, the lack of impact of the wind in the
flower and fruit production of plants in the campus of NCA/
UFMG can be due to the fact that these trees were planted at
lower altitude and closer, 1020 m, to the main building of the
campus that would reduce such effect. This can also be applied
to C. brasiliense plants in the cerrado where the bush and
nearby trees might have reduced the impact of wind on these
trees. Besides, the wind speed had larger impact on flower
production of this species in the pasture because this area had
higher altitude than the campus of the NCA/UFMG, and its
trees are less protected than in the cerrado. Smaller trees and
those with wider crowns (basal leaves of the cannopy) produced
higher number of flowers and fruits, respectively, because they
are better protected from wind (Table 3). Strong winds can
reduce the production of fruits due to the desiccation of leaves,
thus reducing photosynthesis and causing bad formation of
flowers and fruits or even the fall of flowers and of fruits. For
this reason the use of windbreaks can reduce these damages
(Wakefield, 1989; Mowat and George, 1996; Castellani and
dEca-Neves, 2000; Wight and Stuhr, 2002; FAO, 2006).
Besides, the wind speed over 2.0 m/s reduces the visits of bees
to the flowers of the plants as the speed affects their
aerodynamics (Dutra and Machado, 2001). However, we
observed low density of, only, Trigona spinipes Fabr.
(Hymenoptera: Apidae) damaging the flowers of C. brasiliense
(data not presented). We cannot discard the hypothesis that the
wind speed might have negatively affected the night pollinators
of this plant, mainly because the wind speed was over 2.0 m/s,

G.L.D. Leite et al. / Forest Ecology and Management 236 (2006) 286294

thus resulting in lower fruit production in the parts of the plant

more exposed to the wind.
The lower propagation of C. brasiliense in the pasture and
cerrado indicates that the collectors remove majority of its
fruits. On the contary, the largest number of plants, from 1.0 to
1.5 m (vegetative stage), in the pasture (23%) than in the
cerrado (3.41%) can be due to the largest shading and worse soil
conditions (physicochemical) in the second area, thus resulting
in lower germination rate and lower fruit production. The
highest number of trees with wider crowns (reproductive stage)
indicates that the populations of C. brasiliense can be reduced
and this species can disappear from areas of the Brazilian
cerrado. The area of the pasture presents an ideal density of C.
brasiliense, with around 42 trees/ha (Almeida et al., 1998).
However, the size of most plants was about the size to be fed by
bovine cattle (up to 2 m) (Almeida et al., 1998). Therefore, they
are constantly damaged. On the other hand, the cerrado area,
that is an area of permanent conservation, presents an inferior
density (17) of plants than the ideal per hectare. The area with
cerrado vegetation strictu sensu anthropomorphized, and the
pasture area (private property = farm) do not present the fruit
disperseres of C. brasiliense, D. albiventris and C. cristatellus
(Almeida et al., 1998) that make the situation of this plant more
critical in these areas. The involvement of the local
communities that extract the fruits from C. brasiliense through
environmental education represents one of the most promising
way to conserve the natural resources and to protect natural
areas (Aguiar and Camargo, 2004), besides the law. The cerrado
has been the main source of firewood and charcoal in Brazil,
and neither plantations of native species nor sustainable
management of them have been adopted so far in the region. For
this reason, the plantation of native species could help to
conserve these species and to use management strategies (Vale
and Felfili, 2005).
C. brasiliense blooms in the dry period and produces fruits
during the raining season in the north of the State of Minas
Gerais, Brazil. This makes it necessary to use windbreaks to
reduce the fall of their floral buttons, flowers and fruits and to
increase the production of this species. Besides, plants of C.
brasiliense should be maintained in pastures as shading areas
for the cattle, fruit production and as an alternative source of
income for farmers and collectors of fruits of C. brasiliense.
Finally, the cerrado should be conserved because it represents
the natural bank of germoplasm of this species. It is also
recommended to develop researches on education and crop
planning with rural communities to preserve C. brasiliense in
cerrado areas of the State of Minas Gerais, Brazil.
Acknowledgements
The author acknowledges Mr. Celio Barbosa, Oscar
Madureira da Silva and Fabrcio Maurlio Ruas for supplying
climatic data and the Brazilian agencies Fundacao de Amparo
a` Pesquisa do Estado de Minas Gerais (FAPEMIG),
Conselho Nacional de Desenvolvimento Cientfico e Tecnologico (CNPq) and Secretaria de Ciencia e Tecnologia do
Estado de Minas Gerais.

293

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